ISSN 0394-9125

STUDIA GEOBOTANICA An international journal

Voi. 20 2001

EDITORS G. Cristofolini - Bologna Lj. llianic' - Zagreb E. Mayer - Ljubljana P.L. Nimis - Trieste L. Orlòci - London, On. F. Pedrotti - Camerino S. Pignatti - Roma A. Pirola - Pavia L. Poldini - Trieste E. Wikus Pignatti - Trieste MANAGING EDITOR M. Tretiach - Trieste SECRETARY M. Codogno - Trieste

Dipartimento di Biologia Sezione di Geobotanica ed Ecologia vegetale Università di Trieste

Studia Geobotanica. Voi. 20: 3-16 (2001)

MALLEE COMMUNITIES ALONG ROADSIDES IN SOUTH-WESTERN AUSTRALIA

Erika PIGNATTI WIKUS ', Giuseppe PIGNATTI 2 and Stephen D. HOPPER'

' Dipartimento di Biologia, Università di Trieste, Via Giorgieri I O, l-34127 Trieste, ltaly. 2 Istituto Sperimentale per la Selvicoltura, Viale S. Margherita 80, l-52100 Arezzo, ltaly. 3 Botanic Gardens and Parks Authority, Kings Park & Botanic Garden, West Perth, 6005 Western Australia.

Keywords: conservation, Eucalyptus, mallee vegetation, phytosociology, Western Australia.

Abstract: Vegetation with eucalypts of the mallee type along four roadside transects in the South-West of Western Australia has been studied. Mallee communities occur under dry-mediterranean climate conditions. Species composition, diversity and turnover, locai endemism with particular regard to Euca/yptus and Acacia species and habitat conservation are discussed.

Introduction

Mallee eucalypts are many-branched evergreen A von, Roe and Eyre, and is the major agricultural shrubs with woody stems of about 2-4 m height Wheatbelt of the southwest and part of the rising from prominent underground fire-resistant Goldfields, woodlands and pastora] country beyond lignotubers. In areas with dry-mediten-anean clima­ (i.e. Beard's Southwestem Interzone). Of approxi­ te, where rainfall and soil conditions can only sup­ mately 500 species of Western Australian eucalypts, port small trees in natural vegetation, the mallee about 225 are mallees and the above mentioned habit seems an adaptation to above ground damage, three regions are the richest in taxa, with numbers caused more often by frequent fires in ecosystems declining inland and westward (Hopper 1992). As (Beadle 1981). Alternatively, the lignotuber may discussed by Hnatiuk & Maslin (1988), the TRZ is have arisen originally as a simple way of maximi­ the area of highest species richness for Acacia in sing space for access to soil, water and nutrients. Australia. There are two foci of richness, one cen­ Besides the mallee eucalypts many different species tered on Wongan Hills and the other on Newdegate, of the genus Acacia of the phyllodineous type play with in excess of 100 species recorded for each cen­ an important role in mallee communities (Maslin & tre. As with Eucalyptus, the numbers of Acacia spe­ Hopper 1982). cies decline rapidly towards the arid interior and the According to Beard (1990) in southwestern temperate western areas. Western Australia mallee vegetation occurs mainly At present conservation status of species genera­ in the natural regions (Fig. 1) of the Wheatbelt tes concerns due to the land use in the area, where 2 (extent of mallee vegetation 3,596 km ), Mallee extensive farming has changed most of the native 2 Region (60,680 km ) and Esperance Plains (21,410 vegetation into agricultural land (Hopper 1990). 2 km ), developing three physiognomic types: mallee Burgman (1988), who studied the design of appro­ with sclerophyll understorey, mallee with patches of priate reserves specifically for mallee vegetation in woodland and mallee-heath (kwongan). These Western Australia, determined that reserves should regions, placed between the low-rainfall zone of the be located at least every 15 km for the same broad interior of Western Australia and the high-rainfall formations and soil types if they are to conserve of the western and southem coasts, and therefore representative flora, especially rare species. Rapid called Transitional Rainfall Zone (TRZ, Hopper geographical replacement of taxa occurs in the 1979), may be regarded as species rich in relation to Western Australian mallee, and a close network of the other regions (Marchant 1973). The TRZ (300- reserves is needed as a consequence. 800 mm annua! rainfall) includes the four districts Of particular interest in this context is the road­ of the South-West Botanical Province, i.e. Irwin, side vegetation, which, in a heavily cleared agricul-

3 E. PIGNATTI WIKUS e/ al.

300 ?50 South-West Western Australia 350

400 RAINFALL Annua! average in m,llimelres O Predom1nantly matlee areas D Areas where mallee ,s s,gnificanl bui noi predom,nanl

30' 0

4•

O 100 200 300 1200 1000 KILOMETRES

° 112· 11s· 120" 124 128' I Fig. I - Map of the area with natural regions of rnallee cornrnunities in Western Australia (Wheatbelt Region, Mallee Region, Esperance Plains). turai land, plays an important role as a corridor or the regime of winter rainfall, but annua! averages reserve for isolated pockets. Roadside vegetation in vary. In the TRZ, to which the transects of the pre­ Western Australia is protected by rather strict laws sent study belong, precipitations in certain years which means that it should remain untouched and may be extremely reduced or even lacking entirely, preserved as much as possible; farmers are obliged while in others unexpected rainfall may alter the to safeguard and protect these vegetation strips yearly mean values. The criticai climatic factor for along roadsides, around farmland and pastures. vegetation is not the annua! average rainfall but the Even where the present laws are respected, it may length of the dry period and the season of the year be sometimes difficult to interpret what the natural in which it falls (Hnatiuk & Maslin 1988, Beard vegetation might have been Iike, since much may 1992). depend on the surface extension of such areas and The vegetation relevés are located in the extra­ the intensity of human impact in the past. dry mediterranean to dry-medite1i-anean zone with Nevertheless the roadside vegetation pockets are 7-8 dry months and winter precipitation from 300- often the last remains of the former natural vegeta­ 500 mm per year (Fig.1). Vegetation changes in tion, so that it is indeed possible to provide a relia­ composition from W to E (increasing aridity) and ble analysis of the changes in species composition from N to S (increasing humidity). The yearly rain­ in strongly altered areas by human intervention. fall is almost constant from 338 mm (Morawa) and This research aimed to study roadside vegetation 356 mm (Buntine) in the North to 417 mm types and their relation to environmental changes of (Ravensthorpe) in the South. But most striking dif­ mallee communities according to the phytosociolo­ ferences are the number of days with high atmo­ gical method of Braun-Blanquet in the TRZ along spheric humidity (61-70 in the North and 112 in the the direction north-west/south-east. South) and temperatures. Therefore the relevés are Materiai and methods scattered along a climatic gradient from a warmer climate with longer lasting aridity in the North Climate and soils towards cooler and moister conditions in the South (Gentilli 1951). The South-West of Western Australia is under The whole area is characterised by geologica!

4 Studia Geobot. 20 (200 I) - Mallee communities in SW Australia - uniformity with gently undulating surfaces. The cies complexes and hybrids are frequent there landscape is largely flattened and shaped by erosion (Hopper 1992). The natural vegetation of this zone long ago. Only modest orogenic activity has taken is complex due to its history. piace since the Permian. Important rivers are no lon­ By law countryroads in Western Australia have ger active in the area of the transects. to have on both sides a strip of uncleared vegetation The different soil types lie on archaean granites which has to be left rather undisturbed and therefo­ with unfolded metamorphics of the Yilgarn Block. re can represent natural or seminatural conditions. Bedrocks of granites and gneisses are largely cove­ Roadside vegetation may not be the best example to red by sandy and gravelly soils overlying iron- and study plant communities, but in Western Australia, aluminium -rich duricrust, the latter above the pal­ especially in the Wheatbelt area, this often repre­ lid clay horizon. All soil types in the area are sents the only remnants of the natural vegetation. nutrient-poor for plant life. Nevertheless vegetation The possibility of aliens entering such areas can be is interrelated with soil. On the basis of changes in expected, accidental fires may start more easily soil, in generai, catenary sequences of natural vege­ close to roadsides, fire-sensitive species may be tation can be observed i.e. kwongan (scrub-heath) destroyed after frequent fires, seedbanks impoveri­ on sandplains, thickets on ironstone gravels, wood­ shed, and evolution of plant communities and their lands of Eucalyptus loxophleba, Eucalyptus salmo­ normai species composition might be impeded. nophloia on loams, and halophytes on saline soils. Human impact and grazing effects may cause Valley soils usually are red loams. Uplands show a further disturbance. Also extra run-off water and deeply weathered profile and are often covered with dust may affect plant-life along roadsides. Outside sandplain soils. The sand quantity is varying in pro­ roadside strips the natural vegetation has often been portion of ironstone gravels. In generai, woodlands completely destroyed due to agricultural and pasto­ are on valley soils, kwongan is on sandplain, while ra! use or human settlements in the vicinity. To mallee occupies an intermediate position between interpret what the natural vegetation might have valley soils and sandplains (Beard 1976). been like, much depends on the surface extension of Southwest Western Australia has experienced such areas and the intensity of human impact in the no glaciation since the Permian Era. During the late past. Nevertheless often it is indeed possible to pro­ Tertiary and Pleistocene environmental perturba­ vide a reliable interpretation of the natural plant life tions were most pronounced in the TRZ (Bowler based on the actual species composition. 1982, Martin 1990). Study areas and methods Species richness The relevés were chosen from North to South The entire Southwest Province of Western along four roadside transects rich in mallee Australia has undergone a long and complex evolu­ eucalypts. Locations of the four transects are as fol­ tionary history resulting in a rich angiosperm flora lows (Fig. 1): which can be estimated 8,000 and maybe even as A) transect from Mullewa to Pithara: the loca­ high as 9,000 species. As a consequence of environ­ tion is between the Northern Sandplains and the mental perturbations in the TRZ during the late Murchison Region, in the northern part of the Tertiary and Pleistocene the flora has radiated Wheatbelt Region; the transect plots were taken extensively at the species leve! (Acacia about 560 along the road within a length of about 250 km, species in all of Western Australia, Eucalyptusmore from 29.08.1994 to 01.09.1994 (Tab. 1, rels. 1 to 21 than 400 species only in the Southwest Province, and Appendix). Grevillea about 200, Melaleuca about 150, Hakea B) transect from Jilbadge Rock to north of and Caladenia about 100, etc.). The speciation pro­ Hyden: the location is between the Wheatbelt and cess has been concentrated mainly on woody peren­ the Mallee regions, starting southwest of Southern nials (comprising ali types of phanerophytes and Cross; distances between relevés are about 120 km; chamaephytes). After continued improvement of first transect plot on foothill of granite outcrop, the floristic, genetic and phytogeographic knowledge, others along roadsides in direction of Hyden; the number of endemics is stili increasing and may relevés date: 11-12.09.1994 (Table 1, rels. 22 to 27 be even higher than 80% (Hopper 1979). Many rare and Appendix). locai endemie species are concentrated in the TRZ C) transect from east of Lake King to Frank and subspecies, infraspecific variants, cryptic spe- Hann National Park: located in the Mallee Region,

5 E. PIGNATTI WIKUS et al.

Transect plots start about 70 km east of Lake King PH = phanerophytes, i.e. tall trees (PH scap), tall and continue from there into Frank Hann National shrubs (PH caesp); Park; distances from plot 28 to 32 are about 70 km; PHc = woody climbers; relevés date: 13-14.09.1994 (Tab. 1, rels. 28 to 32 PHm = eucalypts of the mallee type; and Appendix). NPH = nanophanerophytes, i.e. small shrubs more D) transect Cascade Road-Oldfield Road: the or less up to 1-1,5 meters high); plots start along the Cascade Road and continue CH = chamaephytes, i.e. dwarfshrubs, lignified along the Oldfield Road; distances in between are creeping and cushion plants; about 80 km; relevés date: 23.09.1994 (Tab. 1, rels. H = hemicryptophytes, i.e. perennial herbs; 33 to 37 and Appendix) G = geophytes, i.e. perennial species with bulbs, Along the four transects 37 plots were studied rhizomes, tubers or corms; according to the Braun-Blanquet method, i.e. areas T = therophytes, i.e. a11nualscomprising aliens. were selected which were homogeneous as to spe­ cies composition and physiognomy and in which the Results vegetation was regularly distributed over the surfa­ ce. As to the extension of the relevés, it has to be The classification of the 37 vegetation relevés mentioned that it was not always possible to have using the euclidean distance algorithm and mini­ the same number of square metres, as roadside mum variance clustering method is shown in Fig. 2. vegetation can sometimes be limited and shows On the cluster diagram three main groups of relevés rapid gradients and changes. at a distance of 1.21 can be traced: group 1 i11cludes Regarding the usual performance of a relevé, the the 21 relevés of transect A (Mullewa-Pithara) plus starting point is always the choice of a homotoneous the first three of transect B (Southern Cross­ surface, the complete list of all given species, their Jilbadge Rock); group 2 is composed by the rem­ cover in % on the same surface and sociability nant relevés of transect B and all those of transect C (shape of growing within the selected surface, (East of Lake King-Frank Hann NP); group 3 is which was neglected in the statistic analysis). For composed by the relevés of transect D (Cascade­ statistically useful data the Braun-Blanquet's cover Oldfield Rd.). values scale was slightly modified according to Ordination of relevés by Principal Components Pignatti (1959). Analysis is shown in Fig. 3 (12% of tota! variance In order to give a11 interpretation to the vegeta­ explained). The three groups of relevés are well tion data, Cluster analysis (Fig. 2) and Principal distinct by the projection of the two first compo­ Components Analysis (Fig. 3) were applied 011 nents: group 1 on top of the left side of the diagram, cover values of species in the relevés, using the groups 2 and 3 on the bottom of this, with transect package MULVA (Wildi & Orloci 1996). D relevés on the highest values of the PCA first For the main groups of relevés obtained by clu­ axis. ster analysis, the life form spectrum, based 011 the The results of this statistic analysis, with sequen­ sum of prese11ces of every species belonging to the ce following the cluster analysis of relevés and spe­ same growth form, was calculated. Life forms are cies ordered in subjective way in order to give "the architecture and shape" (germ.: Bauplan and emphasis to Eucalyptus, Acacia and most discrimi­ Gestalt) species have adopted during the course of nating taxa, is shown in Tab. 1. evolution to be able to survive best during the most 51 families are represented by 139 genera and criticai periods. In a mediterranean type climate it is over 373 species (742 occurrences: with severa! the summer drought species have to cope with. The subspecies and varieties; 2 new species have been Iife form system according to Raunkiaer is apt and discovered (Acacia sp. nov. in transect A, rei. 17 quite sufficient for European parameters but should and Calectasia sp. nov., a stilt plant, in transect D, have some extension of new forms for other conti­ rei. 37). (Table 1, see Appendix). nents and climates (Box 1981). In Western The participation of the most frequent families Australia severa! particular growth forms such as and the genus Acacia with the observed species grasstrees, stiltplants, species with lignotubers etc. number in the four transects is shown in Tab. 2. occur which do not exist in Europe. For the transects Despite the high number of eucalypts recorded dealt with in this paper we used only a slightly ( 46 species or subspecies), there is a certain number enlarged system according to Raunkiaer and distin­ of species with relatively high frequencies (20-60%) guished the following categories: in the three main clusters: E. Joxophleba, E. kochii

6 Studia Geobot. 20 (200 l) - Mallee communities in SW Australia -

o o +

o o 2 3

o o �-

Fig. 2 - Cluster analysis dendrogram with the three groups ofrelevés (1-3). 0,5

0,4

0,3

<> 0,2 <> <> <> Group 1 <> <> <> e <><> <> 0,1 <> -0,5 -0,4 -0,3 0,1 0,2 0,3 &0,1 Gr up 3 Group 2 o o f6o -0,3 -0,4

-0,5

Fig. 3 - PCA with ordination of relevés.

7 E. PIGNATTI W!KUS et al.

Tab. I - Transect relevés with cluster groups A-D.

Number ofrelevé N o .... "' "' a, N � "' N :: � N � � � S? � � .... � "' � a, .., ;;; ;! N N N N gi N g "' ;;; ;:i; � � :Il � Mallee (surface cover in%) o "' o o "' o o o N o o � � N � N R � .., "' N N .... g g g R � Kwongan (surface coverin %) � g o og g g o o 2 aìo g g g g go g g o g o � � .., N g $ g 2 2 .., o o N � g N � g 2 g .... g � .... Herbs (surf ace cover in% ) g "' g g "' o o g g � 2 g 2 2 2 � � g � a, g N � "' N � � � "' g "' g .... "' g N � o o o � "' N o o � o N Speciesnumber N "' a, "' o ...... "' o "' .... a, "' "' "' "' S? � N � ;'!: � ;;; N N ..,N ):j � N N � :: � N N N � N ;ll N N ):j N N N ;;; � S? � N N � Surface (m2 ) o "' "' o 8 "' 2 "' � 2 N � � � § 2 2 2 2 2 2 � N 2 2 2 .., .., N 2 § � � § 2 § § 2 § § § § § Cluster group 1 1 1 1 1 2 2 2 2 2 2 2 2 3 3 3 3 3 Freq. Transect A A A A A A B B B A A A A A A A A A A A A A A AB B B e e e e e D D D D D

Eucalyptusloxophleba + 2 3 4 4 3 + + + + 11 Eucalyptus kochii subsp. kochii 1 + 1 2 2 r + 7 Eucalyptus subangusta subsp. pusilla 2 2 5 Eucalyptussalmonophloia 1 1 2 4 Eucalyptus leptopoda + + + 2 4 Eucalyptus oldfieldii + 3 + 1 5 Eucalyptus wandoo + + 1 3 Eucalyptussalubris 2 + 3 Eucalyptus eudesmioides 1 + 2 3 Eucalyptus erythronernavar. marginata + + 3 Eucalyptus py1iformis + 3 2 Eucalyptus eremophila subsp. erernophila 2 + 1 4 Eucalyptus kochii subsp. pienissima + + 2 Eucalyptus platycorys + + 2 Eucalyptus luteola + 2 2 Eucalyptus densa subsp. densa 2 + 3 Eucalyptus perangusta + + + 3 Eucalyptusgrossa 1 + 2 Eucalyptusconglobata + + 1 3 Eucalyplusleptocalyx + + + 3 Eucalyptustetragona 2 + 3 Eucalyplus flocktoniae + + + 3 Eucalyptus cylindrocarpa + 2 Eucalyptus uncinata + 1 + + 4 Eucalyptus phaenophylla subsp. interjacens 1 2 3 Eucalyptus eremophila s.str. 1 1 2 Eucalyptus platypus 2 + 2 Eucalyptus kessellii subsp. kessellii + 2 2 Eucalyptus sheatiana + Eucalyptus aff. erernophila + Eucalyptus rigidula Eucalyptus ebbanoensis Eucalyptusewartiana 3 Eucalyptus stowardii Eucalyptus diminuta + Eucalyptus phaenophylla subsp. phaenophylla Eucalyptus pileata 2 Eucalyptus obesa + Eucalyptus tereba + Eucalyptusstoatei + Eucalyptus suggrandis subsp. suggrandis + Eucalyptus forrestiana 2 Eucalyptus tetraptera + Eucalyptus aff. conglobata Eucalyptus tumida + Eucalyplus xyphocalyx +

Acacia acuminata + + + + + 2 + + + 11 Acacia acuaria + + + + + 7 Acacia anthochaera + + + 4 Acacia neurophylla subsp. erugata + + 3 Acacia longispinea + + + 3 Acacia aff. anthochaera + + 2 Acacia coolgardiensis subsp. coolgardiensis 2 2 Acacia microbotrya var. borealis + 2 Acacia andrewsii + + 2 Acacia aulacophylla + + 2 Acacia hemit8es + + + + 6 Acacia yorkrakinensissubsp. acrita + + + 4 Acacia colletioides + + + 2 4 Acacia gonophylla + 1 2

8 Studia Geobot. 20 (200 I) - Mallee communities in SW Australia -

Tab. [ - Continued

Acacia tetragonophylla + Acacia merrallii s. I. + Acacia chrysocephala + Acacia fragilis Acacia spec. nov. + Acacia multispicata + Acacia restiacea + Acacia nigripilosa subsp. nigripilosa Acacia daviesioides Acacia cfr. assimilis + Acacia longiphyllodinea + Acacia cfr. erinacea + Acacia assimilis subsp. assimilis + Acacia mutabilis var. angustifolia + Acacia evenulosa + Acacia assimilis subsp. atroviridis +

Waitzia acuminata + + + + + + + + + + + + 13 Stipa compressa 2 + + + + + + 1 g Thysanotus manglesianus + + + + + + + + 8 Avena barbata + 2 + 3 + + + 8 Erodiumcygnorum + + + + + + + + 8 T rachymene pilosa + + + + + + + 7 Hypochoerisglabra + + + + + + 6 Podolepiscapillaris + + + + + 5 Arctotheca calendula + + + + + 5 Hyalospenmum glutinosum + + + + + 5 Gilbertia tenuifolia + + 1 + 5 Rhagodia baccata + + + + 4 Bromussterilis + + 4 4 Trachymene ornata + + + + 4 Ptilotus obovatus + + + 4 Melaleuca websteri + + + 4 Cheilanthes austrotenuifolia + + + + 4 Podolepis lessonii + + + + 4 Alyxia buxifolia + 1 3 Choretrum pritzelii + + + 3 Olearia muelleri + + + 3 Osteospermumclandeslinum + + + 3 Podotheca angustifolia + + + 3 Astrolorna serratifdium + + + 3 Neurachne alopecuroidea + + + 3 Podolepis canescens + + + 3 Ursinia anthemoides + + 3 Daucus glochidiatus + + + 3 Helichrysumdavenportii + + 2 Raphanus raphanistrum + + 2 Brachyscorne iberidifolia + 2 Ctenocephalus pseudevax + + 2 Hakea preissii + + 2 Rhagodia eremaea + 1 2 Cephalipterum drummondii + + 2 Cassia sturtii + 2 Cryptandra leucophracta + + + + 4 Glischrocaryon aureum + + + +_ 4 Gre,;11ea eriostachya + + + 4 Malleostemon tuberculatum + + + + 4 Calytrix superba + + + 3 Psammornoya choretroides + + + 3 Schoenus pleòostemoneus + + + 3 Hakea coriacea + + + 4 Hakea sulcata + + 2 Hibbertia polyclada + + 2 Verticordia cfr. chrysantha + + 2 Brachysema daviesioides + + 2 Cryptandra pungens + + + 3 Melaleuca conothamnoides + + 2 Chamaelaucium parviflorum + + 2 Drosera erythromiza + + 2 Ecdeiocolea monostachya + 2

9 E. PIGNATTI WIKUS et al.

Tab. I - Continued

Velleia rosea + + 3 Gnephosisbrevifolia + + 2 Maireana tomentosa 2 2 2 Velleiacycnopotamica + + 2 Blennospora drummondii + 2 Dodonaea cfr. aptera + + 2 Exocarpos aphyllus + 1 2 Dianella revoluta + + + + + + + + 8 Melaleuca cardiophylla + + 1 + + + + 1 + 9 Allocasuarinacampestris + 2 + + + 6 Helichrysum lindleyi + + + + + 6 Grevillea paradoxa + + + + + + 6 Santalum acuminatum 1 + + + + + 6 Isopogon scabriusculus + + + + + + 6 Cornespermavolubilis + + + + + 5 Allocasuarina thuyoides + + + + 4 Grevilleaarmigera + + + 4 Persooniaquinquenervis + + + + 4 Santalum spicatum + + + 3 Eremophila scoparia + + + 3 Mesornelaenastygia + + + 3 Astrolorna serrat�olium var. horridulum + + + 3 Lysinerna ciliatum + + + 3 Scaevola pulvinaris + + + 3 Banksia elderiana + 2 Banksia lul�itzii + + 2 Goodenia filiformis + + 2 Grevillea petrophiloides + 2 Hibbertia aurea + + 2 Melaleuca uncinata + + + 2 + 1 2 2 + 1 11 Thryptornenetenella + + + + + + + + + 12 Melaleuca scabra s.l. 2 + + + 2 + 1 10 Phebaliumtuberculatum + + + + 5 Phebalium drummondii + + 3 Allocasuarina corniculata 2 2 + + 1 + 6 Hakeameisneriana + + + + + 1 6 Malleosternon roseum + + + + + 6 Callitris verrucosa + + 1 + 4 Orummondita hassellii + + + 3 Grevrneahookeriana + + + 3 Hakea multilineata + + + 3 Eriosternonspicatum + + 2 Scae'v'Ola arenaria 1 + 2 Conothamnus aureus + + + + 4 Leptomeria spinosa + + + 1 5 Melaleuca calycina + + + 3 Gastrolobium spinosum + + + + 1 + + 7 Verticordia chrysantha + + + + + + 6 Leptospermum erubescens 1 + + + 5 Calytrix lechenaultii + + + 3 Loganiabuxifolia + + 2 Cassytha racemosa + 1 3 Hakea nitida + + 2 Grevilleapauciflora + + 1 + + 5 Comesperma spinosum + + + + 4 Hibbertiacfr. rupicola + + + 3 Grelillea pectinata + 1 2 Calothamnus gracilis + 1 2 lsopogon polycephalus + + 2 Lepidosperma tenue + + + 3 Banksia dryandroides + + 2 Banksia media 2 2 2 Baronia albida + + 2

10 Studia Geobot. 20 (200 I) - Mallee communities in SW Australia - subsp. kochii and E. subangusta subsp. pusilla for and 3 (Grevillea pauciflora, Comespenna spinosum, group 1, E. oldfieldii and E. eremophila subsp. ere­ Hibbertia cfr. rupicola). mophila for the groups 1 and 2, E. uncinata and E. The life form spectra in the three main relevés phaenophylla subsp. inte1jacens for group 3. Acacia groups are shown in Tab. 3. It is noteworthy: acuminata and A acuaria reach the highest frequen­ 1) the increasing percentages in mallee eucalypts cies in group 1, A hemiteles, A. yorkrakinensis from group 1 to group 3 (from 14.4 to 26.9%); subsp. acrita and A colletioides in groups 1 and 2. 2) the relevant participation of chamaephytes (15.1 In the herb layer the highest occurrences (= fre­ to 25,8%); quencies) in group 1 were reached by Waitzia acu­ 3) the high percentage of nanophanerophytes in minata, Thysanotus manglesianus, Trachymene group 2 (53%); pilosa, Podolepis capillaris, Hyalospermum glutino­ 4) the high percentage of therophytes (mostly sum, Stipa compressa, now Allstrostipa compressa, aliens) in group 1 (29.1%). Erodillm cygnorwn, nearly all with low cover values. Also some aliens like A vena barbata, Discussion Hypochoeris glabra, Arctotheca calendL1la appear Vegetation patterns in the transects frequently. Some species have a distinct distribution in groups 1 and 2 (Dianella revoluta, MelaleL1ca car­ Although the surfaces of plots in the four tran­ diophylla, AllocasL1arina campestris, Grevillea para­ sects can often be considered not sufficiently big doxa, Santalum aCL1minatum, Isopogon scabriuscll­ and therefore under the minimum area along the lus etc.), only 4-5 species are distributed in all grou­ geographical gradient NW-SE, where mallee com­ ps (MelaleL1ca uncinata, Thryptomene tenella, munities have their highest concentration, the gene­ Mela/elica scabra s.l., Phebalillm tuberculatum, rai trend emerging from vegetation plots shows a Phebalium drummondi1). Some species distinguish clear distinction between relevés of the north­ group 2 (Allocasuarina corniculata, Hakea meisne­ western and south-eastern part. As cluster and scat­ riana, Malleostemon roseum, Conothamnus aureus) ter diagrams reveal, the north-western group of relevés is composed by data collected in transect A Tab. 2 - Most frequent taxa occurring in the relevés. and the first three relevés of transect B, whereas, on the contrary, the remaining relevés of the last tran­ Taxon Specie sect and those of transects C and D form the second number group of relevés (Tab. 1). This division is not sur­ Myrlilceae 85 prising, because along a line in NE-SW direction Proteaceae 52 Acacù1 37 between Southern Cross and Hyden, the border of Asteraceae 26 the two natural regions is placed, i.e. between the Poaceae 19 Wheatbelt in the north and the Mallee Region in the Fabaceae 9 south, characterizing the TRZ. As pointed out by Goodeniaceae 8 Beard (1992), there are differences in climate and Dilleniaceae 8 Liliaceae s.l. 8 soil between the Wheatbelt Region vs. Mallee Chenopodiaceae 7 Region, which cause a remarkable change within Rutaceae 7 the catenary sequence in the two areas. In the Santalnceae 7 Wheatbelt Region the mallee which occurs on Epacridaceae 7 duplex soil comprises a minor portion of the catena Restionnceae 7 which is dominated by woodland and kwongan. In the Mallee Region, however, mallee eucalypts are

Tab. 3 - Percentage of !ife forms within the three groups of dominant in the topographic sequence, extending relevés. along the whole slope and surpassing woodlands and kwongan, which occupy respectively the bot­ Life Form Group I Group 2 Group 3 tom and the top of the slopes. This fact is explained PHm 14.4 15.8 26.9 by the tota! rainfall which is higher in the western PHc o I 3.7 areas and creates more favourable soil moisture NPH 30.7 53 41.7 conditions. CH 15.1 25.8 22.2 The difference in habitat between the two natu­ H 2.5 0.5 o G 8.2 3.5 5.6 ral regions could explain basically the distinct vege­ T 29.1 1.5 o tation features in the groups of relevés, as underli-

11 E. PIGNATTI WIKUS et al. ned in the multivariate analysis (Fig. 3). In fact we sandplains and species-rich, becoming less so in can assume that along the transects a climate gra­ dryer areas (Hill 1990). dient could also be responsible for vegetation chan­ Most mallee stands contain severa! eucalypt spe­ ge: mean temperatures decline in a N-S and W-E cies varying mostly from 2-8 in the four transects. direction, whereas total rainfall is more or less con­ The most significant development of mallee stant in the Wheatbelt, but increases in the Mallee eucalypts in terms of number and species diversity Region in NW-SE direction. The better moisture occurs in southwestern Western Australia where conditions could explain the ordination of relevés extensive speciation and radiation have taken piace. on the first PCA axis, which shows, from left to The mallee habit may have arisen early in history or right, both groups of relevés, the sequence from at a later time; some species appear to have secon­ vegetation data carried out in dryer conditions darily lost the mallee habit and returned to tree (Mullewa area and Hyden area) to those under moi­ forms; severa! moort, marlock and mallet species ster conditions (Dalwallinu area, Cascade-Olfield are of this type (Hill 1990). Rd area). Basically, PCA axis 1 could be interpreted The mallees of the transects are ali sclerophyl­ as related to moisture (increasing moisture with lous shrubs of 1-5 m height, adapted to fires, seaso­ higher x values) and PCA axis 2 to temperature nal drought and extreme nutrient-poor soils. (increasing temperature with higher y values). This Evidently a high speciation rate occurred in may explain the higher occurrences of therophytes eucalypts predisposed to such conditions (Beard (species adapted to dryer conditions) in the northern 1992). relevés (Cluster group 1, Tab. 1). Although climatic and soil factors seem to play The Acacia species of the transects an important role in geographical replacement of vegetation in the mallee communities, biogeo­ Acacias, with 950 species presently recognised, graphical considerations should also be considered. form the largest group of vascular plants in Eucalyptus subangusta subsp. pusilla, Waitzia acu­ Australia (Fig. 4A). 560 species occur in Western minata (distribution area: Austin, Avon, Coolgardie, Australia (Hnatiuk & Maslin 1988, Maslin 1995). lrwin Botanical Districts) and several other taxa can The southwest region is most distinct regarding the be considered as indicator species for the north distribution of Acacia species and shows the highe­ western mallee communities. Others like Grevillea st species richness. Species distribution maps of Fig. pauciflora, Melaleuca calycina etc. seem to be more 4 demonstrate that the area where transect A is frequent in the southern distribution area of mallee situated, is dose to the edge of highest species rich­ communities. Aliens show highest frequencies in ness of Acacia within Australia and the world the relevés of the Wheatbelt Region, where land (Maslin & Pedey 1982). One third of the taxa of the clearing for agricultural purpose was most intense. region are present in the transect relevés. Although The collected data show a geographical replacement they represent all widerspread and rather common of species and explain the main characteristics of species within the area, they are a significant sam­ the Natural Regions, as described on a different ple of northern Wheatbelt taxa of Acacia. While A basis in literature (Beard 1992). In addition they acuminata and A microbot1ya are widespread over point out the potential danger of diffusion of aliens the entire Wheatbelt, A coolgardiensis, A. longispi­ and reduction of the corridor role of native vegeta­ nea and A anthochaera are restricted to the northern tion remnants. part. Very few extend their distribution area to the East (A tetragonophylla). All the rest are basically The mallee species of the transects endemie to the area. Most of the acacias listed in the four transects Present-day occurrences of mallee communities belong to the subgenus Phyllodineae. This group are closely related to edaphic factors dependent on has undergone extraordinary diversification within rainfall, substrate and landform. Many mallees Australia (Maslin 1995, see Fig. 4B) and can beco­ occur over a wide climatic range, and on different me an important element of the landscape, particu­ soil types as extremes of this climatic range (Hill larly in arid and semiarid areas (Maslin & Pedley 1990). For example, Eu. tetragona (now largely 1982). About half of the species listed in transect A pruinoca1pa) and Eu. eudesmioides can occur on belong to the Phyllodineae, the other half to Ju­ lateritic sandplains but also on calcareous coastal liflorae and Plurinerves; none of section Pulchellae dunes. Mallee communities with kwongan vegeta­ was listed. (Sect. Phyllodineae has 1-nerved phyllo­ tion in the understorey are frequent in the SW on des and spicate inflorescences, sects. Juliflorae and

12 Studia Geobot. 20 (200 I) - Mallee communities in SW Australia -

120 ° 126° 132° 138 ° -- 150° 2' � r.A. 12 ,J- 1 _lJ,.... 1 -- 1 1 \ � f\. <.. N 2 '/' :,. pl 2 1 ) . \/ I'--. " ,, 6' , N I 2 16 lr ,; f'._2 i'-,, r--2 zÌ I/ 11, i' 2 • 2 1 iv N r---- 1 2 .J ,· li I 2 1 1 1 2 j/1 • ' • 1 2 2 _v; 2 1 q·1 1\ /1 1 '. 1 '-- __) I 1 ' IC.....[___) l ,) '� O' J 7 J 2 20 • • !l,--i; 2 1 2 1 l 1 ! ' 2 I 2 _/ 1 I, l ,, [\l\ ' .,, ' 7 I • I J 2 2 2 J ' • • J ' 2 ' 1 ' I ' ' 11\ 7 " t 112> ' 2 ' 2 1 2 • • • ' s J 1 J 2 • • -è\ "' I , I 11 ' 7 2 I ' I 1 ' 1 q!j' ) l ; 2 J 1 ' 0 " • i • ® ) • ' ® ° 4 ' . -· 4 . � I ' 1 • 1 2 J 1 1 �- ' ' 7 ' 5 • • J • ) I • (10 " ,, ti , 1 2 I 2 J l 11. 10 12 2 I 7 5 1 ,_ 5 -.)I 17 ' ' • • I , ' . ·-· '----2 ·- • . o " • ,NZ' ' 7 ' I ' I ' Q J ,,,V, I ' 2 -,· 5 • • 2 ' ' ' • ,, ,,. 17 7 7 t I 1 1 �.- 1 7 1 J�\' 5 , I ' 2 )I ' • • ' ' 1 I ' • 7 " !5 \JZ e' " ' . I 21\ 17 7 • 2 . 2 2 1 I • J 1 • 2 ) • ' 1 . �--' • ' 2 .__ 12 " ,. 28 22 ltì 11\ 2 7 l 1 1 1 1 ;e ' l J 2 • • •! ' • ' 1 ' • " 21 I• \ " 5])1 "' • !ili.i 1 • J 1. 2 J • 2 • 2 Il\ ' 2 • • ' 2 " 17 11 • ·�'\ " 5 1 1 '1 J I 1 J J Il 11 • ' ' ' " 12 2' o I 'ì\ .,., • " ° )1 .. 12 11 10 32 131 ·� 22 10 2 I Il 7 I \Il i • " " "' i-,.,,re-I ,, " " ' " I 11 2.J) 7 i• 2 I 1 12 22 ��4-" � 11)1 � 1.i � " (n_ " J., A) 21 ...i�1·iil -,-=-e:-r 'li• t�."{7 t f • 11 _, . 2 " " 21 1 - ..!.' " I 11 ,. • 11 (lo 21 1, 6' � '--=-� -'-" ,12 ,,,. " " " "> 21 ,, 36 11 1_M �· N_l 17 1l ,,.,, � "-L"- ' �,· 12-- ,, -. V 1 ' 1 O' ,_ 1 -'- • '

° 40 ° 126 ° 132° 138° 144° �� 150 , .. 120 •••

° Fig. 4 - Species richness in Australia of the genus Acacia (Fig. A), based on 840 species, and of its section Phyllodineae (Fig. B); I 0x 1.5 grids (after Masi in & Hnatiuk I 988 and Masi in & l-lnatiuk I 995, respectively).

13 E. PIGNATTI WIKUS et al.

Plurinerves have multinerved phyllodes and flowers sive communities of mallee eucalypts on the conti­ are in globules.) nent. In Western Australia about 500 species of Acacias clearly present close correlation to eucalypts occur of which about 225 are mallees, 86 edaphic factors (Maslin & Hnatiuk 1983). A hemi­ species are indicated as rare (following the IUCN teles, A. microbotrya and A anthochaera (all sect. categories), 44 as vulnerable, 3 as endangered and Phyllodineae) grow on heavy soils (i.e. loam and 44 as poorly known. This incredible biodiversity is clay), Acacia coolgardiensis, A. neurophylla, A. fra­ concentrated mainly in southern coastal regions. gilis, A. Jongispinea, and A multispicata (sects. Mallee eucalypts in southern Western Australia Plurinerves and Juliflorae) grow on permeable soils have an extensive distribution from isolated patches (i.e. Iighter, gravelly sands). Anomalous is here A in the high rainfall forests and along the extreme acuminata (from the Juliflorae group) on heavier south western coast, through the heavily cleared soils. Acacias generally do not favour very salty Wheatbelt out into the western deserts (the Great areas (Maslin & Pedley 1982). Victoria Desert is especially rich in mallee commu­ For the geographic pattering of species distribu­ nities). Those taxa most endangered consequently tion in acacias, precipitation (i.e. average quantity are found where destruction of native vegetation has and duration of precipitation) seems more impor­ been greatest, i.e. in the Wheatbelt. In the 1990s, 31 tant than temperature (Hnatiuk & Maslin 1988). (ali mallees) were given special legai protection by Different to transect A, cover values of Acacia being declarated as Rare Flora under provisions of species in transect B are not significant, besides the Wildlife Conservation Act. Of these 31, 12 are these stands show relevés with different species yet to be formally described. 22 are localised ende­ combination on a mixture of soil types. All acacias mics of different parts of the Wheatbelt. Most are widespread species, some of them towards the occupy such small areas that populations are threa­ end of the range (A hemiteles, A. yorkrakinensis, tened by accidental distruction (bulldozing, igno­ A. colletÌoides). Some of these species often occur rance ). around granite outcrops and have greater water con­ Consequently a considerable effort is made by sumption. Those species are common and generally CALM staff to ensure that landowners and the widespread in the area; only A chiysocephala is potential users of bulldozers are informed and linked to the southernWheatbelt area. shown the location of populations. In some cases, Transect C shows within the relevés a rather populations are threatened by major activities high percentage of kwongan species, but acacias destructive of native vegetation such as agricultural (except in rei. 4) are not relevant. This speaks for land clearings (e.g. Eucalyptus goniantha subsp. the fact that in generai acacias are not well repre­ goniantha) or open-cut mining (e.g. Eu. suberea, Eu. sented in kwongan. From 50 acacias gazetted for the Jateriticola). area grid only 2% are present in the plots. Species In ali four transects locai endemics, species with numbers decline rapidly away from centres of rich­ small distribution ranges, endangered species, geo­ ness; grid 276, Newdegate, shows 101 species in the graphical species variation, interpopulational diffe­ south (Hnatiuk & Maslin 1988, Fig. 4). rentiation and polymorphism within severa! genera Acacias in transect D are lacking or with insi­ have been observed and the necessity to preserve gnificant cover values (rei. 33-37). Plants are small, these areas seems obvious. Moreover at least two shrubby and endemie with rather restricted geo­ transects (B and D) are far from Nature Reserves graphic ranges confined to the southern Wheatbelt and National Parks and in these cases along roadsi­ and the adjacent Goldfields. des they are nearly the only strips of natural or Acacia pollen was found in early Miocene. The semi-natural vegetation left in the area. extensive speciation of the genus may thus have occurred at a time when variations in moisture regi­ Conclusions me were the pre-eminent forces in the taxonomic and ecologica! sorting of taxa. Climatic oscillations Mallee communities and related vegetation during the Pleistocene would have accentuated types (woodland and kwongan) have been studied such forces (Maslin & Hopper 1982). along four roadside transects in the Southwest of Western Australia according to the Braun-Blanquet Conservation issues phytosociological method. The 37 relevés affirm a high species turnover (gamma diversity) already As pointed out by Hopper (1990) Western within short distances and many eucalypts of the Australia has the greatest diversity and most exten- mallee type and wattles of the phyllodineous growth

14 Studia Geobot. 20 (200 I) - Mallee communities in SW Australia - form show patchy distribution. No single species is Chamaelaucium ciliatum ( +), Eriostemon tomentellus(+ ), Baeckea Rel. present in ali 37 relevés. 114 of 742 species listed sp. (I), Drnmmondita sp. (+). 9: 65 km south of Mullewa. Gnaphalium indutum (+), Gnephosis erioceplw/a (+), Rhagodi11 occur only once, among them 11 species of acicL1/aris (+), Sonchus asper (+). Rel. 10: 21 km to Morawa. Eucalyptus and 20 species of Acacia. The genera Cassia eremophila (+), Dodonae,1 Jarraeoides (+), Eremaea denti­ Eucalyptus (63) and Acacia (39) are the most fre­ culata (+). Rel. 11: 5 km south of Morawa. Acanthoca,pus preissii quent ones, followed in decreasing order by the (+ ), Atriplex prostrata (I), Co1ynothec11 macrantlrn ( +). Rel. 12: 25 genera Melaleuca, Hakea, Grevillea, Verticordia. km south of Morawa. Aristida stipoides (+), Ptilotus lividus (+), ( +), ( +). Rel. km south The number of Proteacae increases from north-west Stylobasi!lm australe VL1!pia myL1ros 13: 30 of Morawa. Gilruthia osbomei (+), Pentaschistis airoides (+), to south-east in the southernmost transects. The Ptilotus ex11ltatus (+), Daviesia sp. (+). Rel. 14: 5 km south of number of Acacia species is higher in the northern Perenjori. Hibbertia subvaginata (+), Lo/ium temulentum(+), Stipa Wheatbelt area transect than in the southernmost elegantissima (I), Stylidi!lm elongat!lm (+), Tragus 1wstrnlianL1s transect. Therophytes (mostly everlastings and (+). Rel. 15: 15 km south of Perenjori. Actinostrob!ls psammophi­ (+), Rel. 16: aliens) are frequent in the northern Wheatbelt area Ja Danthonia setacea(+), Verticordia grandiflora(+). 30 km south of Perenjori. Cyanostegia angustifolia (+), Hybanthus transect and nearly absent in the southernmost tran­ floribund!ls (+), Thonlllsia purpurea (+), Verticordia cfr. mona­ sect. delpha (1 ), Verticordù1 pict11 (+), Be1wfortia sp. (+). Rel. 17: 51 km In spite of the fact that several relevés are below south of Perenjori. Pelrophile slwttleworthia1l/l (+). Rel. 18: 63 km the minimum surface due to the particular environ­ south of Perenjori. Hakea bucculenta ( +), Phebalium microphyllllm Rel. 19: mental conditions along roadsides, the Braun­ (+). 88 km south of Perenjori. Brachysema tomentos!lm (+), Chorizema acicu/aris (+), Loxoca,ya flex!losa (+), Mallophora Blanquet method has shown its applicability and rngosifo/i11( + ), Mesomefaena tetragona ( + ), Patersoni:1 occidenta­ practical use also in these vegetation types which lis (+), Verticordia polytricha (+), Anarthria sp. (+), Dampiera sp. yet had not been described by phytosociological (+), Verticordia sp. (+). Rel. 20: 123 km south of Perenjori. Drosera pallida(+), Hypocalymma angustifolium (+), Leucopogon approaches. The groups of relevés resulting from r the statistica] analysis are related to environmental striatus (+), Pimele11 imbric11ta (+), T1ymalium cf. Jedifolium(+). Rel. 21: I km south of Pithara (water catchment). Bo1y11 sphaero­ changes (soil, rainfall and habitat) and show that cephala (+), Briza maxima (I), Dampiera linearis (+), Drosera vegetation maintains its ecological value also under macrnntha (+), Eremophi/a maculata (+), Gnephosis gonotricha disturbed conditions. (+), Hydrocotyle blepharoca1p11 (+), Hypoxis occidentalis (+), In a landscape unglaciated for millions of years Loxocarya cinerea(+), Phyllangiwn paradoxum (+), Scaevola spi­ Rel. like Western Australia with particular distribution nescens (+). 22: Near roadside and foothill of Jilbadge Rock; with single high trees plus mallee surface probably a few years patterns of species and rather small population after a fire; 11.09.94. Disphyma crassifoli!lm ( + ), Ha/oragis nodi­ sizes, the conservation of ecosystems with rare flora ( +), Senecio lautus ( +), Lomandra sp. (+ ), Melaleuca sp. (2). environment and species combination must also Rel. 23: Near roadside and foothill of Jilbadge Rock, mallee with take into consideration the preservation of areas out­ few scattered higher trees; light brown sand, gravelly; 11.09.94. side specifically designated Nature Reserves and Amyema miquelii (on Eucalyptus) (+), Grevillea cfr. asparagoides (+), sp. (+), sp. (+). Rel. 24: East of Yilbadge Rock, National Parks. This means that future management Daviesia Sida near roadsicle, mallee with scattered rare taller trees; hard loamy of specifically small and isolated populations which brown sand; I 1.09.94. Callitris roei (I), Lomandra sp. (+), may occur in limited areas Iike roadside vegetation MelaleL1c11 sp. (2). Rel. 25: 39 km south of petrol station near is required. Southern Cross, close to cross section 16 km SW of Moorine Rock on Southern Cross South Road towarcls Aramasco Roacl; I1.09.94. Appendix AJ/ocasuarina obesa(I), Danthonia cfr. caespitosa ( + ), Hakea erec­ /11 (+), Lepidosperma drnmmondii(+), Petrophile ericoides (+), Localitiesand sporadically occurringspecies of the relevés: Rel. 1: Stylidillm repens (+), Verticordia roei (+), Daviesia sp. (+), r 15 km north of Mullewa. Ptilotus divaricatus (+), Sisymbriwn offi­ Hibbertia sp. (+), Hibbertia sp. (+). Rel. 26: 87 km f om Moorine ciniilè (+), Th,yptomene tuberculata (+). Rel. 2: 7 km south of Rock southwards. Mallee on gravelly sand; 12.09.94. Baeckea beh­ Mullewa. Halgania preissii (+), Maireana carnosa(+), Medicago rii (2), Borya nitida(+), Chamaelaucillm brevifolium ( +), D1yandra minima(+), Pimelea cfr. wgentea (+), Ptilotus gaudich11udii (+), cyrsoides (+), D,yandra sp. (+) Hibbertia sp. (+), Persoonia sp. (+), Stipa cfr. hirs!lta (+), Waitzia aL1rea (+). Rel. 3: 18 km south of Persoonia sp. (+). Rel. 27: 140 km South of Moorine Rock in direc­ Mullewa. Caesia parvif1ora (+), Cheiranthera filifolia (+), tion Hyden; gravels; 12.09.94. D,yandra aff. nivea(+), Hibbertia Pittospornm phyllyreoides (+), Stenopetah,m tilifolillm (+). Rel. 4: sp. (I), Hibbertia sp. (+), Phebalium sp. (+).\I Rel. 28: 70 km befo­ Wilroy, 20 km south of Mullewa. Calytrix t7avescen.s (+), re Lake King from direction Hyden; mallee above kwongan, grave! Cy11tochaeta avem,cea (+), Grevil/ea phanerophlebia (+), over loamy sand; I 2.09.94. Allocasuarina acuti va/vis (2), Callitris Onychosepal!lm Jaxif7ornm (+). Rel. 5: 40 km south of Mullewa. drnmmondii(+), Grevil/ea s/wttleworthiana (+), Ciyptandra sp. Ehrharta longiflora (+), Hordeum leporinwn (+), Loli!lm rigidllm (+), Hibbertia sp. (+), Leucopogon sp. (+), Pime/ea sp. (+). Rel. 29: (+), Sonclws olerace!ls (+). Rel. 6: 46 km south of Mullewa. Frank Hann National Park, mallee on loamy sand; 13.09.94. AllocasL1ari1rn lehmanniana (I), Atriplex erilifolia ( + ), Anarthrfa prolifera (+), Daviesia spinosissima (+), Patersonia Codonoca,pus cotinilo/ius (+), Lamarckia aurea(+). Rel. 7: 49 km umbrosa (+), Persoonia tortifolia (+), Beaufortia sp. (+), Hibbertia south of Mullewa. D11mpiera haemathotricha ( +), Westringia rigida sp. (+), Cyperaceae (+). Rel. 30: Frank Hann National Park, mallee (+). Rel. 8: 58 km south of Mullewa. Be/fida graminea (+), 3 m high, white sand; 13.09.94. Allocasuarina humilis (+),

15 E. PlGNATTI WiKUS et al.

Cassytha pomiformis (+), Dryandrn armata(+), Synaphea petiola­ Beard J .S., I 990. The Mal/ee Lands of Western Australia. In: ris (+), Conostylis sp. (+), Dwnpiera sp. (+), Juncus sp. (+), Noble J.C. (ed.), The Mal/ee U1nds. A Conservation Loxoca1ya sp. (+). Rel. 31: Frank Hann National Park, on side­ Perspective. CSIRO. East Melbourne, 29-33. track, reddish-grey loam with iron pebbles on surface; 13.09.94. Bearci J.S., 1992. Plant Lite of WesternAustralia . Kangaroo Press, lsopogon teretifolius (+), Baronia sp. (+). Rei. 32: Towards the Kenthurst, NSW. eastern end of Frank Hann National Park, near main road, kwongan Bowler J.M., 1982. Aridity in the late Terti111y and Quatema1y of with low mallee on yellow sand; 13.09.94. Calothamnus asper (+ ), Australia. In: W.R.Barker & P.W.M. Greenslade (ecls.), Darwinia Juekmannii (+), Grevillea excelsior(+), Hakea cfr. recur­ Evolution of the flora and fauna of arid Australia. Peacock, va (+), Leucopogon sp. (+). Rei. 33: Cascade Rd. - Oldfield Rd. Adelaide, 35-45. Cassytha me/antha (+). Rei. 34: Cascade Rd. - Oldfield Rd. Rei. Box E.O., 1981. Macroclimate and Plant Forms. An introduction of 35: Cascade Rd. - Oldfield Rd. Ci1ustis dioica(+), Chamaexeros predictive modelling in phytogeography. Tasks for Vegetation /Jmbriata (+), Dielsia crispata (+), Hakea cinerea (+), Hakea Science, Junk, The Hague, 258 pp. co1ymbosa (I), Hakea rnscifolia ( +), lsopogon trilobus (+ ), Burgman M.A., 1988. Spatia/ analysis of vegetation pattems in Phymatoca1pus maxwellii (+), Regelia inops (+), Sollya hete­ southem Western Australia: implicLambertia inermis (+). Rei. 37: Cascade Rd. - Oldfield Rd. WesternAustralian Naturalist, 8: I 75-184. Astartea heteranthera (+ ), Calecwsia sp. nov. ( +), Daviesia euphor­ Hill K.D., I 990. Ma//ee Eucalypt Communities: Their Classi­ bioides (+), Hakea laurina (+), Halgania favandulacea (+), Daviesia fication and Biogeography. In: Noble J.C. (ed.), The Mal/ee sp. (+), Leucopogon sp. (+), Cyperaceae (+). Lands. A Conservation Perspective. CSIRO , East Melbourne, Species which could not be determinated exactly are listed 93-108. under "sp."; sporadic species in the phytosociologic sense are spe­ Hnatiuk R.J. & Maslin B.R., 1988. Phytogeography of Acacia in cies which occur only as a single individuai in the plots and whose Australia in re/ation to clinwte and species-riclmess. Austr. occurrence seems to have no importance for the species combina­ Journ. Bot., 36: 361-383. tion of the community. Hopper S.D., 1979. Biogeographic 11spects of speciation in the southwest Australian Flora. Ann. Rev. Eco!. Syst., 10: 339- 422. Acknowledgements Hopper S.D., 1990. Conservation Staws of Ma/Jee Euc,1/ypts in South Western Australia. In: Noble J.C., Joss P.J. & Jones The authors are indebted to Dr. Bruce R. Masi in (Perth) for fruit­ G.K. (eds.), The Mallee Lands. A Conservation Perspective. ful suggestions on the chapter acacias and for their determina­ CSIRO East Melbourne, 21-24. tions, to Dr. John S. Beard (Perth) for comments on a first draft Hopper S.D., 1992. Patterns of plani diversity at the population and and for making available his unpublishecl data; to Dr. Kingsley W. species Jevels in southwest Australfan ecosystems. In: Hobbs Dixon and Mr. Lione! Johnston (Perth) for assistance during field R.J. (ed.), Biodiversity of Mediterranean Ecosystems in excursions, cliscussionof the first resulls ancl generous hospitality Australia. Surrey Beatty & Sons, Sydney, 27-46. in Perth; to Dr. Judith Brown (Wanneroo) for making available her Marchant N.G., 1973. Species diversity in the southwestern flora. field data from the centrai Wheatbelt area; to Mr. & Mrs. John & J. Royal Soc. W. Australia, 56:23-30. Jucly Browne (Breakaway), Mr. & Mrs. Frank & Margaret Merry Martin H.A., 1990. Tertia1y climate and phytogeograpiiy in (Caraboda), to the Vanzetti Family (Moora), to the late Mr. Pate & southeastem Australia. Paleobot. Palynol., 65: 41-55. to Mrs. Thelma Daniell (Esperance), to Mr. Richarcl Pepper & Dr. Maslin B.R., 1995. Systematics and phytogeography ofAustralian Gillian Pepper Craig (Ravensthorpe) for their kind hospitality, to species of ACllcia: an overview. 4 pp. , lnstitute of Foresters of Dr. Paul G. Wilson (Asteraceae) and the Director and staff of the Australia Newsletter 36 (2): 2-5. Western Australian Herbarium for assistance with criticai speci­ Maslin B.R. & Hnatiuk R.J., 1983. The distribution of Acacia mens, to Miss Beatrice von Roemer (New York) for editing the (Leguminosae-Mimosoideae) in Western Australia. Part 4. A English text ancl to Prof. S. Pignatti for critically reading the draft. descriptive Account of Acacia Districts. Western Australian Technical assistance of Mr. F. Bersan (Trieste) is appreciated. Herbarium Research Notes n. 8: 23-55. This study was supporleci by the ltalian M.U.R.S.T. (Ministry Maslin B.R. & Hopper S.D., 1982. Phytogeography of Acacia of Scientific Research and Technology) 40%. (Leguminosae: Mimosoideae) in Centrai Australia. In: Barker W.R. & Greenslade P.J.M. (eds.), Evo/ution of the Flora and Fauna ofArid Australia. Peacock Publications, Adelaide, 301-316. Maslin B.R. & Pedley L., 1982. The distribution of Acacia References (Leguminosae: Mimosoidea) in Australia, part I & 2. Western Australia Herb. Res. Notes, 6: 1-171. Beaclle N.C.W., 1981. The Vegerntion of Australia. Cambridge Pignatti S., 1959. Fitogeografia. In: Cappelletti C. (ed.), Trattato di University Press, Cambridge, 690 pp. Botanica. UTET, Torino, 681-81I. Bearci J.S., 1976. An indigenous term for the Western Austra/ian Wilcli O. & Orloci L., 1996. Numerica/ exploration of community sandplain and its vegetation. J. Royal Soc. Western Austr., pattems. 2nd edition, SPB Academic Publishing, Amsterdam, 59:55-70. 171 pp.

Received September 9, 2000 Accepted July 9, 200 I

16 Studia Geobot. 20 (200I) Studia Geobotanica. Voi. 20: 17-32 (2001)

A PHYTOSOCIOLOGICAL STUDY OF THE HYGROPHILOUS VEGETATION OF SIERRA NEVADA (SOUTHERN SPAIN)

Carlos SALAZAR *, Juan LORITE **, Antonio GARCÌA-FUENTES *, Juan Antonio TORRES *, Eusebio CANO * and Francisco VALLE **

* Opto. Biologia Animai, Biologia Vegetai y Ecologia. Facultad de Ciencias Experimentales. Universidad de Jaén. Campus Las Lagunillas, s/n. 23071-Jaén (SPAIN).E-mail:[email protected] **Opto.Biologia Vegetai. Facultad de Ciencias. Universidad de Granada. Campus Fuentenueva. 18071-Granada (SPAIN).

Keywords: Chorology, Dynamics, Ecology, Edapho-hygrophilous communities, Riparian vegetation.

Abstract: A study on the vegetation related to streams and rivers of the Nevadense sector of Sierra Nevada (Southern Spain) has been carried out by detennining its main structural, floristic, ecologica!, chorologic and dynamic characteristics. A totaI of nine new syntaxa have been highlighted: four associations (Caricetum cam­ posii-cuprinae, Ranunculo granatensis-Coch/earietum megalospermae, Cirsio micranthi-Scùpelum holoschoe­ ni, Carici camposii-Salicetum atrocinereae), two subassociations (Myrrhoidi-Alliarietum petiolatae nepetosum grana/ensis, Carici camposii-Salice/um a/rocinereae salice/osum capreae), two variants (Rubo-Rose/um coiym­ biferae variant with Adenocarpus decorticans, Rubo-Coriarietum myrtifoliae variant with Adenocarpus decor­ licans) and one community (Cratoneuron commutatum and Anagal/is lene/la community).

Introduction

The vegetation of SieITa Nevada, and particu­ some interesting plant commumties related to larly that of the siliceous nucleus (the so-called streams and rivers ofthe Nevadense sector of Sie1Ta Nevadense sector) has been the subject ofnumerous Nevada, developed mainly in areas of the meso­ studies caITied out during the second half of the last Mediterranean and supra-Medite1Tanean them1oty­ (e.g. Quézel 1953; Rivas-Mmifnez 1961; Rivas pes but in some cases reaching the oro­ Goday & Mayor 1966; Prieto 1971; Valle 1985; MediteITanean, which had been completely ignored, Rivas-Martfnez et al. 1986; Losa-Quintana et al. or has only been briefly discussed by previous 1986; Martfnez-Parras et al. 1987a, 1987b; Mota & authors. Valle 1987; Molero-Mesa 1988; Pérez-Raya et al. 1990; Molero-Mesa et al. 1992; Aallali et al. 1998 Study area and Molero-Mesa 1999). However, none of these The Sierra Nevada massif (Fig. 1) is situated in works deals with the hygrophilous vegetation, with the south-eastern Iberian Peninsula and belongs to the exception of Casares et al. (1986), Martinez­ two politica! provinces, Granada and Almeria. It has PaITas et al. (1987c) and Losa-Quintana et al. an ovai shape with an E-W main axis. Altitudes (1987), who studied the cryoro-Mediterranean and range from less than 700 m at the AlpujaITas leve! oro-MediteITanean peat-bog communities (the typi­ up to 3.481 m (Mulhacen peak, the highest moun­ cal Nevadense "borreguil", a high-mountain tain of the Iberian Peninsula). hygrophilous pasture) as well as the riparian com­ The massif, which belongs to the inner areas of munities of the Geni! River basin. the Baetic Mountains, was formed during the Alpine The studies of the riparian vegetation in other folding and is lithologically heterogeneous. The basins with Nevadense tributaries, such as those of peripheral areas ofthe range have a limestone edge, the rivers Andarax (Salinas & Bianca 1996) and whereas the centrai siliceous nucleus is dominated Guadiana Menor (Salazar 1996; Salazar et al. 1999), by mica-schists, quartzites and phyllites, clearly provided new data on traditionally little-studied phy­ defining what is known as the Nevadense sector. tocoenoses, but generally only single floristic, ecolo­ Watercourses flowing through Sierra Nevada gica! and landscape aspects were considered. correspond to two hydrographic basins: the In this study we will discuss data conceming Guadalquivir, which is the main river of the

17 C. SALAZAR et al.

Mediterranean region, gathering over 7% of the Mediterranean flora in just O.O l % of its surface (Bianca et al. 1998). The Sierra Nevada's rate of endemie taxa is outstanding: the massif constitutes one of the most important "bot spots" in Europe, with endemicity rates reaching 30-40% in culminant N areas or even surpassing 80% in certain ecosystems (Bianca et al. 1998). Nearly 6% of the flora (116 taxa) is endangered, in tbe majority of cases being A included in the IUCN's bigher risk categories. Over s 30 species are found in the edapho-hygropbilous vegetation of the Nevadense sector described in this paper. These taxa are partly protected by regional, national and intemational laws on account of their relevance.

Data and methods southern part of the Iberian Peninsula, with the The vegetation was studied following the phyto­ Geni! and Guadiana Menor as main tributaries, and sociological method of the Zurich-Montpellier the South basin, flowing directly into the school (Braun-Blanquet 1979), as modified by Mediterranean Sea, with the Guadalfeo and Géhu & Rivas-Martinez (1982). Relevés of the Andarax as most important rivers. edapho-hygrophilous communities of banks and According to Rivas-Martinez et al. (1997), watercourses were carried out between 1993-1999 Sierra Nevada biogeographically belongs to three in different localities of the study area. sectors of the Baetic province. The limestone edge In the tables, species have been generally cluste­ and the southern slopes belong to the Malacitano­ red into two groups: characteristics of association Almijarense and Alpujarrefio-Gadorense sectors plus higher syntaxa, and accompanying species. respectively, whereas our study area, consisting of Exceptionally, other groups such as characteristics the centrai siliceous part, belongs to the Nevadense of subassociation, cbaracteristics of variant, etc. sector. The latter is divided into two districts: bave been included. Nevadense - mostly towards the West and to which The relevés bave been classified according to Sie1rn Nevada pertains - and Filabrico - mostly phytosociological criteria, revealing the associa­ towards the East, to which the Sierra de Filabres tions present in the area. The relevés, wbich could pertains. Although this study deals with the vegeta­ not be related to any known syntaxon, have been tion of rivers and streams of both districts, the ripa­ described and classified as new associations and rian vegetation is better developed in the Nevadense subassociations. Wbere information was scarce, district due to higher rainfall, whereas in the they have been referred to as "communities". Filabrico it is scarcer and poorer as the water level Each association has been described by taking is slightly lower. tbe following elements into consideration: structure, Bioclimatically, the Sierra Nevada is dominated ecology, physiognomy, ecologica! cbaracteristics, by the Mediterranean macrobioclimate of the pluvi­ chorology (specifying the biogeograpbical distribu­ seasonal type, with continental or oceanic influen­ tion up to the district leve! by using the classifica­ ces (Rivas-Martinez 1996), being characterised by a tion of Rivas-Martinez et al. 1997). Finally, the prolonged period of summer drought and precipita­ study deals witb tbe pbytocoenosis dynamics, dyna­ tions (rain or snow) restricted to winter and spring. mic significance and relationships with other com­ The following thermotypes are present: cryoro­ munities. Mediterranean, oro-Mediterranean, supra-Medi­ Two units, subassociation and variant, bave been terranean, meso-Mediterranean and, occasionally, considered to belong to a lower bierarchical rank thermo-Mediterranean, as well as the hyperhumid, than that of association. As for subassociation, the humid, subhumid and dry ombrotypes. concept of geographical race (Alcaraz 1996) bas From the floristic point of view, the Sierra been used wben tbe floristic combination does not Nevada massif is considered to be the most impor­ hold sufficient cbaracteristics to be considered as an tant area for plant diversity in the western association. On tbe otber hand, wben a community

18 Studia Geobot. 20 (200 l) - Hygrophilous vegetation of Sierra Nevada - is diversifiedwithin the same territory forbioclima­ developed on carbonated substrata and waters tic, edaphic, ecotonal situations, etc., it has been (Cratoneuron .fìlicinum and Anagallis tenella com­ considered as a variant. munity) was described by Rios (1996:521) from The nomenclature of syntaxa follows the Sierra de Segura (Sub-Baetic sector). International Code of Phytosociological Nomen­ In waterlogged soils it comes into contact with clature (Barkman et al. 1988). The phytosociologi­ the helophytic Caricetum camposii-cuprinae and cal typology used up to the suballiance leve! follows Festuca ampia grass communities (Euphrasio­ Rivas-Martinez et al. (1999). Festucetum amplae Martinez-Parras, Peinado et Nomenclature of taxa follows, when possible, Alcaraz 1987). In less humid areas it comes into Flora ibérica (Castroviejo et al. 1986, 1990, 1993a, contact with Juncion acuti/lori rush-beds or with the 1993b, 1997a, 1997b; Mufioz-Garmendia & more sciophilous Aquilegio-Ranunculetum grana­ Navarro 1998; Talavera et al. 1999), otherwise tensis Martinez-Parras, Peinado et Alcaraz 1987 Flora Europaea (Tutin et al. 1964-1980). grass formations. Monographs bave been used for Agrostis (Romero Caricetum camposii-cuprinae Salazar, Lorite, Cano et al. 1988), Carex (Lucefio 1994), Cirsium et Valle ass. nova (Tab. 2, holotypus rei. n. 9) (Talavera & Valdés 1976), Juncus (Fernandez­ An association dominated by large hemicryp­ Carvajal 1981, 1982a, 1982b, 1983) and Sa/ix (Diaz tophytes of the genus Carex occupying slopes Gonzalez & Llamas 1987). which are permanently humid due to water coming The taxa for which different criteria have been from adjacent streams. lt develops on base-poor, followed are indicated herein: peat-bog like substrata that are often removed by Digitalis purpurea L. var. nevadensis (Kunze) animals, which form numerous, permanently floo­ Amo [Flora Fanerogamica de la Peninsula lbérica ded micro-depressions. With optimum in the supra­ III:350 (1872)]. Festuca trichophylla (Ducros ex Mediterranean and oro-Mediterranean thermotypes, Gaudin) K. Richter subsp. scabrescens (Hackel ex it may occasionally be found above the meso­ Trabut) Catalàn & Stace [Anales Jard. Bot. Madrid Mediterranean belt. 50(2): 219 (1992)]. Lotus pedunculatus Cav. subsp. The most characteristic and predominant species granadensis (Zertovà) Molero Mesa & Pérez Raya is the Nevadense endemism Carex camposii, [La Flora de Sierra Nevada: 152 (1987)]. Luzula cam­ accompanjed by Carex cuprina, Juncus articulatus, pestris (L.) DC. subsp. nevadensis P. Montserrat Eleocharis palustris, Cyperus longus, Alchemilla [Anales Inst. Bot. Cavanilles 21(2)]. Scrophularia auriculata Loefl. ex L. [Sp. PI. 620 (1753)]. straminea, Primula elatior subsp. lofthousei, etc. The aggressive livestock action (mainly by cattle) makes it difficult to identify with clarity the abun­ Results and discussion dant different species of Cyperaceae, in many cases A) Spring, amphibious and peat-bog vegetation even preventing the community from reaching nor­ ma! development. Because of the predation of its Cratoneuron commutatum and Anagallis tene/la inflorescences, Carex camposii undergoes strong community (Tab. 1) vegetative propagation, thus increasing its cover. Bryo-cormophytic community in which the Furthermore, livestock activity favours the enrich­ moss stratum (Cratoneur on commutatum and ment in Agropyro-Rumicion crispi elements, above Philonotis seriata) forms a continuous carpet 5-1O ali in the less flooded areas. cm thick, on which abundant plants of Anagallis The new association was previously reported by tene/la develop. The presence of Pinguicula grandi­ Losa-Quintana et al. (1986:228) as "Carex campo­ flora in the community should also be highlighted. sii community", but they did not provide either It occurs in permanently waterlogged areas on relevés or information regarding its ecology and distric gleysols or along watercourses, carpeting floristic composition. It was also cited by Casares stones splashed by water. Its oligotrophic nature is et al. (1986:450) as a new association called revealed by the presence of Calliergonella cuspida­ Laserpitio longiradii-Caricetum camposii Molero­ ta and Parnassia palustris. Mesa et Pérez-Raya (unpublished), but these This community appears at least in the authors report neither relevés nor a description of Nevadense district (its presence in the Filabrico the community. Furthermore, Laserpitium longira­ district has yet to be detected) where it covers small dium was considered as a characteristic species, but areas in the upper meso-, supra- and oro­ this Nevadense endemie is not related to helophytic Mediterranean thennotypes. A similar community formations.

19 C. SALAZAR et al.

Tab. I - Cratoneuron commutatum andAnagallis tenei/a community

Rei. number 1 2 3 4 5 6 7 Altitude (m) 1300 2050 1690 1750 1710 1740 2000 Area (m2) 2 2 1 2 2 2 4 Average height (cm) 15 10 5 10 5 15 5 Cover (%) 100 100 100 100 95 100 75 Number oftaxa 15 15 14 17 18 19 17

Characteristics of association and higher units: Cratoneuron commutatum 1 3 4 3 3 4 1 Anagallis tenella 3 1 3 1 1 Philonotis seriata 3 1 1 1 2 1 Parnassia palustris 2 1 2 + + + Pinguicula grandiflora 2 2 1 + 1 Calliergonella cuspidata 2 1 Pholia cruda 1 Jungermannia exsertifolia cordifolia 1 Reboulia hemisphaerica

Accompanying species: Trifoliumrepens 1 2 1 + 2 + Sagina procumbens 1 2 1 1 2 I luzula campestris nevadensis 1 + + 1 + + Viola palustris paluslris 3 2 + + I Euphrasia wilkommii + 1 + + Ranunculus repens + + + + Prunella vulgaris 1 + Holcus lanatus + + + Lotus pedunculatus granadensis 2 + 1 Carex camposii + + + Juncus articulatus + + + lotus uliginosus 1 1 Hypericum tetrapterum + + Agrostis nevadensis 2 + Festuca tricophylla scabrescens 1 + Cerastium fontanum + + Primula elatior lofthousei + + Festuca ampia + + Epilobium obscurum + + lotus corniculatus 2 Carex lepidocarpa 1 Blackstonia per/oliata 1 Leontodon microcephalus Carex capi/laris

Sporadic specics: Re!. I: Epilobium parviflorum +: Dactylorrhiza elata sesquipedalis +: Cirsium pyrenaicwn micranthum +: Sonchus maritimus aquatilis +. Rel.2: Sedum melanantherum +. Rel.3: Juncus inflexus +: Lythrum portula +. Rei. 6: Leontodon ca,petanus nevadensis +; Mentha spicata+. Rei. 7: Galium sp. +: Gentiana verna sierrae +: Epilobium a/sinifolium, +. Localities: I: Granada: Rio Alhama de Lugros, Dehesa del Camarate, VG7716. 2: Almeria: Abrucena, Barranco de la Campana., WG1005. 3: Almeria: Fiiiana, barranco del Rosai, WF0706. 4: Almeria: Bayarcal, Arroyo Anchuelo, WG0002. 5: Almeria: Fiiiana, barranco del Rosai, WF0706. 6: Almeria: Laujar de Andarax, Barranco del Horcajo, WG0802. 7: Almeria: Abrucena, barranco de Pefia Horadada, WG1205.

20 Studia Geobot. 20 (2001) - Hygrophilous vegetati on of Sierra Nevada -

Tab. 2 - Caricetum camposii-cuprinae Salazar, Lorite, Cano et Valle ass. nova

Rei. number 1 2 3 4 5 6 7 8 9 10 11 Altitude (m) 2 1700 1690 1720 1550 2270 1540 1800 1800 1900 1900 2120 Area (m ) 10 10 10 15 15 10 2 2 50 4 6 Average height (cm) 50 100 50 50 50 50 25 25 25 40 60 Cover (%) 100 100 100 100 100 100 85 90 90 90 100 Number of taxa 16 16 22 15 9 15 10 18 28 10 24 Characteristics of association and higher units: Carex cuprina 4 2 3 4 4 3 1 2 2 1 3 Carex camposii 3 3 2 1 I 2 4 5 5 4 4 Veronica beccabunga 2 + + + I Juncus articulatus 1 1 1 Accompanying species: 1 Holcus lanatus I 2 I 2 + 1 1 2 2 Hypericum tetrapterum 1 + 1 + 1 1 2 1 1 Trifolium repens nevadense 1 1 2 2 2 Juncus injlexus I 1 + + + 1 2 21 1 1 Mentha longifolia 2 2 + Cirsium pyrenaicum micranthum 1 1 + 1 1 2 Lotus u/iginosus 2 2 2 2 Dactylorrhiza elata + + + 1 + Aquilegia vulgaris nevadensis + + 1 + 1 Primula e/atior loflhousei 1 1 1 2 1 Anthoxanthum odoratum 2 + + Agrostis castellana castellana + + + + Carum verticillatum + + + 2 Alchemilla straminea + 2 + I Ranunculus granatensis + 1 I Peucedanum hispanicum + + 2 Ranuncu/us repens 1 + 1 1 11 Geum urbanum 2 + Epilobium obscurum 2 + Anthriscus sylvestris + 1 + Myosotis decumbens teresiana + + + Prunella vulgaris 2 + Trifolium pratense 1 I Cerastium fontanum + + + Carex nigra + + Festuca ampia 2 2 Urtica dioica + + Mentha suaveolens + 1 Juncus ejjùsussubglomeratus 1 +

Sporadic species: Rei. I: Brachypodium sylvaticum +. Rel.2: Rumex conglomerat11s +; Lathyrus pratensis +. Rel.3: Aconitum vulparia neapolitanum +; Epilobium tetragonum tetragonum +; Cirsium x nevadense +. Rel.4: Scirpus holoschoenus +; Mentha spicata +; Rel.5: Nardus stricta l; Potenti/la nevadensis l; Lotlls glareosus 1; Juncus conglomeratus l; Carex lepidocarpa +; Galium verum verum +. Rel.6: Athyrium fìlix-foemina I. Rel.8: Epilobium atlanricum +; Montia fontana fontana +. Rel.9: Scirpus cernuus 2; Cirsium pyrenaicum longespinosum l; Cirsium pyrenaicum pyrenaicum I; Cochlearia megalosperma +; Ligusticum lucidum +; Parnassia palustris +; Juncus conglomeratus +. Rel.10: Stellaria alsine I. Rei. 11: Dryopterisfìlix-mas +; Poa trivialis +: Briza media l. Localities: I: Almeria: Paterna del Rio, Barranco de los Cerezos WG0303. 2 and 3: Almeria: Fifiana, Barranco del Rosai, WF0706. 4: Almeria: Bayarcal, Arroyo del Palancén, VG9801. 5: Almeria: Laujar de Andarax, cabecera del Barranco del Horcajo. WG0704. 6: Almeri a: Beires, ca becera del rio Ohanes, WG I 80 I. 7 and 8: Granada: Dehesa del Camarate, VG7614. 9: Granada: Dehesa del Camarate, VG7613. 10: Granada: Rio Barrio, VG8708. 11: Granada: Rio Dilar, VG6102.

21 C. SALAZAR et al.

The Caricetum camposii-cuprinae is a typical This association can be interpreted as a pioneer association of the Nevadense sector, being widely phase to the setting up of silicicolous Salix atroci­ distributed due to the large amount of gullies where nerea (occasionally, Salix caprea) willows since it peat-bog like soils are fmmed and Carex camposii occupies grave! along streambeds. When it occurs can proliferate. Its presence has been observed in on deeper soils, it may come into contact with other Sierra Nevada (Nevadense district) and in Sierras of Nevadense communities such as rushes ( Cirsio Baza and Filabres (Filabrico district), in the latter micranthi-Juncetum effitsi), megaforbic grass with less hygrophilous endemie species. (Aquilegio-Ranunculetum granatensis) and sedge In more sloped areas with 11011-waterlogged communities of flooded substrata (Caricetum cam­ soils, this helophytic formation comes into contact posii-cuprinae). with the megaforbic association Aconito-Sene­ Myrrhoidi nodosae-Alliarietum petiolatae Rivas­ cietum elodis Quézel 1953. Towards areas closer to Martinez et Mayor ex V. Fuente 1986 nepetosum rivers and streams, on compacted, flat, humid but granatensis subass. nova (Tab. 4, holotypusrei. n. 5) 11011-waterlogged soils, it usually comes into contact Synonym: Alliario petiolatae-Lase1pitietum longi­ with rush-beds of Cirsio micranthi-Juncetum effitsi radii Losa-Quintana 1986 Salazar, Cano et Valle in Salazar et al. 1999 and Scio-nitrophilous herbaceous community of Aquilegia nevadensis-Ranunculetum granatensis clearings within deciduous forests. It develops on grass communities. Under stronger nitrification by deep soils, with a certain nitrification and under livestock it comes into contact with rush-beds of cover of dead leaves. Of limited coverage and ave­ Cirsio-Juncetum inflexi Vigo 1968. Where the asso­ rage size, this community has its optimum in the ciation is directly in contact with streams and subhumid supra-Mediterranean belt. Phenolo­ creeks, these will be carpeted with watercress silici­ gically, it has a clear maximum in spring, when the colous communites (Glycerio declinatae-Apietum most important species (Alliaria petiolata and nodiflori J.A. Molina 1996). Myrrhoides nodosa) bloom; in summer they wither and sprout new shoots. The accompanying species B) Wood-fringe and megaforbic vegetation are Urtica dioica, Anthriscus sylvestris, Chae­ rophyllum hirsutum, Nepeta granatensis, Hera­ Ranunculo granatensis-Cochlearietum megalo­ cleum granatense, etc. spermae Salazar, Lorite, Valle et Cano ass. nova This association was first described in the (Tab. 3, holotypus rei. n. 6) Guadalajara province by Fuente (1986) but it seems to Thick megaforbic grass community developed have a wider distribution with its southernmost limit on siliceous rocks, mainly mica-schists, which corresponding to our survey area. Losa-Quintana et emerges within strongly flowing streams and clean, al. (1986: 181) consider the Alliario-Laserpitietum oligotrophic waters. It can occasionally appear on longiradii Losa-Quintana 1986 association as ende­ more or less developed humid soils, near streams mie to Sierra Nevada, but they had probably identified with a stable water level which never undergo a Laserpitium longiradium for L. gallicum or L. latifo­ deep decrease in their water leve!. With a supra­ lium, since the only localities of longiradium are in Mediterranean optimum, this association, in ce1tain L. cases, can descend to the meso-Mediterranean belt. the Monachi! River gully (Malacitano-Almijarense It occupies shaded areas, being absent in open sector), on scio-nitrophilous stratum limestone of ones. Occasionally, Cochlearia megalosperma evergreen oak groves (Bianca, pers. comm. ). appears in sunnier areas, being integrated with the Therefore, it seems more appropriate to consider this megaforbic grass communities of Senecionion flu­ community as a subassociation, called nepetosum viatilis. Faithful species to this formation are granatensis, suppmted by Baetic elements such as Myosotis decumbens subsp. teresiana, Hypericum Nepeta granatensis and Heracleum granatense, tetrapterum, Senecio jacobaea, Peucedanum hispa­ spreading at least over the Nevadense and Sub-Baetic nicum, Holcus lanatus and Lotus uliginosus, besides sectors, in relation to which similar communities have the two principal species. been already cited (Rios 1996). This association has been observed in Sierra This community, which has a rather short Nevada and could theoretically be found -although growing season, appears in the undergrowth of alder poorer in species- in Sierra de Filabres (Filabrico forests, athrocinereous willows and goat-willows of district), or even in the Serrano-Bacense district Sierra Nevada; when it occurs along streamsides, it (Baza Sierra). It may also be present in North comes into contact with Aquilegio-Ranunculetum Africa, because Cochlearia megalosperma is an granatensis communities or with different rushes or Iberian-North-African endemism. helophytic communities.

Studia Geobot. 20 (200 I) 22 - Hygrophilous vegetation of Sierra Nevada -

Tab. 3 -Ranunculo granatensis-Cochlearietum megalospermae Salazar, Lorite, Cano et Valle ass. nova

1 Rei. number 2 3 4 5 6 7 8 9 Altitude (m) 2 1600 1600 1980 1700 1520 1550 1700 1440 14601 Area (m ) 2 I 4 10 2 1 2 2 Average height (cm) 150 70 120 100 100 100 100 25 100 Cover (%) 90 100 95 90 100 100 100 80 90 Number of taxa 11 11 10 8 9 13 15 10 11 Characteristics of association and higher units: Cochlearia megalosperma 4 3 1 2 4 4 4 2 2 2 Hypericum tetrapterum + 2 2 2 + + 1 Peucedanum hispanicum 2 3 + 1 Senecio jacobaea + 1 1 Ranunculus granatensis 2 + + 2 Scrophularia auriculata l + Accompanying species: Holcus lanatus 2 2 3 4 4 4 3 Myosotis decumbens teresiana 1 2 1 1 + + Lotus uliginosus + l 1 1 + Mentha longifolia 1 l + 2 + Carex composi i 1 2 + Geranium purpureum + 2 + Brachypodium sylvaticum + + + Poa nemoralis 2 Urtica dioica + + Ruhus ulmifolius + + A thyrium jìlix-foemina 1 + Dactylorhiza elata sesquipedalis + + Cardamine jlexuosa l I Anthoxanthum odoratum 1 + Clinopodium vulgare arundanum + + Lonicera peryclimenum hispanica + Anthriscus sylvestris + Mentha suaveolens 3 Epilobium obscurum 2 Agrostis canina canina Juncus injlexus + Apium nodijlorum Juncus effusussubglomeratus + Ranunculus repens 2 Cerastium fontanum + Geranium columbinum 1 Hypochaeris radicata + Lapsana communis + Scrophularia scorodonia + Ligusticum lucidum Cirsium pyrenaicum micranthum + Euphorbia characias + Heracleum sphondylium granatense +

Localities: I: Almeria: Fiiiana,Arroyo de Lubeire, WG0907. 2: Almeria; Bayarcal, Arroyo Anchuelo, WG000 I. 3: Almeria: Paternadel Rio, Barranco de los Murillos, WG0304. 4: Almeria: Paternadel Rio, Barranco del Saltadero, WG040 I. 5: Granada Arroyo del Alhori, VG82 l I. 6: Granada: Arroyo del Alhori, VG8 Il I. 7: Granada: Barranco de las Rozas, VG76 I4. 8: Granada: Arroyo del Alhori, VG83I 2. 9: Granada: Rio Dilar, VG560I.

23 C. SALAZAR et al.

Tab. 4 -Myrrhoidi nodosae-Alliarietum petiolatae Rivas-Martinez et Mayor ex V. Fuente nepetosum granatensis subass. nova.

I Rei. number 2 3 4 5 Altitude (m) 2 1830 1830 1500 1600 1400 Area (m ) 25 25 4 2 4 Average height (cm) 40 40 25 25 30 Cover (%) 95 95 75 8511 80 Number of taxa 12 10 13 15 Charactcristics of association and higher units: Alliaria petiolata 3 3 2 3 2 Urtica dioica 3 3 + 2 2 Myrrhoides nodosa 1 + 1 Anthriscus sy/vestris 2 3 Galium aparine 1 2 Diffcrentialsof subass. nepetosum granatensis: HerrcleumsrJiorrlJJiwn grcndem! 1 2 Nepeta granatensis + 1

Accompanying species: Stellaria media 2 2 2 1 Saxifraga granulata 1 2 Ga/ium 3purium 1 1 Cerastium gibraltaricum + 1 Dactylis glomerata hispanica + 1 Centaurea triumfetti lingulata 1 + Geranium pyrenaicum 1 2 Cynosurus elegans 1 + Geranium molle 2 1 Geranium columbinum 1 1 Veronica an,ensis 2 1 Bunium alpinum 1 + Holcus lanatus + + Lapsanacommunis + + Silene latifolia 1 He//eborus foetidus 1 Nfarrubium supinum 1 Vulpia alopecuroides 1 Arum italicum 1 Doronicum plantagineum + Primula elatior lofthousei + Ranunculus granatensis +

Localities: 1-2: Almeria: Abrucena, Prox. del Barranco del Diablo, WG1206. 3-5: Granada: Dehesa del Camarate, WG71.

24 Studia Geobot. 20 (200 I) - Hygrophilous vegetation of Sierra Nevada -

C) Meadows and pastures thick sarmentous communities protect and offer the necessary shade for the sprouting of new trees, and Cirsio micranthi-Scirpetum holoschoeni Lorite, their degradation mainly leads to the appearance of Salazar, Cano et Valle ass. nova (Tab. 5, holotypus meso-Medite1rnnean rush-beds ( Cirsio micranthi­ rei. n. 8) Scirpetum holoschoeni). Round-headed club rush-bed (Scirpus holo­ There is scant information on this community. schoenus) accompanied by Cirsium pyrenaicum var. We provide the following relevé: micranthum occurs on siliceous substrata in areas of Locality: Granada, Narila, Rio Guadalfeo, VF8390. 2 the meso-Meditenanean thermotype, with soils Altitude: 980 m. Area: 30 m • Average height: 200 drying out in summer. cm. Coverage: 100%. Characteristics of association This new association is endemie to the Nevadense and higher units: Coriaria myrtifolia 4, Rubus sector, though it is rather poor in characteristic spe­ ulmifolius 2, Spartium junceum 2, Tamus communis cies. It is considered to be an edaphic vicariant of the 1, Euphorbia characias l, Brachypodium sylvati­ Cirsio-Holoschoenetum vulgaris Br.-Bl. 1931 (meso­ cum 1, Lonicera peryclimenum hispanica +. Mediterranean basophilic rush-beds). Acidophilous variant differentials: Adenocarpus lt appears in the dominion ofthe Nevadense wil­ decorticans 2, Salix pedicellata 1. Accompanying low-alder forests (Carici-Salicetum atrocinereae). species: Clematisjlammula 1, Do,ycnium rectum +, In drier areas it comes into contact with climatic for­ Mentha suaveolens +, Asparagus acutifolius +, mations, whilst in more humid areas it is related to Equisetum ramosissimum +. the helophytic Glycerio declinatae-Apietum nodi­ Rubo ulmifolù-Rosetum corymbiferae Arnaiz et ori communities. When a strong nitrification fi Rivas-Martinez in Arnaiz 1979 variant with occurs due to increased livestock action, this com­ Adenocarpus decorticans (Tab. 6) munity is possibly substituted by Cirsio-Juncetum Thick bramble communities surrounding supra­ inflexi rush-beds. Meditenanean willow and alder woods. Lianoid, climbing and thomy species such as Rubus ulmifo­ D) Serial shrub and wood-fringe vegetation lius, Crataegus monogyna, Rosa spp., Lonicera pe1yclimenum subsp. hispanica, Clematis vitalba, Rubo ulmifolii-Coriarietum myrtifoliae O. Bolòs Vitisvinifera, etc. are predominant. Other species of 1954 variant with Adenocarpus decorticans climatic vegetation needing great humidity, such as Thick, impenetrable bramble conununity domi­ Rhamnus catharticus, Prunus insititia, Berberis vul­ nated by Rubus ulmifolius together with numerous garis subsp. australis, etc. can also appear. lianoid and climbing species, e.g. Rosa spp., This association is distributed over a large area of Lonicera peryclimenum subsp. hispanica, Clematis the peninsular Northem Sub-Plateau, reaching the jlammula, C. vitalba, Coriaria myrtifolia and Tamus southernmost boundary through the eastern Baetic communis. It surrounds and substitutes the athrocinereous province. As it develops on base-poor substrata, in willow and alder woods in middle river tracks, or the Nevadense sector it is enriched in silicicolous appears along inigation channels. As it appears in species, such as Sa/ix caprea, Alnus glutinosa, areas of meso-Mediterranean thermotype, ther­ Scrophularia scorodonia, Pteridium aquilinum, mophilous species such as Dorycnium rectum, Salix Athyrium filix-foemina, and Adenoca,pus decorti­ pedicellata, Spartium junceum, Calystegia sepium, cans, which although not characteristic ofthe riparian Euphorbia characias, etc. are conunon. These for­ vegetation, is conunonly found in this community. mations are enriched in Adenocarpus decorticans The ecologica! significance and dynamics of which, although not a typical riparian species, is bramble communities within the ripicolous vegeta­ often linked to these bramble communities. tion has been previously discussed. In this case, the The Rubo-Coriarietum myrtifoliae has a wide only exception is that the rush-beds originated distribution in non-continental Ibero-Levantine ter­ through successive degradation and belong to the ritories. This silicicolous variant is rarely seen in the supra-Meditenanean rush-beds of Cirsio micranthi­ Nevadense sector, being more frequent in the Juncetum e[fitsi. Filabrico district or in the vicinity to the Alpu­ ja1Tefio-Gadorense sector, which are characterised E) Mediterranean and Euro-Siberian climactic by higher temperatures. vegetation As bramble patches typically sunound gallery forests, they become more vigorous when the remo­ Carici camposii-Salicetum atrocinereae Salazar, val of tree cover increase the light regime. These Lorite, Cano et Valle ass. nova

25 C. SALAZAR et al.

Tab. 5 - Cirsio micranthi-Scirpet11m holoschoeni Lorite, Salazar, Cano et Valle ass. nova.

Rel. number 1 2 3 4 5 6 7 8 9 Altitude (m) 2 1000 1200 1180 1140 1150 1090 1300 1080 1250 Area (m ) 8 60 15 15 20 12 30 40 24 Average height (cm) 75 70 100 120 80 75 70 120 70 Cover (%) 100 50 100 50 85 95 100 80 90 Number oftaxa 9 16 14 15 14 12 20 24 22 Characteristics of association and higher units: Scirpus holoschoenus 3 3 4 3 4 3 3 4 2 Cirsium pyrenaicum micranthum I + 1 1 2 l 2 2 3 Nfentha suaveo/ens 2 2 2 2 + 3 2 2 Holcus lanatus 1 1 + 1 l 1 1 1 2 l 3 Ranunculus repens 2 + + 1 1 Trifolium repens 1 1 1 3 21 1 Juncus injlexus 2 2 Lotus uliginosus + + 2 2 11 Agrostis castellana castellana + + 2 2 Mentha longifolia 1 l 1 1 Juncus ejfusussubglomeratus + I + Rumex conglomeratus + + + Hypericum tetrapterum I + Pulicaria dysenterica + 1 Dactylorrhizaelata + + Anthoxantum odoratum 2 Sonchus maritimus aquatilis + + Lotus corniculatus 1 2 Accompanying species: Brachypodium sylvaticum 1 1 1 1 1 1 1 Medicago lupulina 2 Euphorbia characias 1 + Epilobium parviflorum + + Apium nodiflorum + + Scrophularia scorodonia 1 + + Sa/ix atrocinerea + + + Artemisia campestris glutinosa I + 1 Festuca scariosa + + I Ononis spinosa + 1 Urtica dioica I l Cirsium vulgare + +

Sporadic species: Rei. l. Dorycnium rectum 2, Trifolium pratense, +, Plantago major +. Rei. 2. Lolium perenne I, Cynosunis elegans 1., Mantisalca salmantica +, Digitalis pwpurea nevadensis +. Rei. 3. Lotus pedunculatus granadensis 1, Cyperus longus 1. Epilobium hirsut11m +, Hypericum perforatum +. Rel. 4. Equisetum telmateia +. Rei. 5. Crepis vesicaria I, Dittrichia viscosa +. Rei. 6. Juncus bufonius +. Rei. 7. Lathyrus pratensis +, Festuca arundinacea +, Carex cuprina +, Carex camposii +, Juncus subnodulosus +. Rei. 8. Equisetum ramosissimum 2, Cynodon dactylon I, Mentha spicata 1, Veronica anagallis-aqua/ica 1. Picnomon acarna +, Senecio malacitanus +, Silene latifolia +, Helleborus foetidus +. Rei. 9. Ranunculus granatensis +, Sa/ix purpurea +. Rubus ulmifòlius +. Localities: I: Granada: Narila, Rio Guadalfeo, VF0998. 4 and 5: Almeria: Abrucena; rio Abrucena, WG 1608. 6: Almeria: Fiììana, rio Nacimiento, WG08 l l. 7: Granada Lugro, Rio Alhama de Lugros, VG77 l 6. 8: Almeria: Abrucena, rio Abrucena, WG 1708. 9: Granada: Lugros; Rio Alhama de Lugros, VG77 I 6.

26 Studia Geobot. 20 (200 I) - Hygrophilous vegetation of Sierra Nevada -

Tab. 6 - Rubo ulmifolii-Rosetum corymbiferae Rivas Martinez et Arnaiz in Arnaiz 1979 variant with Adenocarpus decorticans.

1 2 3 4 5 6 9 10 Rei. number 7 8 Altitude (m) 1620 1200 2 1120 1140 1380 1460 1520 1350 1440 1440 Area (m ) 100 100 100 50 40 30 160 35 20 1505 10 135 20 Slope (°) 45 20 5 Exposition N o NO SE s s N 150 250 200 200 200 200 Average height (cm) 250 250 175 175 Cover (%) 100 90 100 90 100 100 17 10011 759 809 75 Number of taxa 19 19 13 15 14 12 Characteristics of association and higher units: Rubus ulmifolius 4 4 4 3 4 5 2 5 4 3 Rosa corymbifera 2 2 3 3 2 1 4 2 Crataegus monogyna 3 3 3 1 2 Hedera helix 2 + 1 + Rosa canina 1 2 + Sa/ix atrocinerea 1 + 2 lonicera peric�ymenum hispanica 2 1 + Berberis vulgaris australis 2 1 1 Rosa pouzinii 1

Acidophilous variant differentials Adenocarpus decorticans + 2 1 1 2 l 1 + 1 I Athyrium jìlix-foemina + + + 1 Pteridiumaquilinum 1 1 + 2

Accompanying species: Helleborus foetidus + + 1 1 + + 1 + Euphorbia characias + + 1 + 1 Brachypodium sylvaticum + 1 1 Alnus glutinosa 2 1 Urtica dioica + 1 1 1 Rubia peregrina 1 + + 1 Scrophularia scorodonia + 1 + + Mentha suaveolens + + + 1 Festuca scariosa + + + Ballata hirsuta 1 + Helichrysum italicum serotinum + + Asparagus acutifolius + + Cirsium pyrenaicum micranthum + + Geranium purpureum + + Hypericum tetrapterum 1 + 1 Holcus lanatus + Rumex conglomeratus + +

Sporadic specics: Rei. l. Populus nigra italica I, Rumex induratus +, Artemisia campestris glutinosa+. Rel.2. Quercus rotundifolia 1, Pti/ostemon hispanicus +, Thymus mastichina +, Cytisus grandiflorus +. Rei. 4. Prunus ramburii I, Jug/ans regia +, Peucedanum hispanicum +, Festuca elegans +. Rei. 6. Campanula rapunculus +. Rei. 7. Cirsium pyrenaicwn longespinosum +. Rel. 8. Sonchus maritimus aquatilis +, Ranunculus repens +, AgrosNs canina canina 1, Equisetum ramossisimum +. Rel. 9. Prunus spinosa I. Rei. 1 O. C/emalis vitalba+, Rosa micrantha +, Rhamnus catharticus +. Localities: I and 2: Almeria: Abrucena, rio Abrucena, WG 1507. 3: ibidem; Barranco de la Mina, WG 1506. 4: Almeria: Laujar de Andarax, Barranco de los Tejos, WG0403. 5: Almeria: Bayacal, Arroyo del Palanc6n, VF9999. 6 and 7: Almeria: Beires, rio Ohanes, WG 1801. 8: Granada: Lugros, rio Alhama de Lugros, VG7716. 9 and IO: Granada: Arroyo Alhori, VG8312.

27 C. SALAZAR et al.

Tab. 7 - Carici camposii-Salice/um a/rocinereae Salazar, Lorite, Cano el Valle ass. nova

Rei. number I 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 Altitude (m, x 10) 2 146 160 120 140 160 185 168 164 144 115 155 122 150 170 170 162 100 150 Area (m ) 100 150 100 100 125 60 200 160 160 100 150 125 125 75 200 150 150 60 Average height ( cm) 400 600 500 200 900 300 250 900 900 900 900 800 400 900 600 600 600 300 Cover (%) 90 80 100 100 100 100 100 100 75 90 80 75 75 60 100 90 90 100 Numher oftaxa 20 26 14 24 15 22 18 21 16 24 19 14 18 13 23 22 31 38 Characteristics of assoc. and higher units: Sa/ix atrocinerea 4 4 3 5 2 4 4 4 3 2 3 3 3 3 4 4 2 4 + Carex camposii 2 + + 1 2 1 1 I 2 + 1 2 1 1 Alnus glutinosa 4 5 4 2 4 3 3 + Athyriumfilix-foemina 3 + I 1 2 1 + 2 2 2 2 1 Brachypodiumsy/vaticum I 2 2 I 2 + 1 + 2 Scrophularia scorodonia + + + + 1 + + + 1 Hedera helix + 2 + 1 + 1 Helleborusfoetidus + + I + 1 + E11phorbia characias 1 + + + Populus x canadensis 3 2 2 2 Populus nigra italica 2 2 2 2 Lapsana communis + + + + + Poa nemoralis + + Tamus communis + + Clinopodium vulgare + + + Origanum vulgare + + Sa/ix x quercifolia 2 Geum urbanum Sorbus aucuparia + Sa/ix fragilis 1 Origanum virens + +

Differentialsof subass. salicetos,un capreae: Prunus avium 1 1 + + Sa/ix caprea 3 2 2 Lonicera arborea + + + Acer opalus granatense 2 2 Sorbus aria 2 2 Betula pendula fontquen· l Rhamnus catharticus

Accompanying species Rubus ulmifolius + 2 2 3 3 3 2 2 2 2 2 2 Crataegus monogyna 1 2 1 2 2 1 Rosa corymbifera + 2 + + Laicau�hisp111im + + + Berberis vulgaris austra/is + + + Pteridium aquilinum + 2 Rosa pouzinii + Rosa canina + Clematis vitalba + + Spmtium junceum + +

Studia Geobot. 20 (200 I) 28 - Hygrophilous vegetation of Siena Nevada -

Tab. 7 - Continued.

Accompanying species: l Holcus lanatus 2 l I + l + Urtica dioica + I 2 2 2 2 2 + Digitalis purp. nevadensis + + + + + + + + + + l Peucedanum hispanicum 2 l + I 2 + l l Adenocarpus decorlicans + l l + + Hypericum tetrapten,m + + + + + + + Cochlearia megalosperma + + + + + Geranium purpureum + + + + + l Scirpus holoschoenus + + + + + Epilobium parviflorum + l + + Senecio jacobaea + 2 + Lotus uliginosus I + + Men/ha suaveolens 1 + + Ranunculus granatensis + + Cardamine jlexuosa + + l + Dacty/orrhiza elata + + + Cytisus grandiflorus + + 2 Mentha longifolia + Prunellavulgaris + + + .Juncus i1iflexus + + + My:N1isda:umbe.m temiJm + + + Ranunculus repens + Trifolium repens + + Sonchus mmii. aquatilis l + Hemdeum�gfrnlleme + + Dryopteris filix-mas + CiJ�1�1miaarthwn + + Carex cuprina Trifolium pratense + Tanacetum pmthenium + Rubia peregrina Ficus carica + +

Sporadic species: Characteristics of class: Rei. 4. Asplenium onopteris +. Rei. 5. Viola suavis +. Rei. 6. Populus tremula I, Fraxinus angustifolia angustifo/ia 1, Rosa montana 2. Rei. IO. Castanea sativa 2. Rei. 15. Rosa micrantha I. Rei. 18. Quercus pyrenaica +, Equisetum temalteia I. Accompanying species: Rei. I. Anthoxantum odoratum 1, Festuca arundinacea +. Rei. 2. Lathyrus pratensis 1, Agrimonia eupatoria 1, Festuca e/egans +, Agrostis castellana castellana+, Viola reichembachiana +, Rumex conglomeratus +,. Rei. 3. Equisetum ramossisimum +, Scrophularia canina canina+. Rei. 5. Cistus laurifolius +. Rei. 6. Anthriscus sylvestris +. Rei. 7. Potenti/la reptans +. Rei. IO. Apium nodiflorum +, Parietaria judaica +. Rei. 14. Aconitwn burnatii +. Rei. 15. Aquilegia vulgaris nevadensis I. Rei. 17. Bupleurumfruticosum 2, Agrostis stolonifera+, Trachelium caeruleum +. Localities: 1: Almeria: Beires, Rio Ohanes, WG1801. 2: Almeria: Fiiiana, Arroyo de Lubeire, WG6808. 3: Almeria: Laujar de Andarax. Rio Andarax, WF0999. 4:Granada: Capileira, a!luente del Poqueira, VF6891. 5: Granada: Aldeire, barranco de los Tejos, VG9308. 6: Granada: Rio Barrio, VG8708. 7: Almeria: Ohanes, Arroyo de Tices, WG1904. 8: Granada: Aldeire, arroyo de los Pasillos, VG9309. 9: Granada: Jeres del Marquesado, arroyo del Alhori, VG3812. IO: Granada: Bérchules. rio Guadalfeo, VF8391. 11: Granada: Lanteira, Rio Pueblo, VG8610. 12: Granada: Rio Alhama de Lugros, VG7819. 13: Granada: Dehesa del Camarate, VG7715. 14: Granada: Dehesa del Camarate, barranco de las Rozas, VG7614. 15: Almeria: Bayarcal. Arroyo Anchuelo, WG0002. 16: Granada: Rio Barrio, VG8809. 17: Granada: Rio Bermejo, VF7188. 18: Granada: Las Cebadillas, Arroyo afluente Poqueira, VF6894.

29 c. SALAZAR et al. subas. salicetosum atrocinereae (Tab. 7, rels. 1-12. et al. 1990; Molero-Mesa et al. 1992 and Salinas & holotypus rei. n. 11) Bianca 1996), generically referred to as "Salix atro­ subas. salicetosum capreae (Tab. 7, rels. 13-18. cinerea communities" without any precise syntaxo­ holotypus rei. n. 13) nomic arrangement, and in tbe majority of cases Thick, athrocinereous willow and alder forests even without tbe provision of pbytosociological developed on siliceous materials along streams of relevés. oligotrophic, permanently-flowing waters. Their Tbe new association seems to be endemie to tbe optimum is found in very deep gullies of the subhu­ Nevadense sector, where it is widely distributed, mid supra-Mediterranean, descending to the meso­ and is fragmentaryin tbe Filabrico district due to the Mediterranean thermotype, where they become blur­ lower water leve! of its streams. Tbe characteristic red because of the strong anthropic action. physiognomy of these communities, the existence Alder (Alnus glutinosa) is the predominant spe­ of some relevés witb a strong dominance of alder cies of this community when it occurs in close vici­ and the presence (sometimes with low cover values) nity of running water, bowever the greater part of the of species cbaracteristic of Osmundo-Alnion (Alnus biomass is more frequently formed by Salix atroci­ glutinosa, Salix atrocinerea, Scrophularia scorodo­ nerea. Other willows (e.g. Sa/ix fragilis) and poplar nia, Athyriumfilix�oemina) has led us to include the trees (Populus nigra, P tremula) may also be pre­ new association in the latter alliance, and more par­ sent. The species of tbe sbrub and herbaceous layers ticularly in the Osmundo-Alnenion suballiance, which better characterise these alder forests are whicb was previously restricted to tbe Ibero­ Athyrium .filix-foemina, Scrophularia scorodonia Atlantic province. Due to its isolation in relation to (typical of the alliance Osmundo-Alnion), other alder forests of the suballiance, the new asso­ Adenoca,pus decorticans (very common in the ciation can be regarded as very poor, even lacking Nevadense sector), Digitalis purpurea subsp. neva­ typical taxa such as Osmunda regalis, Clematis densis and Carex camposii (endemie). Tbe deep campani/fora, Galium broterianum or Carex brote­ shade and humidity ofthese environments allows for riana. The Nevadense athrocinereous willow and the growth of severa! ferns (Athyrium .filix�foemina, alder-tree communities bave been characterised by Pteridium aquilinum, D,yopteris.filix-mas, etc.). Carex camposii, a faithful taxon to the herbaceous The alder-tree forests of Siena Nevada have substratum present in both the meso- and supra­ been considerably altered by man, and it is often dif­ Meditenanean belts. ficultto find well developed, typical stands. In seve­ The salicetosum capreae subassociation develo­ ra! studies on the vegetation of the Nevadense mas­ ps in more humid situations (generally within the sif they seem to have been neglected. The existence subhumid dominion of the mesophytic Pyrenean of alder forests along the Nevadense riversides has oak series) as on the northern slope of the even been denied by Mmtinez-Parras et al. (1987a, Nevadense district. Interesting and rare taxa are pre­ 1987c). In other areas alder forests are probably of sent, such as Prunus avium, Betula pendula subsp. anthropic origin, but tbere are some evidences sup­ fontqueri, Salix caprea, Acer opalus subsp. grana­ porting their indigenate in Sierra Nevada. For tense, Taxus baccata, Sorbus aria, Lonicera arbo­ instance, the anthracitic remains found in the rea, Rhamnus catharticus, etc. For floristic and bio­ Millares site, which date back to the Copper Age, geographical reasons, this syntaxon can be properly can have no otber origin than the forests along tbe considered a subassociation and not a simple ecolo­ streams and rivers of the study area. At tbe same gica! variant typical of the Nevadense district but time, on the nmthernslope oftbis massifthere are a absent from the Filabrico district. few toponyma related to alder (Los Alisares of the The degradation of these gallery forests changes Pueblo River, tbe Aliso gully in tbe Granada the ecologica] conditions, with the bramble commu­ Alpujarra, etc). Certainly in tbe past alder was mucb nity taking advantage. The following modifications more abundant tban now, and nowadays a large patt are noteworthy: reduction in the pteridoflora, of alder forests can be interpreted as relictual. regression of Origanetalia vulgaris communities However, tbey sbould be considered as tbe climax and increase of Agropyro-Rumicion species as a of tbe Nevadense supra- and meso-Meditenanean result of increased light conditi on and more frequent riparian vegetation. livestock access. Contrary to alder forests, athrocinereous willow This association comes into contact with nume­ communities bave often been reported by previous rous herbaceous communities such as rush-beds autbors (Lasa-Quintana et al. 1986, 1987; Martinez­ ( Cirsio micranthi-Juncetum effi,si), megaforbic Panas et al. 1987a; Molero-Mesa 1988; Pérez-Raya communities (Aquilegio-Ranunculetum granaten-

30 Studia Geobot. 20 (200 I) - Hygrophilous vegetation of Sierra Nevada -

sis, Ranunculo-Cochlearietum megalospermae), Acknowledgements belophytic fo1mations (Caricetum camposii-cupri­ We wish to thank M" Carmen Maestro far her suppor! in the tran­ nae) and nitrophilous vegetation. slation of this paper.

Conclusions References

In the past the phytosociological studies on the Aallali A., L6pez-Nieto J.M., Pérez-Raya F. & Molero-Mesa J., hygrophilous vegetation of Sierra Nevada had tradi­ 1998. Estudio de la vegetaci6n foresta/ en la vertiente sur de Sierra Nevada (Alpujarra a/t11 granadina). lt. Geobot., 11:387- tionally been limited to pastures, peat-bogs, and 402. streams of the upper bioclimatic belts ( oro­ Alcaraz F., 1996. Fitosociologia integrada, paisaje y biogeografia. Meditenanean and cryoro-Meditenanean). In: Laidi, J. (ed.). Avances en Fitosociologia: 59-94. Servicio In this study we discussed some interesting plant Editoria! de la Universidad del Pafs Vasco, 19 I pp. Barkman J.J., Moravec J. & Rauschert S., 1988. C6digo de nomen­ communities, developed mainly in areas of the clatura fitosocio16gica (traducido por J. Izco & M. J. del Arco­ meso-Meditenanean and supra-Mediterranean ther­ Aguilar). Op. Bot. Phann. Comp. 4: 9-74. motypes but reaching in some cases the oro­ Bianca G., Cueto M., Martfnez-Lirola M.J., & Molero-Mesa J., 1998. Threatned vascular flora of Sierra Nevada (Southem Meditenanean, which had been completely ignored, Spain). Biol. Conserv. 85: 269-285. or had been only briefly discussed by previous Braun-Blanquet J., 1979. Fitosociologia. Ed. Blume, Madrid, 820 authors. pp. The following syntaxa (four associations, two Casares M., Pérez-Raya F., Molero-Mesa J. & Lasa-Quintana J.M., 1986. La vegetaci6n riparia en Sierra Nevada, I: la cuenca alta subassociations, two variants and one community) del rio Geni}, Catenas edaficas. Ars Phannac. 27: 447-453. needed to be described as new: Castroviejo S., Lafnz M., L6pez-Gonzalez G., Montserrat P., 1. Caricetum camposii-cuprinae Salazar, Lorite, Muiìoz-Garmendia F., Paiva J. & Villar L. (eds.), 1986. F/orn Cano et Valle ass. nova Ibérica. Plantas Vasculares de la Peninsula lbérica e Islas Baleares. Voi. I: Lycopodiaceae-Papaveraceae. 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Real Jardfn Botanico, community CSIC, Madrid. 320 pp. The majority are endemie to the Nevadense sec­ Castroviejo S., Aedo C., Benedf C., Lafnz M., Muiìoz-Garmendia F., Nieto Feliner G. & Paiva J. (eds.), 1997b. Flora Ibérica. tor, and are better developed in the Nevadense Plantas Vasculares de la Peninsula Ibérica e Islas Baleares. district. They have an important rate of relict and Voi. VIII: Haloragaceae-Euphorbiaceae. Real Jarclfn Botanico, endemie species, and host several species with an CSIC, Madrid. 375 pp. Dfaz Gonzalez T.E. & Llamas F., 1987. Aportaciones al conoci­ arctic-alpine disjunction. miento del género Salix L. (Salicaceae) en la provincia de A suitable interpretation of the vegetation dyna­ Le6n (NW Espaiia). Acta Bot. Malac., 12: I I 1-150. rnics of these little-known plant communities, Fernandez-Carvajal M.C., 1981. Revisi6n del género Juncus L. en which usually occur in fragile, endangered environ­ la Peninsula Ibérica I. Categorfas supraespecfficas y clave para las especies. Anales Jard. Bot. Madrid 38: 79-89. ments, should be considered of major interest. With Fernandez-Carvajal M.C., 1982a. Revisi6n del género Juncus L. en this knowledge, it will be possible to carry out an la Penfnsula Ibérica II. Subgéneros Juncus y Genuini accurate study of the N evadense vegetation series Buchenau. Anales Jard. Bot. Madrid 38: 417-467. Fernandez-Carvajal M.C. 1982b. Revisi6n del género Juncus L. en and geoseries and to offerpossible solutions for its la Peninsula lbérica III. Subgéneros Subulati Buchenau, restoration. Pseuclotenageia Krecz. & Gonstch. y Poiophylli Buchenau.

31 C. SALAZAR et al.

Anales Jard. Bot. Madrid 39: 79-15 I. Rivas-Martfnez S., 1961. Los pisos de vegetaci6n de Sierra Fernandez-Carvajal, M.C. l 983. Revisi6n del género Juncus L. en Nevada. Bo!. Real. Soc. Esp. Hist. Nat. 59: 55-64. la Penfnsula Ibérica IV. Subgéneros Juncinella (Fourr.)Krecz. Rivas-Martfnez S., 1996. Clasificaci6n Bioclimatica de la Tierra. & Gostch., Septati Buchenau y Alpini Buchenau. Anales Jard. Folia Botanica Matritensis 6: 1-32. Bot. Madrid 39: 301-379. Rivas-Martfnez S., Ferniindez-Gonzalez F. & S,fochez-Mata D., Fuente V. de la, 1986. Vegetaci6n or6flla del occidente de la pro­ 1986. Datos sobre la vegetaci6n del Sistem,1 Centrnl y Sierra vincia de Guada/ajara (Espai'ia). Lazaroa, 8: 123-219. Nevada. Op. Bot. Pharm. Comp., 2: 3-136. Géhu J.M. & Rivas-Martfnez S., 1982. Notions fondamenta/es de Rivas-Martfnez S., Asensi A., Dfez-Garretas B., Molero-Mesa J. & Phytosociologie. Ber. Internat. Symp. IA VS, Syntaxonomie: Valle F., 1997. Biogeographical synthesis of Andalusia 1-33. (Southem Spain). J. Biogeogr. 24: 915-928. J.M. M. Losa-Quintana , Molero-Mesa J. & Casares , 1986. El pai­ Rivas-Martfnez S., Fernandez-Gonzalez F. & Loidi J., I 999. sa_je vegetai de Sierra Nevada: la cuenca alta del rio Geni/. Checklist of plant communities of Iberian peninsula, Balearic Secret. Pubi. Univ. Granada, 285 pp. and Canaiy islands to subal/iance leve/. It. Geobot., 13: 353- M., Losa-Quintana J.M., Molero-Mesa J., Pérez-Raya F. & Casares 45 I. 1987. Comunidades higr6filas del piso supramediterràneo en Romero A.T., Bianca G. & Morales C., 1988. Revisi6n del género la cuenca del rio Geni/. Secr. Pubi. Univ. La Laguna. Serie Agrostis L. en la Penfnsula Ibérica. Ruizia 7. CSIC, Madrid, lnformes, 22: 217-222. Tenerife. 160 pp. Luceiio M., 1994. Monogratìa del género Carex en la penfnsula Salazar C., 1996. Aproximaci6n al conocimiento de la vegetaci6n Ibérica e Islas Baleares. Ruizia 14. CSIC, Madrid. 139 pp. J.M., M. riparia de los rios nevadenses en la cuenca del Guadiana Martfnez-Parras Peinado & Alcaraz F., 1987a. Menar (rios Alhama de Lugros y Guadix). Actas I er Comunidades vegeta/es de Sierra Nevada (Espaiia). Serv. Conferencia Internacional Sierra Nevada, Voi. Il: 353-372. Pubi. Univ. Alcala, Madrid, 74 pp. J.M., M. 608 pp. Martfnez-Parras Peinado & Alcaraz F., 1987b. Datos Salazar C., Garcfa-Fuentes A., Torres J.A., Melendo M., Valle F. sobre la vegetaci6n de Sierra Nevada. Lazaroa 7: 5 I 5-533. & Cano E., 1999. Comunidades de Juncus effususL. en Sierra Martfnez-Parras J.M., Molero-Mesa J., Peinado M. & Pérez-Raya Nevada (Espillia). Quercetea, I: 117-122. F., 1987c. La vegetaci6n foresta/ de riberas en la provincia de Salinas M.J. & Bianca G., 1996. Vegetaci6n foresta/ riparia en la Granada. Secr. Pubi. Univ. La Laguna. Serie Informes, 22: 55- provincia de Almerfa (SE. Espaiia).Monogr. F1. Veg. Béticas 66. Tenerife, 397 pp. 9: 57-95. Molero-Mesa J., 1988. 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Received October 18, 2000 Accepted May 28, 2001.

32 Studia Geobot. 20 (200 I) Studia Geobotanica. Voi. 20: 33-39 (2001)

DUE NUOVE ASSOCIAZIONI DI MARGINE STRADALE DELL' ABRUZZO (ITALIA CENTRALE)

GianfrancoPIRONE

Dipartimento di Scienze Ambientali, Università degli Studi, Loc. Coppito, Via Vetoio s.n., I - 67100 L'Aquila

Keywords: Abruzzo, ltaly, roadside vegetation.

Abstract: Two NEW ASSOCIATIONS OF ROADSIDE VEGETATION FROM ABRUZZO (CENTRAL lTALY). This study descri­ bes the perennial nitrophilous and meso-igrophilous vegetation, occurring along the main roads of Abruzzo up to 900 m a.s.l. Two roadside phytocoenoses are described, one with Senecio inaequidens and the other with P/umbago europaea. They are attributed to two new associations of the Artemisietea vulgaris class, Artemisia absinthii-Senecionetum inaequidentis, belonging to the Dauco-Me/ilotion alliance (Onopordetalia acanthii arder) and Galio albi-P/umbaginetum europaeae, belonging to the Inu/o viscosae-Agropyrion repentis alliance (Agropyretalia repentis arder).

Introduzione

La presente ricerca ha per oggetto alcuni aspetti Il presente studio vuole contribuire, quindi, a della vegetazione di margine delle strade statali e colmare le lacune relative agli aspetti più mesofili e provinciali dell'Abruzzo e dei territori confinanti nitrofili della vegetazione dei margini stradali. del Molise. I dati di letteratura relativi a questo tipo di vege­ Materiali e metodi tazione, con riferimento alle regioni adriatiche dell'Italia appe1minica, sono piuttosto frammentari e Lo studio è stato condotto con il metodo fitoso­ riguardano soprattutto le comunità vegetali delle stra­ ciologico (Braun-Blanquet 1964). I rilievi sono stati de di campagna e delle aree urbanizzate, dove il suolo eseguiti in un intervallo attitudinale compreso tra O viene rimosso periodicamente e si affennano fitoce­ e 900 m circa nel periodo autunnale (settembre-otto­ nosi afferenti in modo preponderante alle classi bre), quando questa vegetazione presenta la massi­ Stellarietea mediae e Molinio-Arrhenatheretea (ordi­ ma espressività sia vegetativa che antesica. ne Plantaginetalia majoris), e solo in minor misura Gli spettri biologici e corologici ponderati sono alla classe Artemisietea vulgaris. Ne sono esempi le stati calcolati servendosi dell'indice di ricoprimento associazioni Rumici-Carduetum pycnocephali, specifico, che esprime e valuta per ciascuna specie Hordeetum murini, Avena barbatae-Brometum dian­ presente nella tabella fitosociologica la sua copertu­ dri per le Marche (Hruska 1985, 1989, 1998; Biondi ra nell'ambito dell'associazione. Esso si calcola & Baldoni 1991); Lolio-Plantaginetum, Hordeetum sommando i valori centrali degli indici di abbon­ murini, Brometum sterilis, Aveno-Brometum diandri, danza-dominanza, dividendo per il numero di rilie­ Diplotaxi tenuifoliae-Agropyretum repentis per vi della tabella e moltiplicando questo valore per l'Abruzzo (Pascetti & Veri 1983; Pirone & Ferretti 100 (Braun-Blanquet 1964; Pirola 1970). 1999). La vegetazione indagata in questo studio è al La nomenclatura delle specie è, salvo eccezioni, contrario condizionata dagli interventi di manuten­ quella riportata in Pignatti (1982). zione delle strade, consistenti in particolare nello sfal­ cio periodico. Questa pratica, assieme alla buona Risultati disponibilità idrica dell'habitat favorita dal transito veicolare e dalle cunette laterali di drenaggio, per­ Nel territorio indagato la vegetazione lungo le mette l'affennazione di un elevato numero di piante strade si distingue, sul piano ecologico-floristico, in perennanti (emicriptofite, geofite e camefite) a carat­ due tipologie di cui una è a diretto contatto con la tere più o meno nitrofilo e mesoigrofilo, il cui massi­ sede stradale asfaltata ed è fonnata da una stretta mo sviluppo viene raggiunto in autunno. fascia pianeggiante, l'altra è insediata lungo il ver-

33 Gianfranco PIRONE sante oltre la cunetta (Fig. 1). di manutenzione dei margini stradali favoriscono il li seguente schema chiarisce la posizione sintas­ mantenimento nel tempo di questa particolare sonomica delle fitocenosi descritte. comunità vegetale che costituisce, quindi, un aspet­ to peculiare ed affermato della vegetazione legata a Artemisietea vulgaris Lohm., Prsg. & Tx. 1950 tale tipo di habitat a determinismo antropico. ampi. Riv.-Mart. et a/ii 1991 Per questa vegetazione, sulla base dei rilievi di Onopordetalia acanthii Br.-81. & Tx. 1943 em. Tab. 1, viene proposta una nuova associazione Goers 1966 denominata Artemisia absinthii-Senecionetum inae­ Dauco-Me/ilotion Goers 1966 quidentis (rilievo tipo n. 5 di Tab. 1 ), di cui si indi­ Artemisia absinthii-Senecionetum inaequidentis cano come specie caratteristiche e differenziali ass. nova Senecio inaequidens, Polygonum aviculare, Agropyretalia repentis (Oberd. et a/ii 1967) Cynodon dactylon, Verbascum sinuatum e Muell. & Goers 1969 Onob,ychis viciifolia. In particolare, Polygonum, Jnulo viscosae-Agropyrion repentis Biondi & Cynodon e Verbascum sottolineano il disturbo lega­ Allegrezza 1996 to al calpestio. La vegetazione raggiunge l'optimum Galio a/bi-Plumbaginetum europeae ass. nova di sviluppo in autunno. elytrigetosum athericae subass. nova L'associazione viene inquadrata nell'alleanza elytrigetoswn repentis subass. nova Dauco-Melilotion e nell'ordine Onopordetalia acanthii, che riuniscono le vegetazioni ruderali ter­ A rtemisio absinthii-Senecionetum inaeq uidentis moxerofile da bienni a perenni, legate a suoli scar­ ass. nova samente o mediamente dotati di nitrati (Poldini 1989). La fisionomia della vegetazione a contatto con la Come proposto da alcuni Autori, la vegetazione sede stradale è conferita principalmente da Senecio nitrofila e subnitrofila perenne viene riunita nella inaequidens e, in secondo luogo, da Artemisia classe Artemisietea vulgaris, a distribuzione eurosi­ absinthium, A. vulgaris, Diplotaxis tenui/olia e beriana, che penetra frequentemente nella regione Achillea collina. mediterranea, su suoli freschi e ricchi in sostanza Senecio inaequidens è una neofita originaria del orga111ca. Sudafrica, osservata per la prima volta in Italia pres­ L'associazione è caratterizzata da un buon livel­ so Verona nel 1947 (Pignatti 1982) e diffusasi rapi­ lo di mesofilia, sottolineata dal contingente di spe­ damente in tutto il centro-nord. In Abruzzo è stata cie della classe Molinio-Arrhenatheretea (Dactylis osservata per la prima volta negli anni '70 glomerata, Taraxacum officinale, Plantago lanceo­ (Anzalone 1976) ed è diventata comune negli lata, Arrhenatherum elatius, ecc.), favorito dal ambienti ruderali e nelle aree incolte (Tammaro & periodico sfalcio della vegetazione per manutenzio­ Pirone 1980; Guarrera e Tammaro 1994; Conti ne e dall'acqua proveniente dalla sede stradale 1998; osservazioni inedite dell'Autore). Nel territo­ durante i periodi di pioggia. rio indagato la vegetazione meglio caratterizzata da I rilievi dal n. 4 al n. 10 sono più ricchi di specie Senecio inaequidens è quella dei margini delle stra­ della classe Stellarietea mediae e sottolineano un de, dove spesso domina nettamente e si comporta, più marcato rapporto con la vegetazione di questo generalmente, da camefita suffruticosa. Le pratiche sintaxon.

4 2 2 3 4

Fig. I - Disposizione, lungo i margini stradali, della vegetazione studiata. I: sede stradale; 2: Artemisia absinlhii-Senecionelum inaequiden­ lis; 3: cunetta; 4: Ca/io albi-Plumbagine/11111 europaeae. Schemalic diagram ofthe dislrib11tio11 (Jf/hetwo phytocoenoses along a road seclion. I: road; 2: Artemisio absinthii-Senecionetum inacquidentis; 3: mwgin; 4: Galio albi-Plumbagineturn europaeae.

34 Studia Geobot. 20 (200 I) - Associazioni di margine stradale in Abruzzo -

Tab. 1 -Artemisia absinthii - Senecionetum inaequidentis ass. nova.

Elemento Forma N. del rilievo 1 2 3 4 5• 6 7 8 9 10 CLASSE Indice coro logico biologica Attitudine(m) 800 140 570 645 670 670 470 765 740 570 PRE- DI di Esposizione SEN- PRE- ricòpr. ° Inclinazione( ) ZA SEN- specif. Copert. vegetaz. (%) 100 90 100 100 100 100 100 100 100 100 ZA Superi. rilev.(mq) 4 10 20 3 4 3 6 3 25

Car. e diff. di associazione Slldafrica (nat.) Ch suffr Senecio inaequidens 3 4 4 4 4 4 3 3 4 10 V 5250 Cosmop. T rept Polygonum aviculare + + + + + 7 IV 100 Cosmop. Gmiz Cy nodon dacty lon 2 + 5 lii 235 Euri-Medit. H bienn Verbascum sinuatum + + + + 4 40 Medit. -Mont. H scap O nobrychis viciifolia + + + 3 30

Car. Dauco-M•lilotion/Onopordet1li• 1canthii Eurosib. H scap Picris hieracioides 2 + + + + 8 IV 390 Paleotemp H bienn Daucus carota + + + + + + 7 IV 85 Circumbor. H scap + Artemisia vulgaris 2 2 2 6 lii 635 Subcosmop. Ch suffr Artemisia absinthium 2 2 2 + 6 lii 510 Eurasiat. H scap Unaria vulgaris + + + 3 30 S-Medrt. H scap Foeniculum vulgare + 2 35 Cosmopol. H scap Cichorium intybus 25 Europ. H scap Reseda lutea 10 Europ. H bienn Pastinaca sativa ssp. urens + 10

Car. Arlemisi•t•• vulg•ris C-Europ. H bienn Anthemis tinctoria + + + 7 IV 255 Submedit.-Subatl. H scap Diplotaxis tenuifolia 1 4 11 450 SE-Europ. H scap Achillea collina 2 2 4 350 Asia orient. (nat.) H scap Artemisia verlotorum 2 4 225 Paleotemp. G miz Convolvulus arvensis + + + + 4 40 SL1bcosmop. H scap Rumex crispus + + + 4 55 Pateotemp. H scap Silene alba + + + 3 30 Paleotemp. H scap Verbena officinalis + 3 60 Euri-Medrt. Gmiz Sambucus ebulus + + 2 20 Circumbor. Gmiz Elytrigia repens + + 2 20 Subpontica H scap Chondrillajuncea + + 2 20 Circumbor. Gmiz Equisetum ramosisslmum + 2 35 Euri-Medrt. H scap BaUota nigra 25 Eurasiat. G rad Cirsium aivense + 10 SLtbpontica H scap Atthaea cannabina + 10 Steno-Medit. G rad Plurnbago europaea + 10 Steno-Medit. G miz Arundo pliniana + 10 Eurosiber. H scap Saponaria officinalis + 10

Car. St•ll•ri•t•• medi•• Euri-Medit. T scap + Brom us diandrus 2 2 7 IV 410 Eurosib T scap Malva sylvestris + + + 7 IV 255 E uri-Medn. -S-Siber. T scap Lactuca seniola + + + + + + 6 60 Euri-Medit T scap Avena barbata + + + + 90 Subcosmop. T scap Erodium cicutarium + + + 4 55 Europ. H bienn Echium vulgare + + + + 4 40 E uri-Medit.-T uran. T scap Dasypirum villosum + + 3 45 Circumbor. T scap Atriplex patula 2 + + 3 170 Subcosmop. T scap Chenopodium album + + 2 20 Eurasiat. T scap Sonchus oleraceus + + 2 20 Euri-Medrt. H bienn Echium italicum + + 2 20

Car. Molinio-ArrhenetheNt•• Paleotemp. H caesp Dactylis glomerata 2 2 2 2 7 IV 800 Circumbor. H ros Taraxacum officinale 1 + 2 + 7 IV 270 Euras1at. H ros Plantago lanceolata 2 + 2 5 lii 360 Paleotemp. H caesp Arrhenatherum elatius 2 + + 3 Il 170 Paleotemp. H ros Potentilla reptans + 2 35 Eurosiber. + H scap Trtfolium pratense + 2 20 Car. Fesfuco-Srometea Eurasiat H scap Galium verum + 3 60 Medit.-Mont. H scap Calamintha nepeta + + 2 20 Euri-Medit. T scap Odontites lutea + + 2 20

Altre specie Eurasiat. H scap Medicago sativa + + 8 IV 295 + Euri-Medit NP Rubus ulmifolius + 2 I 20 SE-Europ.-Caucas. T scap Cephalaria transsy!vanica + 2 20 Steno-Medit. H scap Reichardia picroides + 2 35 Europ. -Caucas P Han Clematis vitalba + 2 35 Steno-Medit. H bienn Seseli tortuosum + + 2 20

35 - Ga/io albi - Plumbaginetum europaeae ass. nova o

o �mento Forma N. del rilievo 2 3 4 5 6 7' 8 9 10 11 12 13 14 15 16 17 18 19 20 CLASS Indice � ologico biologica Altrtudine (m) 20 250 270 290 280 480 700 585 840 860 700 520 500 330 460 880 885 825 740 330 PRE- DI di PRE- .o Esposizione s E NE NE NE w sw SE w E E NE SSE SE E w ENE SEN- PRE- ricopr. SEN- Inclinazione C) 30 30 30 15 40 15 10 20 20 30 30 25 20 15 25 30 30 ZA SEN- specif. ZA o Copert. vegetaz. (%) 100 100 100 100 100 100 100 100 100 100 100 100 100 100 100 90 100 100 100 100 (A) ZA (A) (6) '6 Superf. rii. (mq) 8 12 6 5 5 5 5 6 6 6 6 12 6 6 8 5 6 6 15 8 (A)

ff. di associazione edlt. G rad Plumbago europaea 4 3 4 3 4 3 4 3 3 3 4 4 3 3 3 3 4 3 4 4 4 IV 5250 15 H scap Galium album 2 + + 2 2 + 1 + + + + 2 320 11

,ass. elytrigetosum athericae (A) G miz Elytrigìa atherica 2 3 3 2 2 5 V 2150 llt. T scap Raphanus raphanistrum + + + + 4 IV 80 lt Tscap Beta vu lgaris + + + 3 lii 60 T scap Sinapis alba + 3 lii 120 ,drt T scap Sonchus tenerrimus 2 + lii 320

ass. eiytrigetosum repentis (B) "· G rhiz Elytrigia repens 4 5 2 3 50 9 H scap Eryngium campestre + + + + + + + + H scap Galium verum + + + + ,p. H scap Hypericum perforatum + + + + 4 H scap Achillea collina + + + 4 o ,dlt. H scand Convolvulus elegantissimus + + 2 '-O (.) o viscosae-Agropyrion repentis / dalia repentis ,p G miz Convotvulus arvensis + + + + + + + + + + + 4 IV 110 10 i:3 p. H bienn Daucus carota + + 2 + + + + + + + 3 lii 90 8 op. H scap Rumex crispus + + + + + + + + + 20 9 H blenn Anthemis tinctoria + + + 50 5 -Subatl. H scap Diplotaxis tenurfolia + + + + + 20 :a H scap Chondrilla juncea + + + + 4 H rept Agrostis stolonifera 2 + 2 ,, G miz Equisetum ramosissimum 50 H scap Poa trivialis (nat.) Ch suffr Senecio inaequidens +

misietea vulgaris H scap Picris hieracioides + + + + + + lii 120 9 r. H scap Artemisia vulgaris 2 2 2 + 5 H scap Foeniculum vulgare + + + + 3 lii 90 3 op. H scap Urtica dioica + 3 + 4 H scap Unaria vulgaris + + + 3 t. Gmiz Sambucus ebu/us + + + 20 2 H scap Cichorium intybus + + + 2 40 1 Tscap Galium aparine + + + 3 op. Ch suffr Artemisia absinthium 2 2 dlt. H bienn Onopordon iltyricum + + 20 aleotemp. H scap Silene vulgaris + + 20 urasiat. G rad Cirsium arvense + + 20 uri-Medit. H scap Ballata nigra + ,at.) H scap Artemisia ver1otorum + 20 aleotemp. H bienn Cirsiumvulgare + 20 E-Asiat. H bienn lsatis tinctoria urasiat. Hscap Cruciata laevipes + E-Medlt. Ch frut Ballota rupestris 2

ar. Stellarietea mediae uri-Medlt.-S-Sib. Tscap Laducaserriola + + + 2 + + + + + + + + 1 1 + 3 lii 60 13 uri-Medit. Tscap Brom us diandrus 2 2 2 2 2 2 2 2 2 2 2 3 lii 650 12 uri-Medit. Tscap Avena barbata + + + + + + 4 IV 170 7 urì-Medit.-Turan. Tscap Dasypirum villosum 2 2 2 2 100 5 o lrcumbor. Tscap Fallopia convolvulus + + + + 6 N N ircumbor. Tscap Atriplex patula + + + + 5 .o2 ubcosmop. T scap Chenopodium album + + + + + 5 � edit.-Turan H bienn Carduus pycnocephalus + + + 4 .s urosib. T scap Malva sytvestris + + 3 "' urop. H bienn Echium vulgare + + + 3 "1 urastat. T scap Sonchus oleraceus + + + 2 40 1 "O "' -Medlt. T scap Papaver rtioeas + + + 3 b "'CJJ aleotemp. Tscap Mercurialis annua + 2 = 11r. Molinio-Arrhenatheretea i:-- "§i aleotemp. H caesp Dactylis glomerata 2 2 + + + 2 2 2 2 2 5 V 500 13 urasiat. H ros Plantago lanceolata + + + I 20 2 ircumbor. H ros Taraxacum officinale + + + 3 :a aleotemp. H ros Potentilla reptans + + 20 o .N ar. Fes-tuco-Brometea edrt.-Mont. H scap Calamintha nepeta + + + + 50 8 ·;:; o ubatl. H caesp Brachypodiumrupestre so 1 ndem. G rhiz Phleum amblguum 2 2 � ltre specie uri-Medit. NP Rubus ulmifollus + + + + + + 6 E-Eur.-Caucas. T scap Cephalaria transsylvanica + + + + + 40 3 ur.-Cauc:as. P lian Clematis vitalba + + + 4 teno-Medit. H bienn Seseli tortuosum + + + 4 edlt.-Atl. H sc:ap Salvia verbenaca + + + 4 Jri-Medit. T scap Tordlium maximum + + + + 4 :eno-Medit. Hsca.p Reichardia picroides + + + 20 2 :1leotemp. P ca.esp Crataegus monogyna + 3 JfaS�. H scap MedtcagO saUva + + 2 Jr. -Caucas. P caesp Prunus spinosa + + 20 :>smop. G rhiz Cynodon dactylon 50 :tleotemp. NP Rosa canina + + 2 Jbcosmop. H scap Agrimonia eupa1oria + + 2 eno-Medit. H scap Phlomis herba-venti + 2 Gianfranco PIRONE

La comunità con cui l'Artemisio-Senecionetum 2); è l'aspetto più termofilo, relativo ai territori sembra mostrare le maggiori somiglianze è il costieri e basso-collinari; Dauco-Picridetum hieracioidis Goers 1966, subass. elytrigetosum repentis (rill. 6-20), diffe­ dell'Europa centrale. La proposta di formalizzare le renziata da Elytrigia repens, Eryngiun campestre, differenzea livello di associazione vuole sottolinea­ Hypericum perforatum, Galium verum, Achillea re l'autonomia delle cenosi abruzzesi sia floristica, collina e Convolvulus elegantissimus (rilievo tipo n. evidenziata in particolare dalla presenza e/o consi­ 7 di Tab. 2); rappresenta le cenosi della fascia alto­ stenza delle specie caratteristiche e diferenziali, che collinare e montana. ecologica, rimarcata anche dall'elevata espressività fisionomica della vegetazione, mantenuta e favorita Considerazioni sugli spettri biologici e corologici nel tempo soprattutto dalle pratiche periodiche di sfalcio. Nello spettro biologico ponderato dell'Artemisia absinthii-Senecionetum inaequidentis dominano le Galio albi-Plumbaginetum europaeae ass. nova camefite (46,5%) e le emicriptofite (39,8%); anche le terofite sono relativamente ben rappresentate Lungo le scarpate laterali delle strade la fisio­ (1 O, 1 % ), mentre sono molto bassi i valori di geofite nomia della vegetazione è data da Elytrigia repens (3,1%) e fanerofite (0,5%). (= Agropyron repens), Elytrigia atherica (= Agro­ Lo spettro biologico ponderato del Galio albi­ pyron pungens auct. ital.) e Plumbago europaea. Plumbaginetum europaeae mostra invece una netta Quet'ultima è una geofita radicigemmata a distribu­ dominanza delle geofite (70,9% nella subass. elytrige­ zione steno-mediterranea, che forma, negli ambien­ tosum athericae e 57,2% nella subass. elytrigetosum ti studiati, popolamenti più o meno derisi, a fenante­ repentis), seguite dalle emicriptofite (13,8 e 23,8%). Il si autunnale. livello delle terofite (15,1 e 16,1%) è superiore all'as­ Questo secondo tipo di vegetazione viene asse­ sociazione precedente, mentre scarsa è la presenza di gnato, sulla base dei rilievi di Tab. 2, ad una nuova camefite (O e 1,9%) e fanerofite (0,2 e 1,0). associazione denominata Galio albi­ Lo spettro corologico ponderato dell'Artemisio­ Plumbaginetum europaeae (rilievo tipo n. 7 di Tab. Senecionetum è dominato da specie ad ampia distri­ 2), di cui sono specie caratteristiche e differenziali buzione (cosmopolite, subcosmopolite, avventizie) Plumbago europaea e Galium album. che assommano al 52,6%, seguite dalle eurasiatiche L'associazione si inquadra nell'alleanza Inulo s.l. (22,7%) e dalle boreali (14,6%). Le mediterra­ viscosae-Agropyrion repentis, istituita da Biondi e nee s.l. sono presenti con il 6,5% e le atlantiche s.l. Allegrezza (1996) come variante submediterraneo­ con il 3,6%. appenninica del Convolvulo-Agropyrion repentis a Nello spettro corologico del Galio-Plum­ carattere temperato. L'alleanza afferisce all'ordine baginetum la percentuale più alta è raggiunta dalle Agropyretalia repentis (classe Artemisietea vulgaris). specie mediterranee s.l. (83,3% nella subass. elytri­ La minore igrofilìa rispetto all' Artemisio­ getosum athericae e 54,5% nella subass. elytrigeto­ Senecionetum, sottolineata da un minor numero e sum repentis). Seguono, a distanza, le eurasiatiche frequenza di specie della classe Molinio­ s.l. (11,9 e 16,2%), le boreali (2,0 e 19,1%) e quelle Arrhenatheretea, deriva dalla maggiore distanza ad ampia distribuzione (2,1 e 8,3%). Trascurabili rispetto alla strada e dalla acclività delle stazioni che sono i valori delle atlantiche s.l. (0,7 e 0,9%) e delle assicura un più pronunciato drenaggio. endemiche (O e 1,0%). Nutrito è, anche in questo caso, il corteggio delle Dai dati esposti si deduce che l 'Artemisio­ specie della classe Stellarietea mediae e dell'ordine Senecionetum possiede un carattere marcatamente Brometalia rubenti-tectori; la presenza, anche con­ nitrofilo-ruderale e decisamente continentale, sistente, di Bromus diandrus, evidenzia il contatto denunciato dalle elevate presenze delle entità ad con cenosi afferenti all'ordine citato e, in particola­ ampia distribuzione ed eurasiatiche s.l. Tale ecolo­ re, con l'associazione Avena barbatae-Brometum gia deriva dalla posizione di questa vegetazione a diandri o simili. diretto contatto con la strada e dall' assenza di accli­ In Tab. 2 sono stati evidenziati due aspetti, che vità delle stazioni. vengono interpretati come subassociazioni: Nel Galio-Plumbaginetum è invece evidente una subass. elytrigetosum athericae (rill. 1-5), di cui marcata impronta mediterranea, che diventa molto sono specie differenziali Elytrigia atherica, decisa nella subassociazione elytrigetosum atheri­ Raphanus raphanistrum, Beta vulgaris, Sinapis cae. L'associazione mostra anche un non elevato alba e Sonchus tenerrimus (rilievo tipo n. 2 di Tab. livello di ruderalità e nitrofilla, evidenziato dai bassi

38 Studia Geobot. 20 (2001) - Associazioni di margine stradale in Abruzzo

80 ,------, nigrum (+), Mercurialis annua(+), Conyza canadensis (+). 70 a) Tabella 2 - Rii. 1: Pescara, 14.10.1995; Aster squamatus (+), Sorghum halepense (+). Rii. 2: Alanno (PE), 1.10.1995; Picris 60 echioides (+), Consolida regalis (+), Brachypodium sylvaticum 50 (+), Ulmus minor(+). Rii!. 3, 4: Alanno (PE), 1.10.1995. Rii. 5: 40 Alanno (PE), 1.10.1995; Oxalis articulata (+). Rii. 6: Ofena (AQ), 30 2.10.1995; Ailanthus altissima(+), Conyza canadensis (+). Rii. 7: Tra Gioia dei Marsi e Venere (AQ), 8.10.1985. Rii. 8: Pressi di 20 Monticchio (AQ), 19.9.1995; AJThenatherum elatius ( +). Rii!. 9, 10 10: Tra Paganica e Barisciano (AQ), 19.9.1995. Rii. Il: Tra o Navelli e Capestrano (AQ), 19.9.1995. Rii. 12: Trivento (CB), Echium italicum Eryngium amethystinum emicriptofite geofite camefite terofite fanerofite 24.9.1995; (+), (+). Rii. 13: Trivento (CB), 24.9.1995; Erysimum cheiri (+). Rii. 14: Bussi 90 ,------, (PE), 18.9.1995; Equisetum arvense(I). Rii. 15: Capestrano(AQ), 8 18.9.I 995; Odontites lutea (1). Rii. 16: Tra Poggio Picenze e 0 - b) 70 Barisciano (AQ), 11.9.1995; Rii. 17: Tra Poggio Picenze e Barisciano (QA), 11.9.1995; Orlaya grandiflora(+). Rii. 18: S. Pio 60 delle Camere (AQ), 11.9.1995. Rii. 19: Navelli (AQ), 11.9.1995; 50 Spartium junceum (+). Rii. 20: Capestrano (AQ), 11.9.1995; 40 Sanguisorba minor ( +), Scolymus hispanicus (r). 30 Ringraziamenti

La ricerca è stata eseguita con il contributo M.U.R.S.T. ex 60%. medit. eurasiat. boreali ampia atlant. endem. distrib. Bibliografia Fig. 2 - Spettri biologici (a) e corologici (b) dell'Artemisio­ Senecionetum (•), Galio-Plumbaginetum elytrigetosum athericae Anzalone B., 1976. Il "Senecio inaequidens" DC. in Italia. Giom. ( O) e Galio-Plumbaginetum elytrigetosum repentis (__,). Bot. Ital., I IO: 437-438. Life form (a) and chorological (b) spectra of Artemisio­ Biondi E. & Allegrezza M. 1986. Inquadramento fitosociologico di Senecionetum (•). Galio-Plumbaginetum elytrigetosum athericae alcune fonnazioni prative del territoiio coJlinare anconetano. (O) and Galio-Plumbaginetum elytrigetosum repentis ( ). Giorn. Bot. Ital., 130: 136-148. Biondi E. & Baldoni M., 1991. La vegetazione di margine stradale deJJ'ordineBrometalia rubenti-tectori nell'Italia centrale. Ann. valori delle entità ad ampia distribuzione. Tutto ciò Bot. (Roma), Studi sul territorio, voi. 49, suppi. 8: 213-218. dipende da vari fattori: maggiore distanza dal ciglio Braun-Blanquet, 1964. Pflanzensoziologie, Wien. Conti F., 1998. An annoted checklist of che flora of the Abruzzo. stradale, contatto con la vegetazione dei campi Bocconea, 10: 5-275. abbandonati, acclività delle stazioni. Pascetti S. & Veri L., 1983. Alcuni aspetti deJla vegetazione sinan­ tropica della città deJJ'Aquila e dei dintorni (Abruzzo-Italia centrale). Coli. Phytosoc., 12: 429-447. Appendice Guarrera P. & Tammaro F., 1994. La Floradel M. Sirente e zone limi­ trofe (Appennino Abruzzese). Ann. Bot.(Roma), 52: 267-381. Località, data e specie sporadiche dei rilievi: Hruska K., 1985. Contributo alla conoscenza della vegetazione Tabella 1 - Rii. I: Tra Barisciano e Prata d' Ansidonia (AQ), ruderale deJle Marche. Doc. Phytosoc., 9: 359-365. 19.9.1995. Rii. 2: S. Atto di Teramo, 14.11.1997; Sonchus asper Hruska K., 1989. La vegetazione spontanea della città di Urbino. (+), Conyza albida (+). Rii. 3: Pressi di Monticchio (AQ), Arch. Bot. Biogeogr. ltal., 65: 207-221. 19.9.1995; Amaranthus retroflexus (+), Linaria purpurea (+), Hruska K., 1998. Carta della vegetazione della città di CameriJJo Eryngium campestre(+). Rii. 4: Coppito (AQ), 9.10.1995; Salvia (MC). Dipartimento di Botanica ed Ecologia, Università di verbenaca (+), Scabiosa columbaria (1), Galium album (I), Camerino. Brachypodium rupestre (+). Rii. 5: Pressi di Pettino (AQ), Pignatti S., 1982. Flora d'Italia. Edagricole, Bologna, 3 voli. 28.9.1995; Diplotaxis erucoides (+), Fa1lopù1 convolvulus (+), Pirola A., 1970. Elementi di Fitosociologia, Bologna, 153 pp. Trifolium repens (+). Rii. 6: Pressi di Pettino (AQ), 28.9.1995; Pirone G. & Ferretti C., 1999. Flora e vegetazione spontanee della Lolium perenne(+), Poa annua(+). Rii. 7: Ofena (AQ), 2.10.1995; città di Pescara (Abruzzo, Italia). Fitosociologia, 36: 111-155. EcbaJlium elaterium (+), Xanthium italicum (1), Silybum maria­ Poldini L., 1989. La vegetazione del Carso Isontino e Triestino. Ed. num(+). Rii. 8: Caporciano(AQ), 25.9.1995. Rii. 9: Navelli(AQ), Lint, Trieste, 3 J 5 pp. 25.9.1995; Phleum ambiguum (!). Rii. 10: L'Aquila, 19.9.1995; Tammaro F. & Pirone G., 1980. Segnalazioni Floristiche Italiane: Tordilium maximum (+), Lepidium graminifolium (r), Solanum 62-65. Inform. Bot. Ital., 12: 75-77.

Received October 2, 2000 Accepted March 26, 2001

39

Studia Geobotanica. Voi. 20: 41-59 (2001)

CONTRIBUTIONS TO THE LICHEN FLORA OF CRETE: A COMPILATION OF REFERENCES AND SOME NEW RECORDS

Martin GRUBE, Louise LINDBLOM 2 and Helmut MAYRHOFER '

' lnstitut fiir Botanik, Karl-Franzens-Universitat Graz, 1-lolteigasse 6, A-80 I O Graz, Austria. ' Dept. of Botany, Univ. of Bergen, Allégaten 41, N-5007 Bergen, Norway.

Keywords: biodiversity, checklist, Crete, flora, lichenicolous fungi, lichenized fungi, lichens, Mediterranean.

Abstract: An updated checklist of lichens of Crete is presented. A tota! of 440 taxa is reported. The checklist distinguishes 4 administrative subdivisions (prefectures) in Crete. Strong degradation of the vegetation through­ out the island limits the abundance of corticolous lichens, only the woody patches in the western part contain a number of otherwise rare oceanic species.

Introduction

The first lichen collections from Crete were systems, e.g. the Dikti mountains towards the east, reported by Zahlbruckner (1906). Some years later, and on the Sitia periinsula, the Omo ridges and the Steiner (1916, 1919) published on collections made Afendis-Kavousi mountains (1476 m). Amidst the by Ginzberger and those gathered during an excur­ larger mountain systems, three highland plains sion ofthe University ofVienna. A larger number of (polje) are found: the Omalos, Lassithi, and Nida specimens were collected by Rechinger during a plains. The mountains ridges are steep at their longer stay on the island. These collections were southem ridges towards the Lybian sea, and inter­ determined and published by Szatala (1943a). These rupted by deep gorges, with Samaria as the most prior results were the basis for the first checklist by famous one. Severa! small islands surround Crete: Kleinig (1966) who listed some 330 species, inclu­ Gramvoussa, Pontikinisi, Dia, Dyonisiades in the ding 80 new records. Additional information was Nmth, and Gavdos (i.e. the southemmost point of contributed by Rondon (1969), and is scattered Europe), Paximadia, Chrisi, and Koufonisi in the throughout the literature. In this compilation we South. have tried to screen the literature completely, and Crete is the remaining part ofa larger geologica! also include data from various other collectors as folding system originating from the Oligocene, well as some of our own collections. which connected the Peloponnes with the mountains ofWest Anatolia. This is part ofthe European plate, Survey area which is raised while the African plate moved underneath to the north beginning with the Crete has a rather centrai location in the Cretaceous. A geologica! map was provided by MeditetTanean, and is, together with its surrounding Creutzberg et al. (1977). Phyllite-quartzites, with a small islets, the southernmost island complex of richly developed lichen diversity, are the oldest Europe. lt is also the fifth-largest island in the metamorphic rocks on the island (at least 300 m Mediterranean with an area of 8729 square kilome­ years old) and mainly found in the western parts. A ters (after Sicily, Sardinia, Cyprus, and Corsica). completely different lichen flora is found on platey The main island has a significant East-West orienta­ limestones and crystalline limestones, which are tion and measures 254 km in this direction, whereas otherwise dominating in the mountains and contri­ its width is only 12 kn1 at its narrowest and 56 km buting to karst phenomena. The late tertiary to neo­ at its broadest point. The natural subdivision of the genie mari, sandstones and clays, are mainly found island is determined by its mountain systems, the in the lowlands. Natural concretes, such as breccia Lefka Ori (2453 m), the Psiloritis (2456 m), which and conglomerate, can be of any date, even less than have snow cover mostly unti! June, and the lower 50 years old. The unstable structure of these sub-

41 M. GRUBE, L. LINDBLOM, H. MAYRHOFER

'

Fig. I - The island ofCrete and its subdivision in prefectures: C = Nomos Chanion, R = Nomos Rethimnion, l = Nomos lrakliou, L = Nomos Lassithi. strata does not support a well developed lichen influenced by grazing animals, and there is hardly flora. any undisturbed locality. Open vegetation is

Crete is characterized by a typical Medi­ mostly garigue (phrygana) or steppe, and well terranean climate and influenced by different developed maquis is found only in the valleys of winds. In winter, the accompanying rains of the the western parts. Rondon (1969) provided some mainly westerly winds are the reason why Western notes on common lichen habitats in Crete. To these Crete gets most of the precipitation, whereas the we can add the observations that thomy cushion eastern and southern parts are more dry. A warm plants are an interesting microhabitat for lichens at wind from the South, called Natia, which tran­ higher altitudes, and that Castanea plantations in sports dust from Sahara, sometimes interferes in Western Crete are lichenologically interesting, winter. In summer, relatively strong and cool especially in shaded gorges. Although the under­ winds from the North, the etesian winds, may storey and even branches of the trees in these plan­ occur. Frost is rare in the lowlands, whereas the tations are heavily degraded by goat grazing, nota­ mountains get snow down to 1600 m in winter, ble oceanic species, such as Degelia plumbea or which is then persisting often until May. Rainfalls various Bacidia species, are found here. start in September to N ovember, and reach their maximum in December and January. In June to Organization of the list August, even the mountains rarely get any rain. Fog is encountered mainly in northem exposed Four administrative subdivisions (prefectures, parts of the mountains. The often heavy rains, gr. Nomos) are distinguished (Fig. 1): C = Nomos which are not absorbed by the soil, are of little use Chanion, R = Nomos Rethimnion, I = Nomos for the vegetation in general. However, in the val­ Irakliou, L = Nomos Lassithi. RII denotes the areas ley systems in the West, they contribute to higher in the Psiloritis mountains where the corresponding humidity. prefecture can not be assigned. Details of new The vegetation is mostly eumediterranean and, records for a prefecture are given in brackets after today, large closed forests are missing, contrasting the abbreviation of the corresponding prefecture. As with the descriptions of Homer and Theophrastus, a literature reference we included the first publica­ who characterized Crete as an island with 'rich tion of a specimen. Records of lichenicolous fungi woodland'. However, it is difficult to estimate include the name of their hosts. what exactly was understood under that term and, unfortunately, pollen deposits are rare. Rather The list dense chestnut-platane stands, often with oak are still found in the western, more humid part of the Acarospora cervina A. Massal. island. Large parts of the island, including the Szatala 1943b; Kleinig 1966 - R (7 km S understorey of the woodlands are, however, Anogia, 1993, leg. Nordin, UPS), I, L

42 Studia Geobot. 20 (200 I) - A check-Iist of Crete Iichens -

Acarospora glaucocarpa (Ach.) Korb. Aspicilia caesiocinerea (Nyl. ex Malbr.) Amold Szatala 1943b -R Syn. Aspicilia subdepressa (Nyl.) Arnold Steiner 1916; Kleinig 1966, - C, I Acarospora nodulosa (Dufour) Hue Szatala 1943b -C Aspicilia calcarea (L.) Mudd Zahlbruckner 1906 (var. concreta Schaer.); Acarospora schleicheri (Ach.) A. Massa!. Szatala 1943b (f. atomaria (A. Massa!.) Hazsl.); Rondon 1969 - C (lmbros, 1988, leg. Timdal, Kleinig 1966; Rondon 1969; Eigler 1969; Papp hb. Timdal) et al. 1999 -C, I, L Acarospora smaragdula (Wahlenb.) A. Massa!. Aspicilia cheresina (Muli. Arg.) Hue var. microspo­ Zahlbruckner 1906 - I ra (Amold) Clauzade & Cl. Roux Acarospora sulphurata (Arnold)Amold Syn. Aspicilia microspora (Zahlbr.) Hue Kleinig 1966 - R Szatala 1943b; Rondon 1969 -I, L Acarospora umbilicata Bagl. Aspicilia cinerea (L.) Korb. Zahlbruckner 1906; Kleinig 1966; Rondon 1969 Syn. Acarospora eretica J. Steiner, A. percaenoi­ -C, I, L des (Nyl.) Jatta Aspicilia contorta (Hoffm.) Kremp. ssp. contorta Zahlbruckner 1906; Steiner 1916 -I Syn. Aspicilia viridescens (A. Massa!.) Hue Acarospora veronensis A. Massa!. Zahlbruckner 1906; Szatala 1943b - C, RJI, I, L Szatala 1943b -C, RJI Aspicilia contorta ssp. hojfmanniana S. Ekman & Acrocordia gemmata (Ach.) A. Massa!. Froberg R (Adelaki, 12 km S Rethimno, 1993, leg. Syn. Aspicilia hoffmannii auct. Nordin, UPS) Rondon 1969 - I (2 km W Festos, 1993, leg. Nordin, UPS) Amandinea punctata (Hoffm.) Coppins & Scheid. Aspicilia coronata (A. Massa!.) Anzi Syn. Buellia myriocarpa DC. (Mudd) var. virens Szatala 1943b -C, I J. Steiner; Buellia aequata (Ach.) Szatala Aspicilia cupreoglauca de Lesd. Steiner 1916 -I Rondon 1969 Anaptychia ciliaris (L.) Korb. Aspicilia esculenta (Pall.) Flagey Szatala 1943b (var. crinalis (Schleich.) Rabenh., Syn. Aspicilia desertorum (Kremp.) Mer. var. melanosticta (Ach.) Boist.); Kleinig 1966; Szatala 1943b - C Czeczuga et al. 1994 - C, R (Oros Idi, Skinakas, Aspicilia farinosa (Florke) Arnold Zahlbruckner 1906; Szatala 1943b; Rondon 1997, leg. Suppan, GZU), RJI 1969 - I, L Anema prodigulum (Nyl.) Henssen Aspicilia grisea Amold Rondon 1969 Kleinig 1966 - C Arthonia dispersa (Schrad.) Nyl. Aspicilia intermutans (Nyl.) Arnold Szatala 1943b - I (Matala, 1997, leg. Grube, Zahlbruckner 1906; Rondon 1969 -I, L GZU), L Arthonia epiphyscia Nyl. Bacidia bagliettoana (A. Massa!. & De Not.) Jatta L (1,5 km S Kalavros, 1999, leg. Grube, GZU) Syn. Bacidia muscorum (Sw.) Mudd Kleinig 1966 -C, R (7 km SE Garazo, 1993, leg. Host: Physcia scopulorum Nordin, UPS) Arthonia galactites (DC.) Dufour Bacidia delicata (Larbal. ex Leight.) Coppins Zahlbruckner 1906 - I R (7 km S Anogia, 1993, leg. Nordin, UPS) Arthonia meridionalis Zahlbr. Bacidia rosella (Pers.) De Not. Szatala 1943b -L R (6 km N Apostoli, 1993, leg. Nordin, UPS; Arthonia palmicola Ach. Adelaki, 12 km S Rethimno, 1993, leg. Nordin, Szatala 1943b - L UPS) Bacidia subincompta (Nyl.) Arnold Arthonia pinastri Anzi Syn. Bacidia affinis(Stizenb.) Vain. Sundin 1999 - L Szatala 1943b - C Arthothelium ruanum (A. Massa!.) Korb. Bacidina assulata (Korb.) S. Ekman Syn. A. dispersum auct. Syn. Bacidia intermedia (Hepp) Vain. Kleinig 1966 -S Szatala 1943b -C

43 M. GRUBE, L. LINDBLOM, H. MAYRHOFER

Bagliettoa limborioides A. Massa!. Caloplaca chrysodeta (Vain. ex Rasanen) Dombr. Syn. Eagliettoa sphinctrina auct. C (Agia Irini, 1998, leg. Trinkaus & Grube, Kleinig 1966 -L GZU) Bagliettoa parmigera (J. Steiner) Vezda & Poelt Caloplaca circumalbata (Delile) Wunder var. Syn. Verrucaria calciseda DC.; Verrucaria par­ candida (Stizenb.) Wunder migera J. Steiner Wunder 1974 -C, L Zahlbruckner 1906; Steiner 1916 (var. calcivora Caloplaca citrina (Hoffm.) Th. Fr. A. Massa!.); Rondon 1969 -I, L Syn. Caloplaca phlogina (Ach.) Flag Buellia epigaea (Pers.) Tuck. Steiner 1916 -I Trinkaus & Mayrhofer 2000 - L Caloplaca crenularia (With.) J.R. Laundon Buellia stellulata (Taylor) Mudd Szatala 1943a, Kleinig 1966; Rondon 1969; Syn. Buellia maritima (A. Massa!.) Bagl.; Papp et al. 1999 - C, L Caloplaca cretensis (Zahlbr.) Wunder Bue/lia suba/buia (Nyl.) Muli. Arg. Syn. E/astenia cretensis Zahlbr. Steiner 1916; Szatala 1943b - C, I, L Zahlbruckner 1906; Wunder 1974 - I Buellia subdiscijormis (Leight.) Vain. Caloplaca erythrocarpa (Pers.) Zwackh Szatala 1943b; Rondon 1969 -L Syn. Caloplaca !al/avei (Clemente) Flagey Buellia vilis Th. Fr. Zahlbruckner 1906 (as C. arenaria var. lallavei); Kleinig 1966 -L Kleinig 1966; Rondon 1969 -C (lmbros, 1997, Buellia zoharyi Galun (nom. cons. prop.) leg. Mayrhofer & Erti, GZU; Agios Nektarios, Trinkaus & Mayrhofer 2000 - C 1993, leg. Nordin, UPS), L Caloplaca ferruginea (Huds.) Th. Fr. Caloplaca alnetorum Giralt, Nimis & Poelt Zahlbruckner 1906; Kleinig 1966 - L R (Skinakas, 1997, leg. Suppan, GZU) Caloplacajlavescens (Huds.) J.R. Laundon Caloplaca alociza (A. Massa!.) Mig. C ( 1 km S lmbros, leg. Mayrhofer & Erti, GZU) Syn. Caloplaca agardhiana (A. Massa!.) Caloplacajlavorubescens (Huds.) J.R. Laundon Clauzade & Cl. Roux Syn. Caloplaca salicina (Schrad.) Szatala; Zahlbruckner 1906 (f. minuta, f. albomargina­ Caloplaca aurantiaca s. lat. ta); Szatala 1943b (as E/astenia, severa( forms); Zahlbruckner 1906 (Caloplaca aurantiaca var. Rondon 1969; Wunder 1974- C, I, L diffracta); Kleinig 1966; Rondon 1969 - R Caloplaca aractina (Fr.) Hayren (Skinakas, 1997, leg. Suppan, GZU; 6 km NW Caloplaca viridirufa auct. Apostoli, 1993, leg. Nordin, UPS); L Kleinig 1966 - L Caloplaca jlavovirescens (Wulfen) Dalla Torre & Caloplaca arenaria (Pers.) Mi.ili.Arg. Samth. Syn. Caloplaca lamprocheila (DC.) Flagey, Kleinig 1966 -I Caloplaca craspedia (Ach.) Szatala Caloplaca ji,scoatroides J. Steiner Steiner 1916; Szatala 1943a; Rondon 1969 - I Kleinig 1966 - C, L (Karidi, 1976, leg. Mayrhofer, GZU) Cal laca aurantia (Pers.) Hellb. op Caloplaca grimmiae (Nyl.) H. Olivier Syn. Caloplaca callopisma (Ach.) Th. Fr. I (SE Agi a Pelagia, 1997, leg. Grube, GZU) Zahlbruckner 1906; Steiner 1916 (f. orientalis); Host: Candelariella vitellina Kleinig 1966; Rondon 1969-C (1 km S Imbros, Caloplaca haematites (St.-Amans) Zwackh leg. Mayrhofer & Erti, GZU), I, L Kleinig 1966; Rondon 1969 - C, RII, L Caloplaca biatorina (A. Massa!.) J. Steiner Caloplaca herbidella (Hue) H. Magn. Kleinig 1966 - I C (Omalos, 1998, leg. Trinkaus & Grube, GZU) Caloplaca carphinea (Fr.) latta Caloplaca inconnexa (Nyl.) Zahlbr. Papp et al. 1999 -C Rondon 1969 Caloplaca cerina (Ehrh. ex Hedw.) Th. Fr. Caloplaca lactea (A. Massa!.) Zahlbr. Zahlbruckner 1906; Kleinig 1966; Rondon 1969 Zahlbruckner 1906; Kleinig 1966; Rondon - C, R (Skinakas, 1997, leg. Suppan, GZU; 7 kn1 1969; Triebel 1989 - L, I S Anogia, 1993, leg. Nordin, UPS), RII Caloplaca lithophila H. Magn. Caloplaca chalybaea (Fr.) Muli. Arg. Kleinig 1966; Rondon 1969 - C, R Zahlbruckner 1906; Rondon 1969; Wunder 1974 Caloplaca necator Poelt & Clauzade -1, L Rondon 1969

44 Studia Geobot. 19 (2000) - A check-list of Crete lichens -

Caloplaca ochracea (Schaer.) Flagey Candelariella medians (Nyl.) A.L. Sm. Syn. Xanthocarpia ochracea (Schaer.) A. I (2km W Festos, 1993, leg. Nordin, UPS), L Massa!. (5km S Vai, 1976, leg. Mayrhofer, GZU) Kleinig 1966; Hafellner & Poelt 1979 - L Candelariella vitellina (Hoffm.) Muli. Arg. Caloplaca pyracea (Ach.) Th. Fr. Zahlbruckner 1906; Kleinig 1966; Rondon Syn. Caloplaca luteoalba auct. p.p. 1969; Czeczuga et al. 1994; Papp et al. 1999 - Kleinig 1966; Zahlbruckner 1906 (+ f. pyrithro­ C, R (1km S Kouroutes, 1976, leg. Mayrhofer, ma (Ach.) Zahlbr.); Rondon 1969; Czeczuga et GZU), I al. 1994 - C, I, L Candelariella xanthostigma (Ach.) Lettau Caloplaca rechingeri Servit Szatala 1943b - C Kleinig 1966 - I Catillaria chalybeia (Borr.) A. Massa!. var. Caloplaca rhinodinoides J. Steiner chalybeia Kleinig 1966 - L Zahlbruckner 1906; Szatala 1943b; Kilias 1981; Caloplaca rubelliana (Ach.) Lojka Bellemère 1994 - C, I, L Kleinig 1966 - L Catillaria chalybeia var. choropoliza (Nyl.) H. Caloplaca saxicola (Hoffm.) Nordin Kilias Syn. Caloplaca murorum(Ach.) Th. Fr. Kilias 1981 - R Zahlbruckner 1906; Kleinig 1966 - L Catillaria detractula (Nyl.) H. Olivier Caloplaca scotoplaca (Nyl.) H. Magn. Syn. Lecania detractula (Nyl.) Arnold Syn. Caloplaca caesiorufa auct. Szatala 1943b, Triebel 1989 - I, L Steiner 1916 - I Catillaria lenticularis (Ach.) Th. Fr. Caloplaca tenuata (Nyl.) Zahlbr. Szatala 1943b; Rondon 1969 - C, I, L Rondon 1969 Catillaria mediterranea Hafellner Caloplaca variabilis (Pers.) Muli. Arg. C (Omalos, 1998, leg. Trinkaus & Grube, GZU) Syn. Caloplaca paepalostoma (Anzi) Jatta Host: Anaptychia ciliaris Steiner 1916 (f. acrustacea Arnold); Rondon Catillaria nigroclavata (Nyl.) Schuler 1969 (+ f. acrustacea, + var. ocellulata (Ach.) R (3 km ESE Vrises, 1993, leg. Nordin, UPS; 6 Boist.); Wunder 1974 - C, R (6 km Se Nea Kria km NW Apostoli, 1993, leg. Nordin, UPS) Vrisi, 1997, leg. Mayrhofer & Erti, GZU; 14 km Cercidospora lobothalliae Nav.-Ros. & Calatayud W Spili, 1993, leg. Nordin, UPS), I, L (Psichron, ined. 1976, leg. Mayrhofer, GZU) C (Omalos, 500m SW Lakki, 1976, leg. Caloplaca velana (A. Massa!.) Du Rietz var. velana Mayrhofer, GZU) Syn. Caloplaca placidia (A. Massa!.) J. Steiner, Host: Lobothallia radiosa Caloplaca dolomitico/a (Hue) Zahlbr. Cladonia cariosa (Ach.) Spreng. Rondon 1969 - C (Imbros gorge, 1997, leg. Szatala 1943b - C Grube, GZU), R (6 km NW Apostoli, 1993, leg. Cladonia cervicornis (Ach.) Flot. subsp. Nordin, UPS), I (1,5 km N Kounari, 1993, leg. verticillata (Hoffm.) Ahti Nordin, UPS; 2 kn1 W Festos, 1993, leg. Nordin, Papp et al. 1999 - C UPS) Cladonia convoluta (Lam.) Anders Caloplaca velana var. schaereri (Arnold) Clauzade Syn. Cladoniafoliacea (Huds.) Willd. var. con­ & Cl. Roux voluta (Lam.) Vain.; Cladonia endiviaefolia Syn. Caloplaca schaereri (Florke) Zahlbr. (Dicks.) Fr. Rondon 1969 Zahlbruckner 1906; Steiner 1916; Szatala Caloplaca vitellinula auct. 1943b; Rasanen 1944; Rondon 1969; Papp et al. Kleinig 1966 - R 1999- C, I Caloplaca xantholyta (Nyl.) Jatta Cladoniafirma (Nyl.) Nyl. C (1 km S lmbros, 1997, leg. Mayrhofer & Erti, Syn. Cladonia nylanderi Coutinho GZU) Kleinig 1966 - L Candelariella aure/la (Hoffm.) Zahlbr. Cladoniafoliacea (Huds.) Willd. Syn. Caloplaca subsimilis Th. Fr., Candelariella Kleinig 1966 - C, R (6 km NW Apostoli, 1993, subsimilis (Th. Fr.) J. Steiner leg. Nordin, UPS), I, L Steiner 1916; Szatala 1943b; Kleinig 1966 - C, Cladonia furcata (Huds.) Schrad. ssp . .fi,rcata I, L Kleinig 1966 - L

45 M. GRUBE, L. LINDBLOM, H. MAYRHOFER

Cladonia furcata ssp. subrang?formis (Sandst.) Collema nigrescens (Huds.) DC. Abbayes R (Arkadi Monastery, on Quercus macrolepis, Kleinig 1966 -L 1976, leg. H.J.M. Bowen, E) Cladonia pocillum (Ach.) Grognot Collema occultatum Bagl. Steiner 1916; Szatala 1943b; Rondon 1969-1, R R (6 km NW Apostoli, 1993, leg. Nordin, UPS) (6 km NW Apostoli, 1993, leg. Nordin, UPS), L Collema polycarpon Hoffm. Cladonia pyxidata (L.) Hoffm. Syn. Collema stygium Schaer. Szatala 1943b; Kleinig 1966; Rondon 1969; Szatala 1943b; Degelius 1954; Kleinig 1966- C, Papp et al. 1999 - L I, L Cladonia rangiformis Hoffm. Collema ryssoleum (Tuck.) A. Schneider Szatala 1943b - L Kleinig 1966 - C, R (1km S Kouroutes, 1976, Clauzadea immersa (Weber) Hafellner & Bellem. leg. Mayrhofer, GZU), I Syn. Protoblastenia immersa (Weber) J. Steiner, Collema subnigrescens Degel. Biatora immersa (Weber) Hepp Kleinig 1966 - C; R (Adelaki, 12 km S Szatala 1943b; Kleinig 1966 - C, L Rethimno, 1993, leg. Nordin, UPS) Clauzadea metzleri (Kéirb.) Clauzade & Cl. Roux Collema tenax (Sw.) Ach. em. Degel. ex D. Hawksw. Syn. Collema pulposum (Bernh.) Ach. I (2km W Festos, 1993, leg. Nordin, UPS) Szatala 1943b - C, R (6 km NW Apostoli, 1993, Clauzadea monticola (Schaer.) Hafellner & leg. Nordin, UPS) Bellem. Collema undulatum Laurer ex Flot. Syn. Protoblastenia monticola (Schaer.) J. Kleinig 1966 - I Steiner, Lecidea fi1scorubens (Nyl.) Nyl. Zahlbruckner 1906; Steiner 1916 - I, L (5km S Degelia plumbea (Lightf.) P.M. J0rg. & P. James Vai, leg. Mayrhofer, 1976, GZU). Czeczuga et al. 1994 - C (1km E Vlatos, 1998, Collema callopismum A. Massa!. leg. Trinkaus & Grube, GZU), I Syn. Leptogium callopismum (A. Massa!.) Dermatocarpon luridum (With.) J.R. Laundon Harm. Syn. Dermatocarponfluviatile (Weber) Th. Fr. Szatala 1943b -C Kleinig 1966 - C Collema crispum (Huds.) Weber ex F.H. Wigg. Dermatocarpon miniatum (L.) W. Mann Syn. Collema cheilum Ach.; Collema crispum Zahlbruckner 1906 - C, I var. metzleri (Arnold) Degel.; Collema subchei­ Dimelaena oreina (Ach.) Nonnan lum Harm. Obermayer 1997 -R Steiner 1916 (C. metzleri); Szatala 1943b; Diploicia canescens (Dicks.) A. Massa!. Degelius 1954 -C, I Szatala 1943b - C Collema cristatum (L.) Weber ex F.H. Wigg. var. Diploschistes actinostomus (Ach.) Zahlbr. cristatum Zahlbruckner 1906; Rondon 1969 (var. farino­ Syn. Collema granuliferum Nyl. sus (Anzi) Zahlbr.) - I Szatala 1943b; Kleinig 1966; Papp et al. 1999 - Diploschistes diacapsis (Ach.) Lumbsch C, I, L Syn. Diploschistes a/bescens Lettau, Diplo­ Collema cristatum var. marginale (Huds.) Degel. schistes albissimus (Ach.) Dalla Torre & Sarnth. Syn. Collema multifìdum (Scop.) Rabenh. Steiner 1916; Szatala 1943b; Kleinig 1966; Zahlbruckner 1906 (var. marginale); Szatala Rondon 1969; Lumbsch 1989 - C, I, L 1943b (var.jacobaefolium) - L Diploschistes gypsaceus (Ach.) Zahlbr. Collema jlaccidum (Ach.) Ach. Zahlbruckner 1906 - L R (7 km S Anogia, 1993, leg. Nordin, UPS) The old record ofthis species could also refer to Collema ji,rji,raceum (Arnold) Du Rietz D. diacapsis. Sazatala 1943b- C, R (7 km S Anogia, 1993, leg. Diploschistes muscorum (Scop.) R. Sant. Nordin, UPS; Arkadi Monastery, on Quercus Papp et al. 1999 -C, L macrolepis, 1976, leg. H.J.M. Bowen, E) Diploschistes ocellatus (Vili.) Norman Collema multipunctatum Degel. Szatala 1943b; Rondon 1969; Lumbsch 1989 - R (2 km NW Akoumia, 1993, leg. Nordin, UPS; C (1 km S Imbros, leg. Mayrhofer & Erti, GZU, Adelaki, 12 km S Rethimno, 1993, leg. Nordin, Agios Nektarios, 1993, leg. Nordin, UPS), I UPS) (Rogdia, 1965, leg. Degelius, UPS), L

46 Studia Geobot. 20 (200 I) - A check-list of Crete lichens

Diploschistes scruposus (Schreb.) Norman Farnoldiajurana (Schaer.) Hertel Syn. Diploschistes violarius (Nyl.) Lettau Syn. Lecidea jurana Schaer. Zahlbruckner 1906; Kleinig 1966 - R (1km S Kleinig 1966 -I Kouroutes, 1976, leg. Mayrhofer, GZU), I, L Fulgensia bracteata (Hoffm.) Rasanen Diplotomma alboatrwn (Hoffm.) Flot. Zahlbruckner 1906 -L Syn. Buellia alboatra (Hoffm.) Th. Fr.; Buellia Fulgensiajitlgens (S w.) Elenkin subochracea (Zahlbr.) Steiner, Diplotomma Zahlbruckner 1906 - I subochracewn (Zahlbr.) Szatala Fulgensia jitlgida (Nyl.) Szatala Zahlbruckner 1906 (as B. alboatra var. suboch­ Syn. Caloplacafiilgida (Nyl.) J. Steiner racea); Szatala 1943b -C, I, L Kleinig 1966; Westberg & Karnefelt1998 -C, L Diplotomma chlorophaeum (Hepp ex Leight.) Fulgensia schistidii (Anzi) Poelt Szatala Syn. Caloplaca schistidii (Anzi) Zahlbr. Syn. Buellia chlorophaea (Mtill. Arg.) Lettau Kleinig 1966 - C, R Rondon 1969 Fulgensia subbracteata (Nyl.) Poelt Diplotomma epipolium (Ach.) Arno Id Rondon 1969; Poelt 1965; Papp et al. 1999 - C, Syn. Buellia epipolia (Ach.) Mong. I, L Zahlbruckner 1906; Szatala 1943b; Rondon Fuscopannaria ignobilis (Anzi) P.M. J0rg. 1969 - C, R (Somidos, 1997, leg. Grube, GZU), Syn. Pannaria ignobilis Anzi L Vezda 1995 - I Diplotomma nivale (Bagl. & Carestia) Hafellner Fuscopannaria leucophaea (Vahl) P.M. J0rg. Syn. Diplotomma margaritaceum (Somme,f.) Sza­ Syn. Pannaria leucophaea (Vahl) P.M. J0rg. tala, Buellia nivalis (Bag. & Car.) Hertel ex Haf. C (Omalos, 1998, leg. Trinkaus & Grube, GZU) Szatala 1943b -I, L Fuscopannaria leucosticta (Tuck.) P.M. J0rg. Diplotomma pharcidium (Ach.) M. Choisy Syn. Pannaria leucosticta (Tuck.) Nyl. Syn. Buellia pharcidia (Ach.) Malme Zahlbruck:ner 1906; Szatala 1943b - C; R R (7 km S Anogia, 1993, leg. A. Nordin, UPS) Fuscopannaria mediterranea (Tav.) P.M. J0rg. Diplotomma populorum A. Massa!. Syn. Pannaria mediterranea Tav. Syn. Buellia alboatra var. populorum (A. J0rgensen 1978 (map) Massal.) H. Olivier Fuscopannaria olivacea (P.M. J0rg.) P.M. J0rg. Rondon 1969 Syn. Pannaria olivacea P.M. J0rg. Diplotomma venustum Korb. J0rgensen 1978 (map); Vezda 1985 -C, R (7 km Syn. Buellia venusta (Korb.) Lettau S Anogia, 1993, leg. Nordin, UPS; Adelak:i, 12 Nordin 2000 (map of K- chemotype). km S Rethimno, 1993, leg. Nordin, UPS) Dirina cretacea (Zahlbr.) Tehler Syn. Chiodecton cretaceum Zahlbr. Gonohymenia nummularia (Nyl.) Henssen Zahlbruckner 1906; Szatala 1943b; Rondon C (Agios Nek:tarios, 1993, leg. Nordin, UPS) 1969; Tehler 1983; Roux 1991 (map) - R Gyalecta derivata (Nyl.) H. Olivier (Damioni, 13 km SW Spili, 1993, leg. Nordin, R (Arkadi Monastery, on Quercus macrolepis, UPS), I, L 1976, leg. H.J.M. Bowen, E) Dirina massiliensis Durieu et Mont. f. massiliensis Gyalecta truncigena (Ach.) Hepp Syn. Dirina repanda Fr. C (1 km E Vlatos, 1998, leg. Trinkaus & Grube, Szatala 1943b; Kleinig 1966; Rondon 1969; GZU), R (Adelaki, 12 kn1 S Rethimno, 1993, Tehler 1983; Roux 1991 (map). -C, R (6 km W leg. Nordin, UPS) Spili, 1993, Nordin, UPS), I, L Dirina massiliensis f. sorediata (Mi.ili. Arg.) Tehler Haematomma nemetzii J. Steiner Tehler 1983 - C (Agios Nektarios, 1993, leg. C - (Temenia, leg. Mayrhofer, 1999, GZU) Nordin, UPS) Heppia lobatoplicata (de Lesd.) Timdal ined. C (Imbros, leg. Timdal, 1988, O), R (Preveli, Endocarpon pusillum Hedw. leg. Timdal, 1988, O), L (Lasithi, leg. Timdal, R (7 km S Anogia, 1993, leg. Nordin, UPS) 1995, O) Evernia prunastri (L.) Ach. Hymenelia epulotica (Ach.) Lutzoni Zahlbruckner 1906; Szatala 1943b; Kleinig 1966 Syn. Jonaspis epulotica (Ach.) Arno Id -C, R, R/1 Szatala 1943b; Kleinig 1966 - I, L

47 M. GRUBE, L. LINDBLOM, H. MAYRHOFER

Immersaria cupreoatra (Nyl.) Calatayud & Lecanora dispersa (Pers.) Sommerf. Rambold Zahlbruckner 1906 (as L. crenulata f. dispersa); Syn. Aspicilia cupreoatra (Nyl.) Arnold; Steiner 1916; Szatala 1943b; Rondon 1969; Aspicilia olivacea Bagl. & Carestia; Bellemerea Grube & Hafellner1990 - RII, C, I cupreoatra (Nyl.) Clauzade & Cl. Roux Lecanora gangaleoides Nyl. Zahlbruckner 1906 - R/I, I, L Szatala 1943b; Rondon 1969 - I, L (1km W Karidi, 1976, leg. Mayrhofer, GZU) Lecania erysibe (Ach.) Mudd Lecanora garovaglii (Kéirb.) Zahlbr. Steiner 1916; Szatala 1943b; Kleinig 1966 - I, L Kleinig 1966 - I Lecania rabenhorstii (Hepp) Arnold Lecanora graeca J. Steiner I (2 km W Festos, 1993, leg. Nordin, UPS) Kleinig 1966 - I Lecania spadicea (Flotow) Zahlbr. Lecanora hagenii (Ach.) Ach. Szatala 1943b; Rondon 1969; M. Mayrhofer Steiner 1916 (var. saxigena J. Steiner); Szatala 1988; Roux 1991 (map) - C, I, L 1943b; Kleinig 1966; Rondon 1969 - C, I Lecania subcaesia (Nyl.) de Lesd. Lecanora horiza (Ach.) Linds. Szatala 1943b - C, I, L Syn. Lecanora sienae de Lesd. Lecania turicensis (Hepp) Miill. Arg. Syn. Lecania albariella (Nyl.) Muli. Arg. Rondon 1969 - C (1km S Omalos, 1976, leg. Szatala 1943b; M. Mayrhofer, 1988 - C, L Mayrhofer, GZU) Lecanographa grumulosa (Dufour) Egea & Lecanora intumescens (Rebent.) Rabenh. Torrente Szatala 1943b (var. glaucorufà (Mart.) Kéirb.) - Syn. Opegrapha grumulosa Dufour; Lecanactis c d61jleri Zahlbr.; Lecanactis grumulosa (Dufour) Fr. Lecanora muralis (Schreb.) Rabenh. Zahlbruckner 1906; Lettau 1937; Szatala 1943b; Syn. Squamaria muralis (Schreb.) Elenkin; Torrente & Egea 1989 - I, L Squamaria albomarginata (Nyl.) Rasanen; Lecanora agardhiana Ach. Lecanora albomarginata Cromb.; incl. Squa­ Szatala 1943a (var. pachnodes); Szatala 1943b; maria dif.fracta (Ach.) Duby; Squamaria versi­ Kleinig 1966 - I, L color (Pers.) H. Olivier Lecanora albescens (Hoffin.) Branth. & Rostr. Zahlbruckner 1906; Szatala 1943b; Kleinig Szatala 1943b - I, L 1966; Papp et al. 1999 - R, RII, I, L Lecanora allophana Nyl. Lecanora olivascens Nyl. Syn. Lecanora subfi1sca (L.) Ach., p.p. Kleinig 1966 - C Kleinig 1966; Rondon 1969 - I Lecanora polytropa (Ehrh. ex Hoffm.) Rabenh. Lecanora argentata (Ach.) Malme Szatala 1943b - RII Syn. Lecanora su�fi,scata H. Magn. Lecanora pruinosa Chaub. Szatala 1943b; Kleinig 1966; Czeczuga et al. Syn. Lecanora (Placodium) sulphurella (Kéirb.) 1994 - C, R (Adelaki, 12 km S Rethimno, 1993, H. Olivier; Squamaria sulphurella (Kéirb.) H. leg. Nordin, UPS), I Olivier Lecanora bolcana (Pollini) Poelt Zahlbruckner 1906; Szatala 1943b; Rondon Rondon 1969 - R (1km S, Kouroutes, 1976, leg. 1969 - C, R (14 km W Spili, 1993, leg. Nordin, H. Mayrhofer, GZU) UPS), I, L Lecanora campestris (Schaer.) Hue Lecanora pulicaris (Pers.) Ach. Kleinig 1966 - C (0,5 km SW Lakki, 1976, leg. Syn. Lecanora chlarona (Ach.) Nyl.; Lecanora Mayrhofer, GZU), R (1km S Kouroutes, 1976, leg. Mayrhofer, GZU), L pinastri (Schaer.) Choisy Lecanora carpinea (L.) Vain. Szatala 1943b; Kleinig 1966 Szatala 1943b - C Lecanora reuteri Schaer. Lecanora cenisia Ach. Kleinig 1966 - I Szatala 1943b - I Lecanora rugosella Zahlbr. Lecanora chlarotera Nyl. Szatala 1943b - L Kleinig 1966; Rondon 1969 (+ f. rugose/la Lecanora rupicola (L.) Zahlbr. ssp. rupicola (Zahlbr.) Poelt); Czeczuga et al. 1994 - C, R Syn. Lecanora glaucoma (Hoffm.) Ach. (Adelaki, 12 km S Rethimno, 1993, leg. Nordin, Zahlbruckner 1916; Szatala 1943b; Rondon UPS), I 1969 - I

48 Studia Geobot. 20 (200 I) - A check-list of Crete lichens

Lecanora rupicola ssp. sulphurata (Ach.) Leuckert lecidella atrosanguinea (Florke) ined. & Poelt Zahlbruckner 1906 ( as L. enteroleuca var. atro­ Syn. Lecanora sulphurata (Ach.) Nyl. sanguinea) - L Zahlbruckner 1906; Szatala 1943b; Rondon See Santesson 1993 for the uncertain placement 1969; Leuckert & Poelt 1989- RII, I, L of this taxon. Lecanora schistina (Nyl.) Arnold lecidella carpathica Korb. Syn. Lecanora praepostera auct. Syn. Lecidea latypeaAch., Szatala 1943b; Rondon 1969 - L Zahlbruckner 1906; Szatala 1943b; Rondon Lecanora subrugosa Nyl. 1969; Leuckert et al. 1990 - R (7 km S Anogia, Szatala 1943b- C 1993, leg. Nordin, UPS), RII, I Lecanora sulphurea (Hoffm.)Ach. Lecidella elaeochroma (Ach.) M. Choisy Syn. Lecidea parasema auct., Lecidea olivacea Zahlbruckner 1906; Szatala 1943b - I, L (Hoffm) A. Massal., ?Lecidea epipolioides var. Lecanora umbrina (Ach.) A. Massal. cacuminum (1. Steiner) Szatala Szatala 1943b- L Zahlbruckner 1906; Szatala 1943b; Kleinig lecanora varia (Hoffm.) Ach. 1966; Rondon 1969 - C, RII, I, L Kleinig 1966 - C Lecidella euphorea (Flèirke) Hertel lecanora xanthostoma Wedd. ex Cl. Roux (nomen Syn. Lecidea euphorea (Florke) Nyl. illegit.) Szatala 1943b - C C (1 km S Imbros, leg. Mayrhofer & Erti, Lecidella patavina (A. Massal.) Knoph & Leucke1i GZU) Syn. Lecidea endolithea Lynge Lecidea atrobrunnea (Rama ud ex Lam. & DC.) He1iel 1967- I Schaer. lecidella scabra (Taylor) Hertel & Leucke1i Zahlbruckner 1906; Kleinig 1966 - I Syn. Lecidea protrusa Fr. Lecideafuscoatra (L.) Ach. Szatala 1943b - L Syn. Lecidea grisella Flèirke Lecidella stigmatea (Ach.) Hertel & Leuckert Rondon 1969- C (Street to Omalos, 500m SW Syn. Lecidea stigmatea Ach. Lakki, leg. Mayrhofer, 1976, GZU), I, L Szatala 1943b; Kleinig 1966; Hertel 1970b - C, This might also include the following referen­ I ces: Zahlbruckner 1906 (f. mosigii (Ach.) Nyl.); Lempholemma polyanthes (Bernh.) Malme Szatala 1943b (f. mosigii (Ach.) Nyl., f. subcon­ R (14 km W Spili, 1993, leg. Nordin, UPS) tiguella Vain., f. cechumenoides Vain.) Lepraria nivalis J.R. Laundon lecidea lapicida (Ach.) Ach. var. pantherina Ach. I (1,5 kn1 N Kounari, 1993, leg. Nordin, UPS) Syn. Lecidea subgrisella Nyl. Leprocaulon microscopicum (Vili.) Gams Kleinig 1966 - I C (3 km N Omalos, 1993, leg. Nordin, UPS) Lecidea promiscens Nyl. Leptogium biatorinum (Nyl.) Leight. Hetiel 1971, 1995 - R Syn. Leptogium pusillwn Nyl. The species occurs on carbonate-free, siliceous Steiner 1916- I Leptogiumferax (Durieu & Mont.) Rabenh. rocks in alpine situations. The delimitation from Arvidsson 1984 - I Lecidea auriculata is sometimes problematic leptogium gelatinosum (With.) J.R. Laundon (He1iel 1995). Syn. Leptogium sinuatum (Huds) A. Massa!. Lecidea sarcogynoides Korb. Kleinig 1966 - C Szatala 1943b - C Leptogium lichenoides (L.) Zahlbr. Lecidea xylophila Th. Fr. Szatala 1943b; Rondon 1969 (as var. pulvinatum Kleinig 1966 - C (Hoffm.)Zahlbr.) - L, R (7 km S Anogia, 1993, lecidella anomaloides (A. Massal.) Hertel & H. leg. Nordin, UPS, 3 kn1 ESE Vrises, 1993, leg. Kilias Nordin, UPS) Syn. Lecidea goniophila auct. Leptogiwn saturninmn(Dicks.) Nyl. Szatala 1943b- C, I R (7 km S Anogia, 1993, leg. Nordin, UPS) lecidella asema (Nyl.) Knoph & Hertel Leptogium schraderi (Bernh.) Nyl. Syn. Lecidea subincongrua Nyl. C (3 km N Omalos, 1993, leg. Nordin, UPS), I Rondon 1969 (2 km W Festos, 1993, leg. Nordin, UPS)

49 M. GRUBE, L. LiNDBLOM, H. MAYRHOFER

Leptogium subtile (Schrad.) Torss. Muellerella lichenicola (Sommerf.: Fr.) D. R (Adelaki, 12 km S Rethimno, 1993, leg. Hawksw. Nordin, UPS) Triebel 1989 - I Lethariella intricata (Moris) Krog Hosts: Catillaria detractula, Caloplaca lactea C (Gingilos, 1998, leg. J.W. Bjerke, hb. Bjerke) Muellerella pygmaea (Korb.) D. Hawksw. var. Lichenochora constrictella (Mi.ili.Arg.) Hafellner pygmaea L (Mavros Kolimbos, 1997, leg. Grube, GZU) Triebel 1989 - I Host: Fulgensia cf. subbracteata. Host: Acarospora sp. Lichenostigma elongata Nav.-Ros. & Hafellner Mycobilùnbia hypnorum (Lib.) Kalb & Hafellner Navarro-Rosinés & Hafellner 1996 - C Syn. Lecidea fusca (Schaer.) Th. Fr. Host: Lobothallia radiosa Szatala 1943b - L Lindauopsis caloplacae Zahlbr. Mycobilimbia lurida (Ach.) Hafellner & Ti.irk Vouaux 1914 Syn. Lecidea lurida Ach. Host: Caloplaca aurantia Szatala 1943b; Kleinig 1966; Papp et al. 1999 - Lithographa tesserata (DC.) Nyl. C, I, L Kleinig 1966 - L Myxobilimbia lobulata (Sommerf.) Hafellner Lobothallia radiosa (Hoffm.) Hafellner Syn. Toninia syncomista (Florke) Th. Fr. Syn. Lecanora radiosa (Hoffm.) Schaer., Szatala 1943b - RJI Lecanora radiosa var. subcircinata (Nyl.) Zahlbr., Lecanora subcircinata Nyl., Naetrocymbe fraxini (A. Massa!.) R.C. Harris Squamarina subcircinata (Nyl.) Kiffer Syn. Arthopyrenia.fi'axini A. Massa!. Zahlbruckner 1906; Szatala 1943b (as R (Adelaki, 12 km S Rethimno, 1993, leg. Squamaria radiosa (Hoffìn.) Poetsch); Kleinig Nordin, UPS) 1966; Rondon 1969; Navarro-Rosinés & Naetrocymbe punct�formis (Pers.) R.C. Harris Hafellner 1996; Papp et al. 1999 - C ( 1 km S Syn. Arthopyrenia punctiformis (Pers.) A. lrnbros, leg. Mayrhofer & Eitl, GZU), RJI, I, L Massa!. Rondon 1969 Megaspora verrucosa (Ach.) Hafellner & V. Wirth Neofuscelia attica (Leuckert, Poelt & B. Schwartz) Syn. Aspicilia mutabilis (Ach.) Korb.; Aspicilia Essi. verrucosa (Ach.) Korb. Leucke1t et al. 1975 - C (Street to Ornalos, 500 Szatala 1943b; Kleinig 1966- C (Ornalos, 1998, m SW Lakki, 1976, leg. Mayrhofer, GZU), I leg. Trinkaus & Grube, GZU; Xiloskalo, 1993, Neofi,scelia delisei (Duby) Essi. leg. Nordin, UPS), R (7 km S Anogia, 1993, leg. Syn. Parmelia delisei (Duby) Nyl. Nordin, UPS), I, L Szatala 1943b - RJI, I, L The species is common on bark, however, the Neofuscelia glabrans (Nyl.) Essi. variety mutabilis is not distinguished as its value Syn. Parmelia pulla Ach. ssp. glabrans (Nyl.) is unclear. Clauzade & Cl. Roux, Parmelia glabrans Nyl. Melanelia exasperata (De Not.) Essi. Rondon 1969 Syn. Parmelia aspidota (Ach.) Poetsch Neofitscelia loxodes (Nyl.) Essi. Szatala 1943b - C, R (7 km S Anogia, 1993, leg. Syn. Parmelia isidiotyla Nyl. Nordin, UPS), R/I Szatala 1943b, Leuckett et al. 1975 - L Melanelia glabra (Schaer.) Essi. Neofitscelia pokornyi (Korb.) Essi. Syn. Parmelia glabra Schaer. Syn. Parmelia pokornyi (Korb.) Szatala Zahlbruckner 1906; Szatala 1943b - C, R Kleinig 1966 - C (Skinakas, 1997, leg. Suppan, GZU; 7 km S Neofi,scelia pulla (Ach.) Essi. Anogia, 1993, leg. Nordin, UPS), RJI Syn. Parmelia prolixa (Ach.) Carroli Melanelia laciniatula (Flagey ex H. Olivier) Essi. Zahlbruckner 1906; Szatala 1943b; Kleinig Syn. Parmelia laciniatula (Flagey ex Zahlbr.) 1966; Esslinger 1977 (rnap); Papp et al. 1999 - Zahlbr. R (1 km S Kouroutes, 1976, leg. Mayhofer, Kleinig 1966 - C, R GZU), I, L Micarea mise/la (Nyl.) Hedl. Nephroma laevigatum Ach. R (7 km S Anogia, 1993, leg. Nordin, UPS) Kleinig 1966 - C

50 Studia Geobot. 20 (200 I) - A check-list of Crete lichens -

Nonnandina pulchella (Borrer) Nyl. Peltigera polydactyla (Neck.) Hoffm. s. lat. C (1km E Vlatos, 1998, leg. Trinkaus & Grube, Kleinig 1966 - C, I GZU) Peltigera praetextata (F\orke ex Sommerf.) Zopf Vitikainen 1994 (map) Ochrolechia balcanica Verseghy Peltigera rufescens (Weiss) Humb. C (Omalos, 1976, leg. Mayrhofer, GZU) R (7 km S Anogia, 1993, leg. Nordin, UPS) Ochrolechia pallescens (L.) A. Massal. Pertusaria albescens (Huds.) M. Choisy & Werner Szatala 1943b; Kleinig 1966 - R Syn. Pertusaria discoidea (Pers.) Malme, P glo­ Ochrolechia parella (L.) A. Massa!. bulifera (Turner)A. Massa!. Zahlbruckner 1906; Szatala 1943b; Kleinig Szatala 1943b; Kleinig 1966 - C, I, L 1966; Rondon 1969; Verseghy 1962 (var. la-e­ Pertusaria chiodectonoides Bagl. ex A. Massa!. taensis Verseghy) - I, L Syn. Pertusaria inquinata (Ach.) Th. Fr. Ochrolechia tartarea (L.) A. Massa!. Szatala 1943b - L Zahlbruckner 1906 - R, L Pertusaria dalmatica Erichsen Ochrolechia turneri (Sm. & Sowerby) Hasselrot Szatala 1943b; Schmitz et al. 1994 - C R (7 km S Anogia, 1993, leg. Nordin, UPS) Opegrapha calcarea Sm. Pertusaria jlavicans Lamy Syn. Opegrapha chevallieri Leight. Szatala 1943b; Rondon 1969; Hanko 1983 - L Zahlbruckner 1906; Szatala 1943b; Rondon Pertusaria hemisphaerica (Florke) Erichsen 1969 - I, L Szatala 1943b - C, I Opegrapha durieui Mont. Pertusaria hymenea (Ach.) Schaer. Syn. Enterographa durieui (Mont.) Redgr. Syn. Pertusaria lecanorodes Erichsen, Pertu­ Szatala 1943b; Roux 1991 (map) - L saria wulfenii DC. Opegrapha rupestris Pers. Szatala 1943b; Kleinig 1966- C, R (Adelaki, 12 Syn. Opegrapha saxatilis DC.; Opegrapha saxi­ km S Rethimno, 1993, leg. Nordin, UPS), RII, I cola Ach. Pertusaria leucostoma A. Massa!. Kleinig 1966 - I (1,5 kn1 N Kounari, 1993, leg. Syn. Pertusaria leioplaca DC. Nordin, UPS), L Szatala 1943b - L Opegrapha varia Pers. Pertusaria pentelisi J. Steiner R (3km ESE Vrises, 1993, leg. Nordin, UPS) Syn. Melanaria pentelisi (J. Steiner) Erichsen Opegrapha variaefonnis Anzi ("pentelici") Kleinig 1966 - L Szatala 1943b; Rondon 1969 - I Pertusaria pertusa auct. Parmelia sulcata Taylor Szatala 1943b; Kleinig 1966; Hanko 1983 - C, Kleinig 1966 - C RII, I, L Parmelina carporrhizans (Taylor) Poelt & Vezda Pertusaria pluripuncta Nyl. R (Adelaki, 12 km S Rethimno, 1993, leg. Syn. Pertusaria gallica De Lesd. Nordin, UPS) L (1 km W Karidi, 1976, leg. Mayrhofer, GZU) Parmelina pastillifera(Harm.) Hale Pertusaria rupicola (Fr.) Harm. C (Omalos, 1993, leg. Nordin, UPS), R Szatala 1943b; Rondon 1969; Leuckert et al. (Skinakas, 1997, leg. Suppan, GZU; 7 kn1 S 1969; Hanko 1983 - I Anogia, 1993, leg. Nordin, UPS) Parmelina tiliacea (Hoffm.) Hale Petractis clausa (Hoffm.) Kremp. Syn. Parmelia scortea (Ach.) Ach. Zah\bruckner 1906; Steiner 1916 (var. eradiata Zahlbruckner 1906; Szatala 1943b; Kleinig J. Steiner); Vezda 1965 - I, L 1966; Rondon 1969; Czeczuga et al. 1994 - R Phaeophyscia hirsuta (Mereschk.) Essi. (Adelaki, 12 kn1 S Rethimno, 1993, leg. Nordin, R (Adelaki, 12 km S Rethimno, 1993, leg. UPS), I, S Nordin, UPS) Peltigera canina (L.) Willd. Phaeophyscia orbicularis (Neck.) Moberg Kleinig 1966; Vitikainen 1994 (map). - C, RII C (Agios Nektarios, 1993, leg. Nordin, UPS), R Peltigera collina (Ach.) Schrad. (7 km S Anogia, 1993, leg. Nordin, UPS) Syn. Peltigera scutata (Dicks.) Duby Physcia adscendens (Fr.) H. Olivier Kleinig 1966 - RII Kleinig 1966; Rondon 1969 - C, I, L

51 M. GRUBE, L. LINDBLOM, H. MAYRI·IOFER

Physcia aipolia (Ehrh. ex Humb.) Hampe Placolecis opaca (Fr.) Hafellner Kleinig 1966 - R (Skinakas, 1997, leg. Suppan, Syn. Astroplaca opaca (Fr.) Bagl. GZU), L Szatala 1943b - C Physcia albinea (Ach.) Nyl. Placidium lachneum (Ach.) Breuss Szatala 1943b - L Syn. Catapyrenium lachneum (Ach.) R. Sant., Physcia biziana (A. Massal.) Zahlbr. Dermatocarpon lachnewn (Ach.) A.L. Sm. Szatala 1943b; Kleinig 1966; Rondon 1969- C, Kleinig 1966 - C, I, L R (6 km Se Nea Kria Vrisi, 1997, leg. Mayrhofer Placidium lacinulatum (Ach.) Breuss & Erti, GZU; 2 km NW Akoumia, 1993, leg. Syn. Catapyrenium lacinulatum (Ach.) Breuss Nordin, UPS), L (W ofExo Lakonia, leg. Degel., R (7 km SE Garazo, 1993, leg. Nordin, UPS) UPS) Placidium pilosellmn (Breuss) Breuss Physcia caesia var. caesiella (de Lesd.) Clauzade & Syn. Catapyrenium pilosellum Breuss Cl. Roux Breuss 1990 - L R (1km S Kouroutes, 1976, leg. Mayrhofer, Placidium squamulosum (Ach.) Breuss GZU) Syn. Dermatocarpon hepaticum (Ach.) Th. Fr., Physcia leptalea (Ach.) DC. Catapyrenium squamulosum (Ach.) Breuss Syn. Physcia semipinnata (J.F. Gmelin) Moberg Szatala 1943b; Kleinig 1966; Breuss 1990 - C, Zahlbruckner 1906 (stellaris var. leptalea Nyl.); I, L Szatala 1943b; Kleinig 1966; Rondon 1969; Placynthium nigrum (Huds.) Gray Vezda 1995; Moberg 1994 - C, R (Skinakas, Kleinig 1966 - C, I 1997, leg. Suppan, GZU; 6 km NW Apostoli, Platismatia glauca (L.) W.L. Culb. & C.F. Culb. 1993, leg. Nordin, UPS) C (Gingilos, 1998, leg. J.W. Bjerke, herb. Physcia scopulorum (Lambinon & Vezda) Poelt & Bjerke) Nimis Pleopsidium_flavum (Bellardi) Korb. L (1,5 km S Kalavros, 1999, leg. Grube, GZU; Syn. Acarospora oxytona (Ach.) A. Massa!. filed under Arthonia epiphyscia) Kleinig 1966 - C, RII Physcia stellaris (L.) Nyl. Pleurosticta acetabulum (Neck.) Elix & Lumbsch Szatala 1943b (var. rosulata (Ach.) Hue); Syn. Parmelia acetabulum (Neck.) Duby Kleinig 1966- C, R/I Zahlbruckner 1906; Szatala 1943b; Kleinig 1966 Physcia wainioi Rasanen - C, R (Skinakas, 1997, leg. Suppan, GZU), RII, Papp et al. 1999 - C R (7 kn1 S Anogia, 1993, leg. Nordin, UPS), L Physconia detersa (Nyl.) Poelt Polyblastia terrestris Th. Fr. Kleinig 1966 - C Physconia distorta (With.) J.R. Laundon C (Agios Nektarios, 1993, leg. A. Nordin, UPS) C (Gingilos, 1998, leg. J.W. Bjerke, hb. Bjerke; Polycoccum marmoratum (Kremp.) D. Hawksw. Imbros gorge, 1998, leg. J.W. Bjerke, hb. Syn. Microthelia mormorata (Kremp.) Korb. Bjerke) Szatala 1943b; Kleinig 1966 - C, RII Physconia grisea (Lam.) Poelt Polycoccum sp. Kleinig 1966 - C R (Agios Pavlos, leg. Grube, GZU) Physconia pulverulenta (Anzi) Jatta Host: Melanelia sp. Syn. Physcia pulverulenta (Schreb.) Nyl. f. Polycoccum sp. argyphaea and var. subvenusta This species is distinct from any described spe­ Zahlbruckner 1906 - RII, L cies on Physciaceae, including the species men­ Physconia servitii (Nadv.) Poelt tioned by Matzer et al. ( 1997), which is probably R (Adelaki, 12 km S Rethimno, 1993, leg. identica! to Didymosphaeria echinospora Nordin, UPS) Faure!. Physconia subpulverulenta (Szatala) Poelt Host: Rinodina obnascens Poelt 1966 - C Polysporina simplex (Davies) Vezda Physconia venusta (Ach.) Poelt Syn. Sarcogyne simplex (Davies) Nyl. Syn. Physcia venusta (Ach.) Nyl. Zahlbruckner 1906; Szatala 1943b; Rondon Szatala 1943b (+ var. subvenusta (Nyl.) Oliv.); 1969- C, L Kleinig 1966 - C, R/I, R (7 kn1 S Anogia, 1993, Porina linearis (Leight.) Zahlbr. leg. Nordin, UPS) Kleinig 1966; Rondon 1969 - I

52 Studia Geobot. 20 (2001) - A check-list of Crete lichens -

Porpidia albocaerulescens (Wulfen) Hertel & Psora testacea (Hoffm.) Ach. Knoph Syn. Lecidea testacea (Hoffm.) Ach. Syn. Lecidea albocaerulescens (Wulfen)Ach. Papp et al. 1999 - C (2 km N Omalos, 1988, leg Zahlbruckner 1906 (+f. alpina Schaer.); Rondon Timdal, hb. Timdal), R (7 km Se Garazo, 1993, 1969- L leg. Nordin, UPS), I (Iouhtas, 1995, leg. Rui & Porpidia crustulata (Ach. ) Hertel & Knoph Timdal, hb. Timdal), L Hertel 1992, Papp et al. 1999- C, I Psora vallesiaca (Schaer.) Timdal Porpidia macrocarpa (DC.) Hertel & A.J. Schwab Syn. Lecidea deceptoria Nyl. Syn. Lecidea macrocarpa (DC.) Steud., Lecidea Kleinig 1966 - C, L (between Tzermiadhes and contigua auct. Neapolis, 1995, leg Rui & Timdal, hb. Timdal) Zahlbruckner 1906 (var. platycarpa (Ach.) Th. Psorotichia schaereri (A. Massa!.) Arnold Rondon 1969 - C (Agios Nektarios, 1993, leg. Fr.); Kleinig 1966- C, I, L Nordin, UPS) Protoblastenia calva (Dicks.) Zahlbr. Pyrenula chlorospila (Nyl.) Arnold Kleinig 1966- C, R, I, L C (1km E Vlatos, 1998, leg. Trinkaus & Grube, Protoblastenia incrustans (DC.) J. Steiner GZU) Kleinig 1966 - L Protoparmelia badia (Hoffm.) Hafellner Ramalina breviuscula Nyl. Syn. Lecanora badia (Hoffm.) Ach. L (1 km W ofK.aridi, 1976, H. Mayrhofer, GZU) Kleinig 1966- C, L Ramalina calicaris (L.) Fr. Protoparmelia montagnei (Fr.) Poelt & Nimis ? Ramalina graeca Mtill. Arg. Syn. Lecanora montagnei (Fr.) Schaer. Szatala 1943b; Kleinig 1966 - RII Szatala 1943b - I, L (1km W Karidi, 1976, leg. Ramalina farinacea (L.) Ach. Mayrhofer, GZU) Szatala 1943b (var. gracilenta Ach.); Kleinig Protoparmelia psarophana (Nyl.) Sancho & 1966 - I Crespo Ramalinajastigiata (Pers.) Ach. Syn. Lecanora psarophana Nyl. Kleinig 1966 - C, RII, I Szatala 1943b (var. reagens Servit.); Rondon Ramalinafraxinea (L.) Ach. 1969 ( + var. reagens)- L Kleinig 1966 - C (Omalos, 1976, leg. Pseudeverniaji1rfi1racea (L.) Zopf var. fitrfuracea Mayrhofer, GZU), R, RII Syn. Parmeliafurfi1racea (L.) Ach. Ramalina pollinaria (Westr.) Ach. Kleinig 1966- C Rondon 1969 Pseudevernia fitrfitracea (L.) Zopf var. ceratea Ramalina polymorpha (Lilj.) Ach. (Ach.) D. Hawksw. Szatala 1943b (f. emplecta Ach.)- S Syn. Parmelia olivetorina (Zopf) Zopf; Ramalina requienii (De Not.) Jatta Parmeliafurfùracea var. olivetorina Szatala 1943b; Rondon 1969; Vezda 1969 - I, L Zahlbruckner 1906; Szatala 1943b - C, R Rhiwcarpon distinctum Th. Fr. Psora decipiens (Hedw.) Hoffm. var. decipiens Zahlbruckner 1906; Szatala 1943b; Feuerer 1991 - I, L Syn. Lecidea decipiens (Hedw.) Ach. Rhiwcarpon geographicum (L.) DC. ssp. geo­ Zahlbruckner 1906; Steiner 1916; Szatala graphicum 1943b; Kleinig 1966; Rondon 1969; Schneider Zahlbruckner 1906 (f. contiguum Fw.); Szatala 1979, Papp et al. 1999 - C, R (Psiloritis, 1988, 1943b (f. contiguum Fw., f. atrovirens (L.) leg. Timdal, hb. Timdal), I, L Massal.); Kleinig 1966; Rondon 1969 - C, R (7 Psora decipiens var. galactina Zahlbr. km S Anogia, 1993, leg. Nordin, UPS), RII, I, L Zahlbruckner 1906; Kleinig 1966; Schneider Rhiwcarpon geographicum ssp. kittilense (Ra­ 1979; Timdal 1984 - C, I sanen) Sant. ad int. Psora gresinonis de Lesd. Syn. Rhizocarpon riparium Rasanen Syn. Lecidea gresinonis (de Lesd.) Zahlbr. Szatala 1943b - RII Kleinig 1966; Timdal 1984 - L Rhizocarpon macrosporum Rasanen Psora pseudorussellii Timdal Runemark 1956 - RII C (2 km N of Omalos, 1988, leg. Timdal, hb. Rhiwcarpon polycarpum (Hepp) Th. Fr. Timdal) Szatala 1943b- I

53 M. GRUBE, L. L!NDBLOM, H. MAYRHOFER

Rhizocarpon tinei (Toma6.) Runemark Rinodina trachytica (A. Massa!.) Bagl. & Carestia Runemark 1956, Rertel 1983 - I Mayrhofer 1984 (map); Mayrhofer & Leuckert Rhizocarpon viridiatrum (Wulfen) Kor6. 1985; Mayrhofer & Poelt 1979; Mayrhofer et al. Kleinig 1966 - L 1992 (map) - R Rimularia insularis (Nyl.) Ram6old & Reriel Rinodina tunicata R. Mayrhofer & Poelt Syn. Lecidea intumescens (Florke) Nyl. O6ermayer 1999; Reims et al. 2001 - C Szatala 19436; Rertel 1970a (map); Rertel & Rinodinella controversa (A. Massa!.) R. Mayrhofer Ram6old 1990 - R/1, I & Poelt Rinodina alba Metzler ex Amold Syn. Rinodina crustulata (A. Massa!.) Arnold Szatala 19436; Mayrhofer 1984 (map); Mayr­ Szatala 19436 (as Rinodina atrocinerea (Rook.) hofer & Poelt 1978; Reims et al. 2001 - C, I Kor6.); Mayrhofer 1984 - L Rinodinella dubyanoides (Repp) R. Mayrhofer & Rinodina bischojjii (Repp) A. Massa!. Poelt Mayrhofer & Poelt 1979; Mayrhofer 1984 (map) Mayrhofer& Poelt 1978; Mayrhofer 1984 (map) - C (1 km S lm6ros, leg. Mayrhofer & Erti, - I, C (1 kn1 S lmbros, leg. Mayrhofer & Erti, GZU), L GZU) Rinodina calcarea (Amold) Amold Roccella phycopsis Ach. Poelt & Mayrhofer 1979; Mayrhofer & Poelt Syn. Roccella.fì1coides Vain. 1979; Mayrhofer 1984 (map) - L Zahl6ruckner 1904; Zahlbruckner 1906; Szatala Rinodina confragosa (Ach.) Kor6. 19436; Kleinig 1966; Rondon 1969 - C, I, L Szatala 19436 (+ var. fi1mosa (Wedd.) R. Olivier); Mayrhofer 1984 (map) - I, L Sagiolechia protuberans (Ach.) A. Massa!. Rinodina eretica R. Mayrhofer Kleinig 1966 - L Mayrhofer 1984 - I, L Sarcogyne privigna (Ach.) A. Massa!. Rinodina dubyana (Repp) J. Steiner C (3 km N Omalos, 1993, leg. Nordin, UPS), R Mayrhofer 1984 (map) - L (7 km S Anogia, 1993, leg. Nordin, UPS) Sarcogyne regularis Kor6. Rinodina exigua (Ach.) Gray Syn. Sarcogyne pruinosa auct. R (7 km S Anogia, 1993, leg. Nordin, UPS) Steiner 1916; Szatala 19436; Kleinig 1966 - C, Rinodina gennarii Bagl. R, I, L Syn. Rinodina demissa auct. Solenopsora candicans (Dicks.) J. Steiner Zahl6ruckner 1906; Szatala 19436; Mayrhofer Syn. Placolecania candicans (Dicks.) Zahl6r. 1984 (map) - R, L Zahl6ruckner 1906; Szatala 1943b; Kleinig Rinodina immersa (Kor6.) Amold 1966; Rondon 1969 - C (road from Voukolies to Syn. Rinodina bischo.ffi (Repp) A. Massa!. var. Paleochora, 1998, leg. Rui & Timdal, h6 immersa Kor6. Timdal), I, L Zahl6ruckner 1906; Szatala 19436; Rondon Solenopsora cesatii (A. Massa!.) Zahl6r. 1969; Mayrhofer & Poelt 1979; Mayrhofer1984 Syn. Placolecania cesatii (A. Massa!.) Zahl6r.; (map) - C (1 km S Im6ros, leg. Mayrhofer & Solenopsora bagliettoana Tav. ined.; Soleno­ Erti, GZU), I, L psora cesatii (A. Massa!.) Zahl6r. var. grisea Rinodina lecanorina (A. Massa!.) A. Massa!. (Bagl.) Nimis Syn. Rinodina ocellata (Roffm.) Amold Zahl6ruckner 1906; Szatala 19436 - C, R (14 Mayrhoferet al. 1990; Mayrhofer& Poelt 1979; km W Spili, 1993, leg. Nordin, UPS), I, L The variety grisea is not distinguished, as inter­ Mayrhofer 1984 (map) - I, L mediate forms links it with the type variety. Rinodina mi/vina (Wahlen6.) Th. Fr. Solenopsora marina ined. Szatala 1943b; Kleinig 1966; Mayrhofer 1984 C (road from Voukolies to Paleochora, 1998, leg. (map) - 1/R Rui & Timdal, h6. Timdal) Rinodina obnascens (Nyl.) R. Olivier Solenopsora olivacea (Fr.) R. Kilias I (Mate, 1997, leg. Gru6e, GZU). Syn. Placodiella olivacea (Fr.) Szatala; Catil­ Rinodina pyrina (Ach.) Arnold laria olivacea (Fr.) Zahl6r. Szatala 19436; Kleinig 1966 - C Zahl6ruckner 1906; Rondon 1969; Szatala Rinodina sophodes (Ach.) A. Massa!. 19436; Roux 1991 (map)- C, R (14 km W Spili, Kleinig 1966; Giralt & Mayrhofer 1995 - C 1993, leg. Nordin, UPS), I, L

54 Studia Geobot. 20 (200 I) - A check-list of Crete lichens -

Solenopsora vulturiensis Bagl. Tephromela atra (Huds.) Hafellner var. atra L (Karidi, 1976, leg. Mayrhofer, GZU) Syn. Lecanora atra (Huds.) Ach. Squamarina cartilaginea (With.) P. James f. carti­ Zahlbruckner 1906; Szatala 1943b; Kleinig laginea 1966; Rondon 1969 (+ var. disco/or (Duby) Syn. Squamarina crassa (Huds.) Poelt Schaer.) - R ( 1 km S Kouroutes, 1976, leg. Zahlbruckner 1906 (as Lecanora); Steiner 1916 Mayrhofer, GZU), RII, I, L (as Lecanora, f. dealbata); Szatala 1943b (var. Tephromela atra (Huds.) Hafellner var. cypria caespitosa (Vili.) Rabenh., var. liparia (Ach.) (Korb.) Nimis C (1 km S Imbros, leg. Mayrhofer & Erti, GZU), Boistel); Kleinig 1966; Rondon 1969, Papp et al. I (Dia island, 1942, Rechinger, W; Nida plain, 1999 - C, R, I, L 1942, Rechinger, W), L (Street from Adravasti to Squamarina cartilaginea f. iberica (Mattick) Karidi, 1976, Mayrhofer & Polzl, GZU) Clauzade & Cl. Roux Thelenella modesta (Nyl.) Nyl. Syn. Squamarina crassa f. iberica (Mattick) Syn. Microglaena modesta (Nyl.) A.L. Sm. Poelt Szatala 1943b; Mayrhofer & Poelt 1985; Kleinig 1966 - C Mayrhofer 1987 - C Squamarina concrescens (Muli. Arg.) Poelt Thelidium creticum J. Steiner Rondon 1969; Poelt & Kriiger 1970 (map ); Papp Steiner 1916; Szatala 1943b - I et al. 1999 - C (Agios Nektarios, 1993, leg. A. Thelopsis isiaca Stizenb. Nordin, UPS), I (louhtas, 1995, leg Rui & Szatala 1943b; Vè'zda 1968 - C Timdal, hb. Timdal), R, L (4km N Mirtos, 1976, Toninia albilabra (Dufour) H. Olivier leg. Mayrhofer, GZU) Timdal 1991 (map) - C (between Vouklies and Squamarina gypsacea (Sm.) Poelt Kandanos, 1988, leg. Timdal, hb. Timdal) Syn. Squamarina fragilis (Scop.) Szatala Toninia aromatica (Sm.) A. Massa!. Kleinig 1966; Rondon 1969; Szatala 1943b; Steiner 1916; Szatala 1943b; Kleinig 1966; Papp et al. 1999 - C, I (louhtas, 1995, leg Rui & Timdal 1991 (map) - C, R (2.5 km S of Koxare, 1988, leg. Timdal, hb. Timdal), I Timdal, hb. Timdal), R, L (between Adravasti Toninia candida (Weber) Th. Fr. and Karidi, 1976, leg. Mayrhofer & Polzl, GZU) Szatala 1943b; Rondon 1969; Timdal 1991 Squamarina lentigera (Weber) Poelt (map); Papp et al. 1999 - C (Agios Nektarios, Syn. Lecanora lentigera (Weber) Ach. 1993, leg. Nordin, UPS), R, I (hill just S of Steiner 1916; Szatala 1943b; Poelt & Kriiger Fodele, 1988, leg. Timdal, hb. Timdal), L 1970 (map ); Papp et al. 1999 - C, R, I, L Toninia cinereovirens (Schaer.) A. Massa!. Squamarina oleosa (Zahlbr.) Poelt Timdal 1991 (map) - I (near Epanosifi Rondon 1969 Monastery, 1988, leg. Timdal, hb. Timdal) Staurothele fuscocuprea (Nyl.) Zschacke Toninia eretica Timdal Syn. Staurothele clopima (Wahlenb.) Th. Fr. Timdal 1991 (map) - C (2 km N of Omalos, Szatala 1943b - R 1988, leg. Timdal, hb. Tirndal), R (Psi!Òritis, Staurothele hymenogonia (Nyl.) Th. Fr. 1988, leg. Timdal, hb. Timdal) Syn. Staurothele ventosa (A. Massa!.) P. Syd., Toninia dijfracta (A. Massa!.) Zahlbr. Staurothele extabescens (Nyl.) Zahlbr. C (Omalos, 1976, leg. Mayrhofer, GZU), R Steiner 1916; Szatala 1943b - I, L (Psiloritis, 1988, leg. Timdal, hb. Timdal), I Staurothele immersa (A. Massa!.) Dalla Torre & (Matala, 1997, leg. Grube, GZU; filed under Sarnth. Stigmidium tabacinae) Toninia lutosa (Ach.) Timdal Rondon 1969 C (Agios Nektarios, 1993, leg. Nordin, UPS) Stigmidimn tabacinae (Arnold) Triebel Toninia massata (Tuck.) Herre I (Matala, 1997, leg. Grube, GZU) Timdal 1991 (map) - I (Rogdia, 1965, leg. Host: Toninia diffi-acta Degelius, UPS) Strigula affinis (A. Massa!.) R.C. Harris Toninia opuntioides (Vili.) Timdal Syn. Porina affìnis (A. Massa!.) Zahlbr. Timdal 1991 (map) - C (lmbros, 1985, leg. Szatala 1943b - C Hoiland, O), R (Psiloritis, 1988, leg. Timdal, hb. Synalissa symphorea (Ach.) Nyl. Timdal), L (Gournia, 1988, leg. Timdal, hb. Szatala 1943b - C Timdal)

55 M. GRUBE, L. LINDBLOM, H. MAYRHOFER

Toninia philippea (Mont.) Timdal Usnea articu/ata (L.) Hoffm. Syn. Catillaria philippea (Mont.) A. Massa!.; Szatala 1943a (ssp. mediterranea Motyka) -C Catillaria lutosa A. Massa!. Usnea filipendula Stirt. Zahlbruckner 1906 - L Syn. Usnea dasypoga auct. Not on Crete ace. to Timdal 1991 (maps) Kleinig 1966 - C Toninia physaroides (Opiz) Zahlbr. Timdal 1991 (map) Verrucaria acrotella Ach. Toninia sedifolia (Scop.) Timdal Kleinig 1966 -L Syn. Tonii1ia coeruleonigricans Th. Fr. Verrucaria adriatica Zahlbr. Steiner 1916; Szatala 1943b; Kleinig 1966; Szatala 1943b - L Rondon 1969; Timdal 1991 (map); Papp et al. Verrucaria attica J. Steiner 1999 - C, R,I, L Steiner 1916,Kleinig 1966 - I Toninia squalida (Ach.) A. Massa!. Verrucaria aspiciliico/a R. Sant. Syn. Toninia acervulans (Nyl.) Oliv. Syn. Verrucaria. aspiciliae Zehetl. Rondon 1969 Zehetleitner 1978 -L Not mapped on Crete in Timdal (1991) Host: Aspicilia calcarea Toninia taurica (Szatala) Oxner Verrucaria caerulea DC. Timdal 1991 (map) - C (ca. 2.5 km N of Syn. Verrucaria plumbea Ach., lnvolucrothele Omalos, 1998, leg. Rui & Timdal, hb. Timdal), plumbea (Ach.) Servit R (Psiloritis, 1988,leg. Timdal,hb. Timdal) Kleinig 1966-C Toninia tristis (Th. Fr.) Th. Fr. ssp. tristis Verrucaria cyanea A. Massa!. Timdal 1991 (map)-C (Agios Nektarios, 1993, Syn. Verrucaria decussata Garov. Zahlbruckner 1906 -I leg. Nordin,UPS) Verrucaria foveolata (Florke) A. Massa!. Toninia tristis ssp. asiae-centralis (H. Magn.) Syn. Verrucaria dolomitico/a (A. Massal.) Timdal Kremp. C (Imbros,1988, leg. Timdal,hb. Timdal) Rondon 1969 - C (Agios Nektarios, 1993, leg. Toninia tristis ssp. pseudotabacina Timdal Nordin, UPS) R (2 km E of Preveli Monastery, 1988, leg. Verrucariafuscella (Tumer) Winch Timdal, hb. Timdal), L (Goumia, 1988, leg. Zahlbruckner 1906 -L Timdal,hb. Timdal) Verrucariaji,scula Nyl. Toninia tristis ssp. thalloedaemiformis (Szatala) Syn. Dermatocarpon insulare (A. Massa!.) Mig. Timdal Rondon 1969 Syn. Lecidea thalloedaeformis Szatala Verrucaria hochstetteri Fr. var. mastoidea (A. Szatala 1943b; Timdal 1991 - C (Imbros, 1988, Massa!.) Clauzade & Cl. Roux leg. Timdal, hb. Timdal), R (Fourfouras, 1985, Syn. Amphoridium mastoideum A. Massa!. leg. H0iland, O), I (hill just S of Fodele, 1988, Steiner 1916-I leg. Timdal,hb. Timdal),L Verrucaria macrostoma Dufour ex DC. Toninia tumidula (Sm.) Zahlbr. Steiner 1916; Kleinig 1966; Rondon 1969 - I C (1 km W ofKakopetros,1988, leg Timdal,hb. Verrucaria marmorea (Scop.) Amold Timdal), I (Iouhtas, 1995, leg. Timdal, hb. Syn. Amphoridium marmoreum (Scop.) Baroni Timdal) Zahlbruckner 1906; Kleinig 1966; Rondon 1969 Toninia verrucariae (Nyl.) Timdal -C,L I (Kounari, leg. Nordin, 1993) Verrucaria mortarii Lamy Host: Bagliettoa parmigera Rondon 1969 Topelia heterospora (Zahlbr.) P.M. 10rg. & Vezda Verrucaria muralis Ach. Syn. Clathroporina heterospora Zahlbr. Syn. Verrucaria rupestris Schrad. non (Scop.) Szatala 1943b; Vezda 1968 -I Weber ex. F.H. Wigg. Tornabea scutellifera (With.) J. R. Laundon Zahlbruckner 1906; Steiner 1916 (var. I (Iouhtas,1995, Rui & Timdal,O) hypophaeaSteiner & Zahlbr.); Rondon 1969- I, Trapelia obtegens (Th. Fr.) Hertel L Syn. Lecidea obtegens (Th. Fr.) Vain. Verrucaria murina Leight. Szatala 1943b -L Steiner 1916 -I

56 Studia Geobot. 20 (200 I) - A check-list of Crete lichens -

Verrucaria nigrescens Pers. Collema sublimosa J. Steiner Szatala 1943b; Rondon 1969 - C, I (2 km W Steiner 1916 - I Festos, 1993, leg. Nordin, UPS), L Probably a member of the C. tenax group. See Verrucaria ruderum DC. Degelius 1954: 461 (materiai not seen) Rondon 1969 Lecanora coerulata (Ach.) Szatala Szatala 1943b - I, L Xanthoparmelia conspersa (Ehrh. ex Ach.) Hale Lecidea ocellulata (Schaer.) Th. Fr. Syn. Parmelia conspersa (Ach.) Ach., Parmelia Szatala 1943b - I isidiata (Anzi) Gyeln. This taxon could belong to Lecidea .fiiscoatra Szatala 1943b; Kleinig 1966 - I (L.) Ach. Xanthoparmelia somloensis (Gyeln.) Hale Syn. Parmelia somloensis Gyeln.; Parmelia ste­ Acknowledgments nophylla (Ach.) auct. Thanks are due to Roland Moberg (Uppsala) for access to the her­ Rondon 1969 barium database of Fytoteket Uppsala. Einar Timdal and Jarle Werner Bjerke are acknowledged for providing further informa­ Xanthoparmelia tinctina (Maheu et Gillet) Hale tion. Brian Coppins, Ulli Grube, Josef Hafellner, and Pier Luigi Papp et al. 1999 - C Nimis for discussions and comments on the manuscript. Xanthoparmelia verrucigera (Nyl.) Hale References Syn. Parmelia conspersa var. verrucigera (Nyl.) Boistel; Parmelia verrucigera Nyl. Arvidsson L., 1984. Two new records of Leptogium ferax. Steiner 1916 - I Lichenologist, 16: 91-92. Bellemère A., 1994. Documents et commentaires sur l'ultrnstruc­ Xanthoria calcico/a Oxner ture des asques polysporés des Acarospora, de que/ques genres Syn. Xanthoria aureola (Ach.) Erichsen de la famille des Acarosporaceae et de genres similaires. Bui!. Kleinig 1966; Rondon 1969 (+ var. ectanea Soc. Linn. Provence, 45: 355-388. Breuss O., 1990. Die Flechtengattung Catapyrenium (Ach.) Kickx.) - C, R (6 km Se Nea Kria Vrisi, (Verrucariaceae) in Europa. Slapfia, 23: 1-153 + 22 unnumbe­ leg. Mayrhofer & Erti, GZU; 2 km W Festos, red pages. 1993, leg. Nordin, UPS), L Creutzberg N., Drooger C.W., Meulenkamp J.E., Papaslamaliou J., Xanthoria mediterranea Giralt, Poelt & Nimis Sannemann W., Seidel E. & Talares A., 1977. Genera/ geolo­ gica/ map of the is/and Crete, I: 200 000. Alhen. Syn. Xanthoria isidioidea auct. pro parte Czeczuga B. Amezquila M.M., Grillo M., Breuss O. & Jacobsen Kleinig 1966; Giralt et al. 1993 - C, I P., 1994. Carotenoids in lichens from some Mediterranean Xanthoria parietina (L.) Th. Fr. isfands. Ann. Mus. Goulandris, 9: 99-112. Degelius G., 1954. The lichen genus Collema in Europe: mo1pho­ Zahlbrnckner 1906 (+ var. retirugosa J. Steiner); Jogy, taxonomy, ecology. Symb. Bol. Upsal., I 3: 1-499. Kleinig 1966 C (Omalos, 1976, leg. Eigler G., 1969. Studien zur G/iedenmg der Flechtengattung Mayrhofer, GZU), I Lecanora. Diss. Bot., 4: 1-195 + 30 tabs. Esslinger Th.L., 1977. A chemosystematic revision of the brown Parmeliae. J. Hattori Bot. Lab., 42: 1-211. Zwackhiomyces inconspicuus Grube & Hafellner Feuerer T., 1991. Revision der europiiischen Arten der Grube & Hafellner 1990 - RII F/echtengattung Rhizocarpon mit nicht gelbem Lager und Bibliolh. Lichenol., 139: 1-218. Host: Lecanora dispersa. vie/zelligen Sporen. Girali M. & Mayrhofer H., 1995. Some cortico/ous and lignico/ous species of the genus Rinodina (lichenized Ascomycetes, Unclear taxa Physciaceae) lacking seco11da1y lichen compounds and veget<1- tive propagules in Southem Europe and adjacent regions. Biblioth. Lichenol., 57: 127-160. Aspicilia bricconensis Hue Girali M., Nimis P.L. & Poelt J., 1993. Studien iiber einige Arten Rondon 1969 der F/echtengattung Xanlhoria mit isidiiformen vegetativen A dubious taxon, which might refer to Dirina. Diasporen. J. Hattori Bot. Lab., 74: 271-285. e Aspicilia lactea A. Massal. Grube M. & Haf llner J., I 990. Studien ,m t7echtenbewohnenden Pi/zen der Sammelgattung Didymella (Ascomycetes, Kleinig 1966 - R Dolhideales). Nova Hedwigia, 51: 283-360. Clauzade & Roux 1985 list this taxon under Hafellner J. & Poelt J., 1979. Die Arten der Gattung Caloplaca mit Lecania erysibe (Ach.) Mudd. plurilocularen Sporen (Meroplacis, Triophthalmidium, Xanthocarpia). J. Hattori Bot. Lab., 46: 1-41. Aspicilia reticulata Kremp. Hanko B., 1983. Die Chemotypen der F/echtengattung Pertusaria Steiner 1916 (var. contortoides); Szatala 1943b in Europa. Biblioth. Lichenol., 19: 1-297, Figs 1-59, maps 1-8. (+ ammotropha (Hue) Szatala) - RII, I, L Helms G., Friedl Th., Rambold G. & Mayrhofer H., 2001. Jdentiflc!ltion of photobionts from the lichen family Aspicilia r. ammotropha is a synonym of Physciaceae using a/gal-specific ITS rDNA sequencing. Aspicilia intermutans (Nyl.) Arnold Lichenologist, 32: 73-86.

57 M. GRUBE, L. LINDBLOM, H. MAYRHOFER

Hertel H., 1967. Revision einiger ca/ciphiler Formenkreise der Mayrhofer H., Scheiclegger Ch. & Sheard J.W., 1992. On the taxo­ F/echtengattung Leeidea. Beih. Nova Hedwigia, 24: 1-155. nomy offlve s,1xicolous species of t/Je genus Rinodina (liche­ Hertel H., I 970a. Parasitische lichenisierte Arten der nized Ascomycetes). Nord. J. Bot., 12: 451-459. Sammelgattung Lecidea in Europa. Herzogia, I: 405-438. Mayrhofer M., 1988. Studien iiber die saxicolen Arten der Hertel H., 1970b. Beitriige zur Kenntnis der F/echtenfamilie F/echtengattung Lecania in Europa II. Lec,1nia s. str. Biblioth. Lecideaceae HL Herzogia, 2: 37-62. Lichenol., 28: I- I 33. Hertel H., 1971. Beitriige zur Kenntnis der F/echtenfamilie Moberg R., 1994. Lichenes Se/ecci Exsiccati Upsalienses. Fase. 5 Lecideaceae IV. Herzogia, 2: 231-261. & 6 (Nos 101-150). Thunbergia, 20: 1-17. Hertel H., 1983. Leeideaceae exsiceatae fase. 5 (no. 81-100). Navarro-Rosinés P. & Hafellner J., 1996. Lichenostigma elongata MUnchen. spec. nov. (Dothideales), a lichenicolous Ascomycete on Hertel H., 1992. Lecideaceae exsiccatae fase. I 3 (no. 241-260). Lobothallia and Aspieilia species. Mycotaxon, 57: 211-225. Arnoldia, 2: 1-12. Nimis P.L., 1993. The lichens of Italy. An annotateci catalogue. Hertel H., I 995. Schliissel f'iir die Arten der Flechtenfamilie Museo Regionale di Scienze Naturali, Monografie, XII. Lecideaceae in Europ,1. Biblioth. Lichenol., 58: 137-180. Torino. 897 pp. Hertel H. & Rambold G., 1990. Zur Kenntnis der Familie Nordin A., 2001. Buellia species with pluriseptate spores and the Rimulariaceae (Lecanorales). Biblioth. Lichenol., 38: 145- Physciaeeae (Lecanorales, Aseomycotina). Symb. Bot. Upsal., 189. 33: 1-117. forgensen P.M., 1978. The lichen family Pannariaceae in Europe. Obermayer W., 1997. Lichenes Graecensis fase. 5 (Nos 81-100). Opera Bot., 45: 1-123. Fritschiana, 11: 1-6. Kilias H., 1981. Revision gesteinsbewohnender Sippen der Obermayer W., 1999. 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Nov. Regni Yeg. Beihefte, 69: Poelt J. & KrUger U., 1970. Die Verbreitungsverhiiltnisse der 1-250. F/echtengattung Squamarina in Europa. Fedcles Repertorium, Leuckert Ch., Poelt J., Schultz I. & Schwarz B., I 975. 8 I: 187-201. Chemotaxonomische und stammesgeschichtliche Differenzie­ Poelt J. & Mayrhofer H., 1979. Studien iiber Ascosporen-Typen rung des Formenkreises van Pannelia prolixa in Europa der Flechtengattung Rinodina. Beih. zur Syclowia, 8: 312-331. (Lichenes, Parmeliaeeae). Decheniana, 127: 1-36. Rasanen V., 1944. Griechische F/echten gesammelt van Dr. phil. Leuckert Ch. & Poelt J., 1989. Studien iiber die Lecanora rupicola C. Regel in den Jahren /931-1939. Nova Hedwigia, 81: 229- Grnppe in Europa (Lecanoraceae). Nova Hedwigia, 49: 121- 233. 167. Rondon Y., 1969. Contribution à l'étude des lichens de /'ile de Leuckert Ch., Guse K. & Poelt J., 1969. Zur Chemotaxonomie der Créte. Rev. Fae. Ciencias Lisboa, 2. ser. C, 16: 105-117. Pertusaria hymenea-Gruppe. 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Die F/echtengattungen Rinoclina und Pertusariaeeae (lichenized Ascomyeotina). Acta Bot. Fenn., Rinodinella in der a/ten Welt. J. Hattori Bot. Lab., 55: 327- 150: 153-160. 493. Schneider G., I 979. Die F/echtengattung Psora sensu Mayrhofer H., 1987. Monogrnphieder F/echtengattungThelenella. Zahlbruckner. Versuch einer Gliedenmg. Biblioth. Lichenol., Biblioth. Lichenol., 26: 1-106. 13: 1-291 + 84 Abb. Mayrhofer H. & Leuckert Ch., 1985. Beitriige zur Chemie der Steiner J., 1916. F/echten, van Dr. Ginzberger auf Kreta gesam­ F/echtengattung Rinodina (Ach.) Gray III. Herzogia, 7: I 17- melt. bsterr. Bot. Z., 66: 376-386. 129. Steiner J., 1919. Beitriige zur Kenntnis der Flora Griechenlands. Mayrhofer H. & Poelt J., I 978. Rinoclinella - eine neue Gattung Bearbeitung der anliiB/ich der zweiten Wiener 191 I der Flechtenfamilie Physciaceae. Hoppea, Denkschr. Regensb. Universitiitsreise im Aprii in Griechenland gesammelten Bot. Ges., 37: 89-105. Pflanzen. C. Lichenes. Verh. Zool.-Bot. Ges. Wien, 69: 52- Mayrhofer H. & Poelt J., 1979. Die saxicolen Arten der 101. F/echtengattung Rinodina in Europa. Biblioth. Lichenol., 12: Sundin R., 1999. Phylogenetic and U1xonomic studies within 1-186. Arthonia Ach. (Ascomycetes, Arthoniales). Botaniska Mayrhofer H. & Poelt J., 1985. Die F/echtengattung Microglaena lnstitutionen, Stockholms universitet, 85 pp. + 23 unnumberecl sensu Zahlbruckner in Europa. Herzogia, 7: 13-79. pp. Mayrhofer H., Scheidegger Ch. & Shearcl J.W., 1990. Rinodina Szatala b.,.1943a. Lic/Jenes. In K. H. Rechinger, Florn Aegaea. lecanorina and R. luridata, two c/ose/y related species on cal­ Flora der Jnseln und Halbinseln des Agiiischen Meeres. Akad. ciferous rocks. Biblioth. Lichenol., 38: 335-356. Wiss. Wien, Math.-Naturwiss. Kl., Denkschr., 105/1: 16-58.

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Szatala b., 1943b. Lichenes. In K. H. Rechinger, Neue Beitriige Vezda A., 1969. Lichenes selecti exsiccatae, fase. 3. Brno. wr Flora van Kreta. Akad. Wiss. Wien, Math.-Naturwiss. KI., Vezda A., 1985. Lichenes selecti exsiccatae, fase. 81. Brno. Denkschr., 105/2: 27-47. Vezda A., 1995. Lichenes rariores exsiccatae, fase. 17. Brno. Timdal E., 1984. The delimitation of Psora (Lecideaceae) and refa­ Vitikainen O., 1994. Taxonomic revision of Peltigera (lichenized ted with notes on some species. genera, Nordic J. Bot., 4: 525- Ascomycotina) in Europe. Acta Bot. Fenn., 152: 1-96. 540. Vouaux M., 1914. Synopsis des champignons parasites de Iichens. Timdal E., 199 I. A monograph of the genus Ton inia (Lecideaceae, Ascomycetes). Opera Bot., I IO: 1-137. Bull. Soc. Mycol. France, 30: 1-64, 282-329. Torrente P. & Egea J.M., 1989. La familia Opegraphaceae en e/ Westberg M. & Karnefelt l., 1998. The genus Fulgensia A. Massa/. area Mediterranea de la Peninsula Iberica y Norte de Africa. & De Not., a diverse group in the Teloschistaceae. Biblioth. Lichenol., 32: 1-282. Lichenologist, 30: 5 I5-532. Triebel D., 1989. Lecideico/e Ascomyceten. Biblioth. Lichenol., Wunder H., 1974. Schwarzfriichtige, saxico/e Sippen der Gattung 35: 1-278. Caloplaca (Lichenes, Teloschistaceae) in Mitteleuropa, dem Trinkaus U. & Mayrhofer H., 2000. Revision der Buellia epigaea - Mittelmee1gebiet und Vorderasien. Bibl. Lichenol. 3: 1-186. Gruppe (lichenisierte Ascomyceten, Physciaceae) J. Die Arten Tafeln 1-9. der Nordhemisphiire. Nova Hedwigia, 74: 271-314. Zahlbruckner A., 1904. Schedae ad "Kryptogamas exsiccatas". Verseghy K., 1962. Die Gattung Ochrolechia. Beih. Nova Ann. K.K. Naturhist. Hofmus. 19: 379-427. Hedwigia, I: 1-146 + 12 tables. Beitrag zur Flechtenf1ora Kret11s. Vezda A., 1965. Flechtensystematische Swdien I. Die Gattung Zahlbruckner A., 1906. Petractis Fr. Preslia, 37: 127-143. Sitzungsber. Kaiser!. Akad. Wiss. Wien, Math.-Nat. KI., l 15: Vezda A., 1968. Taxonomische Revision der Gattung Thelopsis 503-523. Nyl. (Lichenisierte Fungi). Folia Geobot. Phytotax., 3: 363- Zehetleilner G., 1978. Uber einige parasitische Arten der 406. Flechtengattung Verrucaria. Nova Hedwigia, 29: 683-734.

Received October 4, 2001 Accepted December 5, 200 I

59

Studia Geobotanica. Voi. 20: 61-66 (2001)

STUDIA LICHENOLOGICA IN ITALIA CENTRALE. III. SPECIE NUOVE PER L'UMBRIA

Valerio GENOVESI', Domenico PUNTILLO 2 e Sonia RAVERA'

'Dipartimento di Biologia Vegetale, Università di Roma "La Sapienza", Piazzale Aldo Moro 5, 1-00185 Roma. ' Orto Botanico, Università della Calabria, 1-87030, Arcavacata di Rende (CS).

Keywords: Centrai ltaly, Flora, Lichens, Umbria.

Abstract: STUDIA L!CHENOLOGICA IN CENTRAL ITALY. III. SPECIES NEW rn UMBRIA. 31 infrageneric taxa are repor­ ted as new to the lichen flora of Umbria (Centrai Italy). Notes conceming their ecology and distribution are given. The new combination Phaeoca/icium mildeanum (Hepp) Puntillo is proposed.

Introduzione Per contribuire alle conoscenze lichenologiche Il paesaggio vegetale delle fasce medio-alte col­ dell'Italia centrale, ancora poco esplorata, gli autori linari è rappresentato da boschi a sclerofillesempre­ stanno effettuando da alcuni anni ricerche di tipo verdi e dai querceti submediterranei che, sui versan­ floristico e fitogeografico nell'Umbria centro-meri­ ti più favorevoli, si spingono fino a c. 1.100 m dionale. Questo territorio, nonostante la posizione s.l.m .. La fascia montana è ricoperta dai boschi di baricentrica rispetto al resto d'Italia, ha condiviso il faggio, monospecifici o misti ad Acer sp. pi. e destino delle regioni adriatiche, poco studiate per­ Sorbus aria, talvolta con potenzialità per Abies ché lontane dai principali istituti botanici (Nimis & alba. Nei fondovalle sono frequenti i consorzi Tretiach, 1999). Nella prima stesura della checklist mesofili a Carpinus betulus e C01ylus avellana e le dei licheni d'Italia (Nimis, 1993) venivano attribui­ boscaglie ripariali a ontani Alnus glutinosa e Salix te all'Umbria solamente 47 segnalazioni, 39 delle sp. pi., miste a pioppi ibridi e robinie. Di notevole quali riferite al Duomo d'Orvieto (Nimis & Monte, interesse lichenologico sono i vecchi impianti di 1988). Una sistematica esplorazione del territorio castagno la cui distribuzione, date le caratteristiche (Ravera, 1998; 1999; 2000a; 2000b; Nimis & del substrato e la parassitosi che li ha colpiti, è Tretiach, 1999; Panfili, 2000a, 2000b; Genovesi & molto ridotta. Ravera, 2001) ha in seguito portato a 415 il numero di taxa noti. Dati e metodi In questo lavoro si segnalano 31 nuove specie per la regione, con qualche commento di interesse Per la detem1inazione delle specie sono stati lichenologico, ecologico, fitogeografico su ogni consultati Clauzade & Roux ( I 985), Fryday & entità. Coppins (1997), Puntillo (1989), Purvis et al. (1992), Timdal (1991), Vitikainen (1994), Wirth Area di studio (1995). La nomenclatura delle specie segue Nirnis L'Umbria è una regione in prevalenza collinare e (2000), quella delle fanerogame Pignatti (1982), montuosa: il clima varia dalla continentalità delle l'abbreviazione degli autori Brurnmitt & Powell zone altomontane alle condizioni miti, di tipo medi­ (1992). terraneo, dei settori collinari (Orsomando et al., Le valutazioni di rarità, rischio di estinzione, la 1999). distribuzione fitogeografica e le caratteristiche eco­ Le litologie affioranti che interessano le località logiche sono desunte da Nirnis (1993, 2000); l'ap­ di raccolta (v. oltre), sono riunite nel complesso car­ partenenza sintassonomica da Wirth (1995). bonatico formato da rocce di origine calcarea che Di ogni specie sono indicati: luogo del ritrova­ attualmente costituiscono l'ossatura di tutti i rilievi mento, tipo di substrato, altitudine, data, e l'iniziale umbri (C.R.I.D.E.A., 1999). del cognome del raccoglitore (V. Genovesi = G, G.

61 V. GENOVESI et al.

Fig. I. - Mappa schematica dell'Umbria con l'indicazione delle località in cui sono stati raccolti i licheni, numerate come nel testo Survey area with geographic distribution ofsampling stations, numbered as in the text.

Massari = M, D. Puntillo = P, S. Ravera = R). Seslerieti con rocce affioranti, calcare bianco e bian­ I campioni sono conservati nell'Erbario del co avorio, 1.400-1.600 m s.1.m., 2/5/99, leg. GMR. Dipartimento di Biologia Vegetale dell'Università 3)* M.te Eremita, lungo il sentiero da Gavelli. In di Roma "La Sapienza" (RO), presso l'Orto faggeta, calcari grigiastri e plumbei o nocciola, Botanico. 1.120 m s.l.m., 1/5/99, leg. GMR. 4) M.te Galenne, Le località di raccolta (Fig. 1) sono indicate con dintorni del Casale dei Cappuccini, in ceITeta, rocce il nome del rilievo o del centro abitato più vicino. affioranti, calcare bianco e bianco avorio, compatto L'asterisco (*) evidenzia i biotopi di rilevante inte­ e ben stratificatocon lenti o strati di selce, ca. 1.000 resse vegetazionale meritevoli di conservazione, m s.l.m., 28/4/00, leg. GMR. 5) M.te Serano, marne segnalati dalla Società Botariica Italiana (1971, mandorlate rosse e verdi, alternate a calcari rossi, 1979). 1.100-1.200 111 s.l.m., 8/4/00, leg. G. 6)* M.ti Provincia di Perugia: 1) Montebibico, a pochi Martani, Parco Regionale. Monte Martano e Monte chilometri dal centro abitato. Vecchio castagneto da Castro, margini di faggeta. Scaglia rossa e calcare frutto a contatto con un bosco a Ca,pinus betulus e bianco e bianco avorio, compatto e ben stratificato Corylus avellana, 860 m s.l.m., 2/10/00, leg. con lenti o strati di selce, 800-930 111 s.l.m., 1/5/00, GMPR. 2)* M.te Coscemo, sopra Forca della Spina. leg. GMR. 7)* Pani.cale. Bosco di roverella di

62 Studia Geobot. 20 (2001) - Licheni nuovi per l'Umbria -

Panicarola, ca. 300 m s.l.m., 28/10/98, leg. R. 8)* Calicium adspersum Pers. Sassovivo. Nei pressi della sorgente minerale, Molto raro, proprio di conifere e vecchi alberi marne rosse e verdastre in alternanza con calcari di decidui con scorza acida e profondamente fessu­ colore verde chiaro, ca. 600 m s.1.m., 27/4/00, leg. rata, in nicchie protette dalla pioggia. L'unica GMR. segnalazione per l'Italia centrale riguarda la Provincia di Terni: 9) Ferentillo, lungo il Fosso Toscana (Jatta, 1909-11). Caratteristica del d' Ancaiano, su Fraxinus ornus, 325 m s.l.m., Chrysotrichetum candelaris Mattick 1937 ex 6/10/00, leg. GMPR. 10) M.te Petano, pressi di Barkm. 1958. (1), su legno di Castanea sativa. Salto del Fosso e Dogana. Pareti fortemente inclina­ Caloplaca cerina var. chloroleuca (Sm.) Th. Fr. te o verticali. Scaglia rossa, ca. 900 m s.1.m., 3- Tipicamente sopra la linea degli alberi, nell'area di studio si trova su substrato calcareo nell' am­ 4/10/00, leg. GMPR. 11)* M.te Solenne, pressi di bito di seslerieti a Sesleria nitida e S. apennina. Loreno, su affioramenti di calcari marnosi rossi e (2, 12), su muschio. bianchi con selce grigia in lenti ed arnioni ed inter­ Caloplaca .fiavovirescens (Wulfen) Dalla Torre & calazioni di calcari detritici, 825 m s.l.m., 14/2/00, Sarnth. leg. GMR. 12) M.te La Pelosa, margini di faggeta, Tipica di silicati basici e arenarie, più raramente calcari grigi e grigi scuri, ben stratificati, ca. 1.300 su calcari compatti. Si segnala dal Monte m s.l.m., 5/3/00, leg. G. 13) Melezzole, in castagne­ Galenne, su substrato calcareo. (4, 5). to, scaglia rossa, ca. 700 m s.l.m., 6/10/00, leg. Catapyrenimn cinereum (Pers.) Korb. GMPR. 14) M.te San Pancrazio, radure ai margini Specie rara, localmente comune solo nella fascia di lecceta, rocce affioranti di calcare massiccio, ca. subalpina, al suolo. (2). 1.000 m s.l.m., 15/7/00, leg. G. Catapyrenimn imbricatmn (Nyl.) Clauzade & Cl. Roux Risultati Rara, tipica di roccia e suoli calcarei. Segi1alata da Sommier (1910) e Jatta (1909-11) per la Agonimia tristicula (Nyl.) Zahlbr. Toscana e da Nimis & Tretiach (1993) per la Tipicamente su muschi terricoli, si trova alla Sardegna. (2), al suolo. base di vecchi alberi, anche nel Lobarion. La sua Catapyrenium rufescens (Ach.) Breufl presenza in Italia è apparentemente sottostimata. Specie rara in Italia, xerofitica, calcicola. (14), al (10), su muschio. suolo. Arthonia cinnabarina (DC.) Wallr. Chaenothecaferruginea (Sm.) Migula Poco comune nel nostro Paese, in via di estin­ Specie molto rara, propria di scorza e legno di zione. Si conferma l'affinità per la scorza liscia conifere e vecchi alberi decidui, in situazioni di Fraxinus, all'interno di boschi umidi. umide e ben illuminate ma protette dalla pioggia. Caratteristica dell' Opegraphetum rufescens Caratteristica del Chaenothecetum ferrugineae Almb. 1948 ex Klem. 1955. (9). Barkm. 1958, è tra le specie segnalate nella Lista Arthonia pruinata (Pers.) A. L. Sm. Rossa dei licheni d'Italia (Nimis, 2000). (1), su Poco comune e in declino, è rish·etta ad ambien­ legno di Castanea sativa. ti umidi ed ha affinità tirrenica. Tipica di vecchi Chaenotheca Jitrfuracea (L.) Tibell alberi isolati, in nicchie protette dalla pioggia. Igrofitica, tipica di radici affioranti o nicchie protette dalla pioggia in situazioni umide. È stata Caratterizza l'Arthonietum pruinatae Almb. raccolta su radice di Corylus avellana nell'ambi­ 1948. (10), su radice. to del Carpino betuli-Coryletum avellanae Aspicilia parasitica de Lesd. Ballelli, Biondi & Pedrotti 1980. Caratterizza il Poco comune in Italia. Xerofitica, cresce preva­ Chaenothecetumfiu:furaceae Kalb. 1969. (1). lentemente su licheni crostosi silicicoli che colo­ Chaenotheca trichialis (Ach.) Th.Fr. nizzano superfici esposte e fo1temente inclinate. Poco comune, acidofila, igrofitica, più frequente (6), parassita su Aspicilia calcarea (L.) Mudd. sulle Alpi. (1), su legno di Castanea sativa. Bacidia bagliettoana (A. Massa!. & De Not.) Jatta Fulgensiafulgens (Sw.) Elenkin È localmente comune nelle regioni alpina, subal­ Specie epigea, tipica di suolo calcareo e praterie pina e sulle montagne mediterranee; si trova tipi­ aride, è stata raccolta al limite di una cerreta. camente su muschio, in fessure di roccia calca­ Caratteristica del Toninio-psoretum decipientis rea, dove è stata raccolta. (11, I 2). Stodieck 1937. (10).

63 V. GENOVESI et al.

Lecanora epibryon (Ach.) Ach. Peltigera polydactyla (Neck.) Hoffm. Specie epibriofitica a distribuzione circumborea- Localmente comune nella regione montana, rac­ le (-polare) artico-alpina, lungo l'Appennino colta su suolo e muschio. (3). nota solo per le vette abruzzesi più elevate Phaeocalicium mildeanum (Hepp) Puntillo comb. (Grillo & Romano, 1987; Nimis & Tretiach, nov. hoc loco 1999). Nell'area di studio è stata raccolta sul Basionimo: Calicium mildeanum Hepp apud versante del rilievo dove è segnalata la presenza Rabenh., Lichen. Eur., nr. 718, 1864. di Vaccinium myrtillus (Francalancia & Sinonimi: Stenocybe mildeana (Hepp) Jatta in Orsomando, 1981). (2). Sylloge Lich. Ital., 1900: 478; Calicium ornico­ Lecanora sambuci (Pers.) Nyl. lum Stein. apud Zahlbr. in Ann. Naturhist. Poco comune in Italia, generalmente su sambu­ Hofmus. Wien 25 (1911): 239; Mycocalicium co (Sambucus nigra) e pioppo (Populus sp.), non ornicolum (Steiner) Nadv.; Mycocalicium mil­ si osserva in ambienti fortemente antropizzati. deanum (Hepp) Nadv. Caratteristica del Lecanoretum sambuci Wirth La specie è da attribuire al genere Phaeocalicium 1980. (10, 7), su scorza di Sambucus nigra. A. Schmidt poiché possiede spore ellissoidali Lepraria nivalis J.R. Laundon 1 (3) settate e non fusiformi plurisettate, come Lichene tipico di calcari e dolomie, su superfici tipico del genere Stenocybe (Nyl.) Korb. Inoltre è molto inclinate e protette dalle precipitazioni molto vicino a Phaeocalicium populneum dirette. I campioni sono stati analizzati mediante (Brond. ex Duby) A. Schmidt da cui differisce cromatografia su lastra sottile presso per le spore più grandi (13-29 x 7-9 µm). l'Università di Trieste (supervisione: Prof. C. Estremamente rara e in declino, è nota esclusiva­ Leukert, Berlin), rilevando la presenza di acido mente per il Trentino-Alto Adige (Amold, 1869; Dalla To1Te & Samthein, 1905) e per le Marche protocetrarico. ( l O, 8). (Nimis & Tretiach, 1999) su Fraxinus. Si confer­ Leptogium gelatinosum (With.) J.R. Laundon ma l'affinità di questa specie per Fraxinus ornus Più acidofilo dell'affine leptogium lichenoides e per gli ambienti umidi. È tra le specie segnala­ (L.) Zahlbr. (6), su suolo e muschi. te nella Lista Rossa dei licheni d'Italia (Nimis, Megaspora verrucosa (Ach.) Hafellner & V. Wirth 2000). (10). var. verrucosa Rhizocarpon obscuratum (Ach.) A. Massa!. Epibriofitica, a distribuzione circumboreale­ Specie silicicola e pioniera, morfologicamente e montana (-alpina), rara lungo gli Appennini. chimicamente variabile. (13). Nella stessa stazione di Lecanora epib,yon. Sarcogyne fallax H. Magn. Caratterizza il Megasporion verrucosae Kalb Rara, è nota in Italia per l'Emilia-Romagna 1970. (2). (Magnusson, 1935-1936), la Toscana (Coppins Microcalicium arenarium (A. Massa!.) Tibell in Nimis, 1993; Tretiach & Nimis, 1994) e Rara, si tratta della prima segnalazione per l'Abruzzo (Nimis & Tretiach, 1999). Tipica di l'Italia centrale. Tipicamente parassita rocce intermedie, quali arenarie e scisti calcarei, Psilolechia lucida (Ach.) M. Choisy o popola­ in boschi decidui aperti. ( 1 O). zioni di Stichococcus e Cystococcaceae ma si Sarcogyne privigna (Ach.) A. Massa!. riscontra anche su radici esposte e su legno in Poco comune in Italia; pioniera. Raccolta su sca­ situazioni umide ed ombrose. Nella stazione è glia rossa. Secondo Nimis & Tretiach (1999), la stata raccolta su radici esposte in una nicchia del var. calcico/a, diffusa nel versante adriatico suolo. Caratteristica dello Psilolechietum luci­ della Penisola e nell'Italia meridionale, ha scar­ dae Schade 1934 ex Klem. 1950, è tra le specie so valore tassonomico. (13). segnalate nella Lista Rossa dei licheni d'Italia Staurothele hymenogonia (Nyl.) Th. Fr. (Nimis, 2000). (1). Poco comune in Italia. Specie pioniera, in Parmelia somloensis Gyelnik Umbria si segnala su rocce calcaree affioranti. Propria di roccia silicea e silicati basici, si segna­ (2, 5). la su suolo calcareo. Caratterizza il Parmelietum Toninia opuntioides (Vili.) Timdal somloensis Klem. 1955. (12). Poco comune in Italia. (6, 10, 11, 12). Peltigera neckeri Mi.ili.Arg. Trapelia coarctata (Sm.) M. Choisy Più comune in faggeta. In ambienti umidi colo­ Rara, silicicola e prevalentemente tirrenica. nizza direttamente il suolo, come nel caso delle Caratterizza il Porpidietum crustulatae Klem. stazioni umbre. (6, 10). 1950. (12, 13), su selce.

64 Studia Geobot. 20 (2001) - Licheni nuovi per l'Umbria -

Discussione Brummitt R.K. & Powell C.E. (ecls.), I 992. Authors of plant names. Royal Botanic Garden, Kew, 732 pp. Il continuo aumento in questi ultimi anni del Clauzade G. & Roux C., 1985. Likenoj de Okcidenta Europa. numero delle specie riportate per l'Umbria sembra Ilustlita detenninlibro. Bull. Soc. Bot. Centre-Ouest n. s., nr. confermare la presenza di condizioni adatte per lo spéc. 7. 893 pp. C.R.I.D.. E.A., 1999. Relazione sullo stato dell'ambiente in Umbria. sviluppo di una flora lichenica ricca e diversificata. (www.regione.umbria.it/ criclea/relazione/clefault. htm). La prevalenza delle montagne, la bassa densità della Dalla Torre K.W. & Sarnthein L. (eds.), 1905. Die Pilze (Fungi) popolazione, l'assenza di importanti insediamenti van Tiro!, Voralberg und Liec/�testein (bearbeitet van Dr. P. industriali ( con qualche eccezione, come la Cònca Magnus). Wagner. lnnsbruck. 716 pp. ternana) hanno limitato la concentrazione e la diffu­ Francalancia C. & Orsomanclo E., I 981. Carta della vegetazione sione degli agenti inquinanti (Marchetti et al., 1999; del Foglio Spoleto. Ist. Bot. Univ. Camerino. C.N.R. Collana del Programma Finalizzato "Promozione della Qualità Bargagli et al., 2000), con evidente vantaggio per la del!'Ambiente" AQ/1/184. 25 pp. conservazione della diversità lichenica. FryclayA. & Coppins B., 1997. Keys to sterile, crustose saxicolous Le nuove segnalazioni offrono lo spunto per and terricolous lichens occurring in the British Isles. alcune considerazioni. Per quanto riguarda i licheni Lichenologist, 29: 301-332. epifiti, Calicium adspersum, Chaenotheca ferrugi­ Genovesi V. & Ravera S., 2001. Studia lichenologica in Italia Centrale. Il. Specie epilitiche nuove per l'Umbria. Webbia., nea, Ch. trichialis, Microcalicium arenarium fanno 56(2) in stampa. parte del corteggio floristico dei castagneti. La scor­ Grillo M. & Romano E., 1987. I Licheni del Parco Nazionale za subacida e profondamente fessurata degli esem­ d'Abruzzo. Primo contributo floristico. Arch. Bot. !tal., 63: plari più vetusti di Castanea è un habitat ideale per 136-152. le Caliciales: si formano nicchie protette dall'azio­ Gruppo di lavoro per la Conservazione della Natura della Società ne diretta delle precipitazioni, in ambienti con umi­ Botanica Italiana, 1971 e 1979. Censimento dei biotopi di rile­ vante interesse vegetazionale meritevoli di conservazione in dità atmosferica piuttosto elevata e luce diffusa. La Italia. Camerino, Tip. Succ. Savini-Mercri, 2 voli. presenza delle Caliciales è indicatrice di continuità Jatta A., 1909-1911. Flora Italica Ciyptoganrn, p11rs III. Lichenes. ecologica e conferma l'importanza dei castagneti la Tip. Cappelli. Rocca di S. Casciano. 958 pp. cui conservazione, non soltanto per motivi licheno­ Magnusson A.H., 1935-1936. Acarosporaceae und Thelocar­ logici, deve essere fortemente sollecitata. paceae. In: Rabenhorst's Kryptogamenflora 2 Aufl., 9, 5: 1- 318. Un altro punto riguarda i licheni epilitici di sub­ Marchetti E., Anzini L., Fabri S., Giovannini G., Viola V. & strati intermedi ( quali Aspicilia parasi ti ca, Castellani M., 1999. Andamento dei valori dell'inquinamento Caloplaca jlavovirescens, Parmelia somloensis, atmosferico riscontrati nella Conca Ternana nel/'<111110 /995. ecc.) che in Umbria si trovano con una certa fre­ Biologi Italiani, 6: 33-40. quenza anche su calcari duri e compatti. Questo Nimis P.L., 1993. The Lichens of Italy. An annotated catalogue. Museo Regionale Scienze Naturali Torino, Monografie, XII. fenomeno, già segnalato per l'Italia adriatica e cer­ 897 pp. tamente legato ali' influenza di fattori climatici Nimis P.L., 2000. Checklist of the Lichens of Italy 2.0. University (Nimis & Tretiach, 1999), deve essere ancora cor­ of Trieste, Dept. of Biology, IN2.0/2 (http://clbios.univ.trie­ rettamente interpretato (Salvadori & Tretiach, ste.it/). 2001). Nimis P.L. & Monte M., 1988. The lichen vegetation on the c11the­ dral of Orvieto (Centrai Italy). Studia Geobot., 8: 77-88. Nimis P.L. & Tretiach M., l 993. A contlibution to Lichen f1oristics Riassunto in Italy. Boli. Museo Reg. Sci. Nat. Torino, 11: 1-45. Nimis P.L. & Tretiach M., 1999. Itinera Adriatica. Lichens from In questo contributo alla conoscenza della flora lichenica dell'Italia the Eastern Part of the Italian peninsula. Studia Geobot., 18: centrale sono riportate 31 specie nuove per l'Umbria e viene pro­ 51-106. posta la nuova combinazione Phaeocalicium mildeanum (Hepp) Orsomando E., Catorci A., Pitzalis M. & Raponi M., 1999. Carta Puntillo. Alla luce delle nuove segnalazioni, alcune delle quali Fitoclimatica dell'Umbria. Regione clell'Urnbria, Università cli d'interesse ecologico e fitogeografico, si rileva: 1) l'importanza Camerino e Università cliPerugia. della salvaguardia dei castagni pluricentenari, che costituiscono il Panfili, 2000a. Contributo alla conoscenza dei licheni dell'Italia substrato d'elezione per molte specie di Caliciales; 2) il rinveni­ centrale: specie nuove o interessanti nell'area della Provincia mento su substrati calcarei di alcune specie silicicole, analogamen­ di Perugia. Provincia di Perugia, Perugia, 5 pp. te a quanto osservato in altri distretti appenninici. Panfili, 2000b. I licheni del parco scientifico didattico Isola Po/vese. Provincia di Perugia, Perugia, IO! pp. Bibliografia Pignatti S., 1982. Flora d'Italia. Edagricole. Bologna, 3 voli. Puntillo D., 1989. Chiavi analitiche delle Caliciales italiane Arnold F., 1869. Lichenologische Ausfiiige in Tirai. IV. Der (Licheni). Webbia, 43: 145-168. Schlern. Yerh. zool.-bot. Ges. Wien, 1869: 606-656. Purvis O.W., Coppins B.J., Hawksworth D.L., James P.W. & Bargagli R., Massari G. & Ravera S., 2000. Biomonitornggio di Moore D.M. (ecls.), 1992. The Lichen Flora of Gre11t Britain elementi in tracce con il lichene Xanthoria parietina in and Ireland. Nlltural History Museum Publications in associa­ Umbria. Biologi Italiani, 8: 42-54. tion with the British Lichen Society, Lonclon. 71 O pp.

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Ravera S., 1998. Contributo alla conoscenza dei licheni d'Italia: la Medite1ranean region. Biblioth. Lichenol., in stampa. florula epifitica dello Spoletino (Regione Umbria). Biologi Sommier S., 1910. La flora dell'isola di Pianosa. N. Giornale Bot. Italiani, 6: 17-24. !tal. 17: 123-164. Ravera S., 1999. Licheni nuovi e interessanti per l'Umbria. Timdal E., 1991. A monograph of the genus Toninia (Lecideacee, Allionia, 36: 101-103. Ascomycetes). Opera Bot., 110: J-137. Ravera S., 2000a. Contributo alla conoscenza della flora lichenica Tretiach M. & Nimis P.L., 1994. Una collezione di licheni dalle epifitica dell'Umbria. Studia Geobot., 19: 49-55. Foreste Casentinesi (Camaldoli, Toscana). Not. Soc. Lich. Ravera S., 2000b. Flora e vegetazione lichenica epifitica !tal., 7: 25-32. dell'Umbria. Tesi di Dottorato. Dipartimento di Biologia Yitikainen O., 1994. Taxonomic revision of Peltigera (lichenized Vegetale, Università di Roma "La Sapienza". Ascomycotina) in Europe. Acta Bot. Fennica, 152: 1-96. Salvadori O. & Tretiach M., 2001. Thallus-substratum relationshi­ Wirth V., 1995. Die Flechten Baden-Wiirttembergs, Teil I & 2. ps of silicico/ous lichens occurring on carbona tic rocks of the Ulmer & Co., Stuttgart. 1006 pp.

Received February 28, 2001 Accepted May 21, 2001

66 Studia Geobot. 20 (200 I)

Direttore responsabile: Dr. Mauro TRETIACH Decreto del Tribunale di Trieste n. 735 del 30.5.1988 Finito di stampare il 28 dicembre 2000 Tipografia Villaggio del Fanciullo - Opicina (Trieste)