(Poaceae) in SW Europe (Iberian Peninsula)

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Anatomical plasticity in species of Deschampsia P. Beauv. (Poaceae) in SW Europe (Iberian Peninsula) a

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Trinidad Ruiz Téllez , Juan Antonio Devesa & Josefa López

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Departamento de Biología y Producción de los Vegetales, Botánica, Facultad de Ciencias, Universidad de Extremadura, Avda. Elvas s/n, E-06071, Badajoz Published online: 27 Apr 2013.

To cite this article: Trinidad Ruiz Téllez , Juan Antonio Devesa & Josefa López (1998) Anatomical plasticity in species of Deschampsia P. Beauv. (Poaceae) in SW Europe (Iberian Peninsula), Acta Botanica Gallica, 145:4, 281-305, DOI: 10.1080/12538078.1998.10516308 To link to this article: http://dx.doi.org/10.1080/12538078.1998.10516308

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Acta bot. Gallica, 1998, 145 (4). 281-305.

Anatomical plasticity in species of Deschampsia P. Beauv. (Poaceae) in SW Europe (Iberian Peninsula) by Trinidad Ruiz Tellez, Juan Antonio Devesa and Josefa Lopez Departamento de Biologia y Produccion de los Vegetates, Botimica, Facultad de Ciencias, Universidad de Extremadura, Avda. Elvas sin, E-06071 Badajoz

Summary.-

The utility of leaf anatomy in the taxonomy of the genus Deschampsia is discussed. To this end, the inter- and intra-populational variability of this character is analyzed in the taxa of the genus which are represented in the Iberian Peninsula (Deschampsia cespitosa (l.) Beauv. s./., D. setacea (Huds.) Hack. and D. flexuosa (l.) Trin.), as well as in Holcus grandiflorus Boiss. and H. caespitosus Boiss., two Spanish endemic species which some authors include in Deschampsia. A critical analysis of their observed anatomical plasticity is given, evidencing the great intra-individual variability when wild plants are grown under experimental conditions.

Resume.- On discute ici de l'utilite de l'anatomie des feuilles dans Ia taxonomie du genre Deschampsia. On analyse ainsi Ia variabilite de ce caractere au sein de differentes populations, ainsi que dans les taxa du genre representes dans Ia Peninsula lberique (Deschampsia cespitosa (l.) Beauv. s./., D. setacea (Huds.) Hack. et D. flexuosa (l.) Trin.) enfin dans les taxa Holcus grandiflorus Boiss. et H. caespitosus Boiss., endemiques espagnoles parfois places dans le genre Deschampsia. L'analyse critique de Ia plasticite anatomique trouvee met en relief un taux eleve de variabilite intra-individuelle quand on fait croitre les plantes sauvages dans des conditions experimentales de culture.

Key-words : Angiosperms

- Poaceae- Aveneae - Deschampsia - Holcus Homalachne- Homoiachne- anatomy - phenotypic plasticity - adaptation.

I. INTRODUCTION

The study of the anatomical features of Poaceae has traditionally been regarded as of great interest in the taxonomy of the family. Since Avdulov ( 1931) this information, specially that relating to leaf anatomy, has been useful to support the segregation of the five subfamilies which many authors today recognize (Watson et a/., 1985; Watson and Dallwitz, 1988, 1992; Ellis, 1987). Outstanding among the contributions to knowledge of the leaf anatomy of the Poaceae are those of De Wet ( 1958), Prat ( 1932, 1936, 1960), Brown


282

( 1958), Ellis ( 1976, 1979), Renvoize ( 1981, 1982-1987) and, above all, of Metcalfe ( 1960), in their most part fundamentally descriptive. With respect to the genus Deschampsia there are abundant anatomical antecedents, including together with some of the aforementioned works, the contributions of Altenkirch ( 1894 ), Burr and Turner ( 1933 ), Buschmann ( 1950), Davies (1959), Duval-Jouvc ( 1875 ), Grob ( 1896 ), Lewton- Brain ( 1904 ), Martin ( 1955 ), Pcc-Laby ( 1898) and Zemann ( 19061907). In the Iberian Peninsula the species of Dcschampsia have also been studied anatomically (for a review, see Devesa, 1992 ), and the characters of this type have been much used to recognize new taxa (Hackel, 1880; PaLmero, 1956; Vivant, 1978; Cervi and Romo, 1981 ). Nevertheless, leaf anatomy features must be used with caution in the taxonomy of the family since a number of authors have pointed out their remarkable variability and their consequent limited usefulness (vide Connor, 1960; Aiken and Lefkovich, 1984; Dube and Morissct, 1996 ). Currently, of the approximately 40 species making up the genus (Clayton and Renvoize, 1986; Nicora and Rugolo, 1987; Watson and Dallwitz, 1988, 1992), there arc three represented in the Iberian Peninsula with many infraspccific variants (see Table 2 in the Discussion). The knowledge of their taxonomy has been due above all to the contributions of Willkomm ( 1861 a), Henriques ( 1905 ), PaLmero ( 1956 ), Vivant ( 1978 ), Cervi and Romo ( 1981 ), Clarke ( 1980), Romo ( 1986) and Bayer and Lopez ( 1994), in which there has been great unevenness in the taxonomic treatment they have received. There has been a recent contribution from Garcia ct a/. ( 1997) which gives abundant details of the differences between the taxa of Dcsclwmpsia and those of A1·cnc/la and Aristavena, genera that arc included in the former. Notable among these contributions because of their taxonomic treatment of the genus are those of Henriques (loc. cit.), Vivant (loc. cit.), and Cervi and Romo (loc. cit.). These authors base the description of new taxa above all on the anatomical characteristics of the leaves. This character has been called into question since the work of Burduja and Tomas ( 1971) on Dcsclwmpsiafle.ruosa. Of importance is the work of Bayer and Lopez ( 1994 ), who clarify the status of the genus in the Peninsula on the basis of analysis of earlier works and of their own studies. They do not, however, take up any position concerning the possibility of including the cndcmisms of southern Spain described under the genus Holcus (Holcus cacspitosus Boiss. and H. grandiflorus Boiss.) in Desclwmpsia. Many authors include these under Holcus (e.g. Tutin, 1980), others in Deschampsia (see Clayton, 1984 ), and others even segregate them into independent genera (Homalachne and Homoiachnc; Pilger, 1949). The present work approaches the study of intra and inter-populational variability of leaf anatomy characters in several species of the genus Desclwmpsia represented in the Iberian Peninsula (D. .flcxuosa (L.) Trin., D. ce.1pitosa (L.) Bcauv. s.l., and D. setacea (Huds.) Hack.). The objectives were to: (I) typify the existing variability; (2) relate the said variability- if it exists- to habitat characteristics; (3) test the usefulness of the character in the taxonomy of the genus; and (4) verify the anatomical affinity of Hole us grandiflorus and H. cac.1pitosus with the taxa of the genus Dcschampsia.

II. MATERIAL AND METHODS Live plants were collected from natural populations (see Annex 1). Some were dried and archived in the herbarium (UNEX). and others of the same population were transported alive in pots (in a 50% Bures peaVsand mixture) to the


283 experimental garden (University of Extremadura, Campus of Badajoz, Spain) where they were grown under identical conditions of irrigation (200 cc water/week) and environmental temperature (yearly average temperature. T = 16,8°C). In both types of plant, leaves were taken from the basal third for transversal section anatomical study. For the live examples, the said study was repeated after 12 months of cultivation under experimental conditions, in order to detect any possible anatomical variation under these new conditions. Similarly, the inter-populational variability was investigated by studying in the same way several herbarium specimens of each taxon from diverse points of their area of distribution in the Iberian Peninsula (see Annex 1). The methodology used for obtaining leaf sections was basically that used by Lopez and Devesa (1991 ). The dry material was cut by hand with a razor and the resulting sections were softened and decoloured in a 50% lactic acid solution. The sections were stained either with a tincture of safranine or with cotton blue, followed by successive rinses with distilled water and 70% alcohol solution. Finally the sections were mounted in 50% lactic acid.

III. RESULTS AND DESCRIPTIONS

A. Deschampsia cespitosa (L.) Beauv., Agrost. 91, tab. 18, fig. 3 (1812) Aira ccspitosa L., Sp. Pl. 64 (1753) a. subsp. ccspitosa A taxon typical of the cold-temperate and wet regions of both hemispheres which reaches Arctic regions, N and S America (US. Canada, and the Patagonian Andes). Northern Europe and Asia, the Mediterranean region (except the E), New Zealand. and African mountains ( Bor, 1960; Dogan, 1985 ). In the Iberian Peninsula it inhabits the mountainous regions of N Portugal (Tras os Montes) and N Spain (provinces of Orensc. Lugo. Conn1a. Leon, Cantabria. Burgos. Yizcaya. Navarra. Huesca. Lerida and Gerona). living in peat bogs ( OY\'Cocco- Sphagnctca), wet meadows (Molinio- Arrcnathcrctm) and alder or beech thickets (A/no - U/mion, Fagion), between (490-) 900 and 1900 (-2500) m a.s.l. Edaphically indifferent. (2n = 26. Hucsca; 2n = 26+2B. Lerida; Ruiz incd. 2n = 26. 26+ lB. 26+ lB+ l t~ Portugal. Kawano. 1963 ). Lea( ana tonn · Deschampsia cespitosa subsp. ccspitosa presents a highly constant pattern of leaf anatomy over the whole of its area of distribution in the Iberian Peninsula (Table l; Fig. l A). Among its most notable characteristics is the thickness and flatness of its transversal section showing highly developed ribs on the adaxial surface, and above all the particular arrangement of the sclcrcnchyma. This tissue in the wild material always appears arranged in strands towards the apex of the ribs. as well as in strands facing those on the abaxial surface. with the same orientation but at the level of the valleys. This was the pattern observed in all the populations studied from the provinces of Yizcaya. La Corui'ia. Hucsca. Leon. Lerida, and Navarra (sec Annex I). After cultivation in the experimental garden of plants from the provinces of Huesca (UNEX 21434) and Lerida (UNEX 21437 and UNEX 21438). major changes were noticed in the anatomical pattern. Individuals. whose leaves collected in situ showed the general anatomical pattern described above. after one year of cultivation under experimental conditions. produced leaves which in the anatomical study showed substantial changes. above all in the arrangement of the sclcrcnchyma. In particular. the individuals of the population UNEX 21434 (Fig. I A') presented after one year of cultivation leaves in which the sclerenchyma was arranged so as to form a continuous subepidermal band on the abaxial surface. and generally with less breadth and a more conduplicatcd lamina. Also. the cultivated individuals of the Lerida populations UNEX 21437 and UNEX 21438 presented first-year leaves either with strands of larger size or with


284

8'

~c

,!A

~D

• E'

Fig. 1.- Evolution of the anatomy of Oeschampsia under cultivation. Wild material A, B, C, 0 and E. The same material after 12 months of cultivation A', B', C' and E'. Oeschampsia cespitosa subsp. cespitosa (A-A', UNEX 21434, Huesca). 0. cespitosa subsp. subtriflora var. subtriflora (B-B', UNEX 21442, Cuenca). 0. cespitosa subsp. subtriflora var. congesta (C-C', UNEX 21451, Avila). 0. cespitosa subsp. media (0, SEV 6264, Valencia; E-E', UNEX 21452, Lerida). (All sections x 33). Fig.1.- Evolution anatomique des Oeschampsia cultives. Materiel sauvage A, B, C, 0 et E. Le meme materiel apnis 12 mois de culture A', B', C' et E'. Oeschampsia cespitosa subsp. cespitosa (A-A', UNEX 21434, Huesca). 0. cespitosa subsp. subtriflora var. subtriflora (B-B', UNEX 21442, Cuenca). 0. cespitosa subsp. subtriflora var. congesta (C-C', UNEX 21451, Avila). 0. cespitosa subsp. media (0, SEV 6264, Valencia; E-E', UNEX 21452, Lerida). (Tousles schemas x 33).

285

the strands touching each other to form a more or less continuous abaxial subepidermal band.


b. subsp. suhtriflora (Lag.) E. Bayer & G. Lopez, Anales .lard. Bot. Madrid 52 (I): 56 ( 1994) A ira suhtriflora Lag., Varied. Ci. 2 ( 19 ): 39 ( 1805 ); Gen. Sp. Nm·. 3 ( 1816)

a. var. suhtrijlora Endemic to theN of Portugal (Tras os Montes, Minho and Douro litoral), and theN, C and E of Spain (provinces of Alava, Albacete, Avila, Burgos, Cantabria, Castell6n, Ciudad Real, Cuenca, Gerona, Granada, Guadalajara, Huesca, .Jaen, La Conu'ia, Le6n, Lcrida, Logrono, Lugo, Madrid, Murcia, Navarra. Palencia, Salamanca, Segovia, Soria, Tarragona. Tcrucl, Valencia, Yalladolid, Yizcaya, Zamora, Zaragoza). It lives in wet meadows (Molinio-Arrenatheretca), between (360-) 800 and 1800 (-2020) m a.s.l. Preferentially basophile. (2n = 26, Albacete; Ruiz incd). Lea(anatom\' Desclwmpsia suhtriflora var. .1uhtrijlora is a Mediterranean optimum taxon whose leaf anatomy is characterized fundamentally by the presence in the lower leaves of a continuous subepidermal band of sclerenchyma towards the abaxial surface (Table I. Fig. I B). This gives them a notable consistency and rigidity. The morphological pattern is completed also by the presence of sclerenchyma reinforcements beneath the adaxial ribs, which arc observed in the sections as sclcrcnchymatic strands (Fig. I B). Slightly different models were observed in only 3% of the cases studied, as in the plants from Lc6n (LEB 14813 and LEB 26639) and Huesca (SEV 125684) in which the basal leaves showed a markedly thinner sclerenchymatic band, including fragmentation into abaxial subepidermal sclcrcnchymatic strands, a variation presented by plants in a Lconcsc population (SEV 125869 and LEB 13848) which were growing together with others showing the "normal" anatomic pattern. Similarly, it is notable that the model developed by the lower leaves is not found in the upper leaves, in which the subepidermal band of sclcrenchyma is discontinuous. On transplanting individuals of this taxon from their natural habitats to the experimental garden, with stable conditions and optimal irrigation, it was found that the anatomic models became noticeably modified. Thus, leaves originating after one year of cultivation showed marked anatomical differences with respect to those studied directly from individuals under natural conditions. In particular, it was found that the continuous sclerenchymatic band was maintained only in the populations UNEX 21441 and UNEX 21439, while in most cases (populations UNEX 21440, UNEX 21442, UNEX 21445 and UNEX 21449) it was replaced by strands of abaxial sclerenchyma facing the adaxial ribs (Fig. I B'), and the apical strands in the latter may even have disappeared. [3. var. congcsta (Font Quer) Ehr. Bayer & G. Lt'lpcz, Analcs .lard. Bot. Madrid 52 (I): 64 ( 1994) Aim media var. conge.1·ta ["codgcsta"] Font Quer, Bot. Real Soc. Hist. Nat. 25: 266 ( 1925) Endemic in the C of the Iberian Peninsula (provinces of Avila, Ciceres and Salamanca). In high pastures (Nardi on) on si liccous soils, between 1800 and 2300 m a.s.l. ( 2n = 26, A vi Ia, Ruiz incd).


286

Lea( ana tom_1 · The pattern of leaf anatomy that characterizes this taxon is defined by the existence of sclerenchymatic reinforcements in the apical region of the adaxial ribs and in the zones opposite these on the abaxial surface (Table I; Fig. I, C). This is a model which in essence is very similar to that of Desclwmpsia cespitosa, but in a species whose leaves are always narrower, with a conduplicate section and which present fewer and less marked ribs than in that other taxon (Fig. I A). In essence, the model does not change after transplantation to experimental conditions, except for a greater degree of development, in one case that was studied (UNEX 21451; Fig. I, C'). The plants from the province of A vii a, from Puerto de Pen a Negra ( MA 253895 ), differed from this pattern. They presented the sclerenchyma arranged continuously in the abaxial subepidermal region. This is a frequent character in the peninsular species of the genus, such as Deschampsia subtrijlora. c. subsp. media (Gouan) Husn., Gram. 34 ( 1897) A ira media Gouan, Ohs. Bot. 3 ( 1773) Taxon of the W and C of the Mediterranean region (C Europe, W Mediterranean region and Balkans, Asia Minor; Rozhevits, 1934 ). In the Iberian Peninsula it is restricted to the mountains of the N and E of Spain, (provinces of Gerona, Barcelona, Lerida, Tarragona, Navarra, Teruel and Valencia), where it inhabits hygrophilous meadows (Desclwmpsion mediae), between (800-) 1300 and 1600 m a.s.l. Calcicolous. (2n = 26, Barcelona; Garcia eta/., 1997). Lea( ana tom\· Deschampsia cespitosa subsp. media presents an anatomic pattern characterized by a lack of development and small number of ribs (and of vascular bundles), as well as the existence of a continuous subepidermal band towards the abaxial surface (Table I; Fig. I D). This is the model presented by the material from SW France, from the region of Montpellier, where the typus was described. The said anatomical model is repeated in the peninsular populations that were studied from Teruel (VAB 902363) and Valencia (SEV 6264). In these, however, the sclerenchymatic band was slightly thicker (Fig. I D). In contrast, the plants of this subspecies which reach the Central Pyrenees, and which therefore live under cooler and wetter conditions, present a discontinuous sclerenchymatic band ( UNEX 21452; Fig. I E). There is also a notable lack of thickness of the leaves, which are morphologically similar to those of Desclwmpsia setacea, and a deep folding of the lamina, specially in the populations from the warmer areas. In general lines, the anatomical pattern is similar to that of the individuals of Desclwmpsia ce.1pitosa subsp. suhtriflom, except in general for a smaller size. With respect to the response to experimental cultivation, it is noteworthy that in the only population studied from Llivia ( Leridian Cerdai1a UNEX 21452 ), the model observed (Fig. I, E') was very similar to that of the original population.

B. Deschampsia.flexuo.m (L.) Trin., Bull. Sci.Acad. Imp. Sci. Petersb. 1:66 (1836) Airajlexuosa L., Sp. Pl. 65 ( 1753) Taxon with representation in America (US, Chile, Argentina), Eurasia, and NW Africa, with citations known for Japan, New Guinea, and New Zealand since it has been introduced into many parts of the world ( Bor, 1960; Dogan, 1985 ). In the Iberian Peninsula it is restricted to the massifs of the N, C, and S (provinces of A lava, Almeria, Asturias, Avila,


287

Barcelona, Burgos, Caceres, Cantabria, Castellon, Cuenca. Gerona, Granada, Guadalajara, Guipt:1zcoa, Huesca, Leon, Lerida, Logrono, Lugo, Madrid Navarra, Orense, Palencia, Pontevedra, Salamanca, Segovia, Soria, Teruel, Valencia, Vizcaya, Zamora, Zaragoza, and the portuguese Algarve, Alto Alentejo, Beira Alta, Beira Litoral, Estremadura and Minho). There it inhabits wet meadows associated with oak thickets and woodland gaps ( Quercctalia robori-pctraeac), in stands of pine (Pino-Juniperetea), etc., between (55(}-) 900 and 2000 m a.s.l. Preferentially, acidophilous. (2n = 28, Huesca, Ruiz ined. 2n = 56, Portugal, Albers, 1972 and Queiros, 1978, sub D. strictu). Lea( ana to 1m· Deschampsia.flnuosa is an Euro-Siberian optimum taxon, which entered the Iberian Peninsula via the major mountain chains, reaching the southernmost mountainous areas and the NW of Africa in the Rif mountains of Morocco. In the territory studied it presents a high degree of anatomical polymorphism, which affects fundamentally the sclerenchymatic reinforcements, as is the case with other peninsular taxa of this genus (Table I; Fig. 2 ). In general, all of the populations studied present very thin leaves, without adaxial rib differentiation except for that opposite the central vascular bundle, and with transversal sections showing a lamina of almost circular or open U-shaped outline. These characteristics (Table I) clearly distinguish the populations from those of the rest of the taxa of the genus that are represented in the Iberian Peninsula. With respect to the arrangement of the sclerenchyma, three basic geographically delimited patterns of variation (patterns A, B, C) can be recognized (Table 2; Fig. 2). To pattern A belong plants in whose leaves the sclerenchyma is arranged in a thick 38 cells deep continuous subepidermal band towards the abaxial surt~1ce. This confers on the leaves a greater rigidity and adaptation with respect to changes in turgescence and dehydration (Table 2; Fig. 2 A). This model was detected in some populations of N Spain (Cantabria and the Pyrenees), and in many of the centre and south of the Peninsula in both the Iberian and Central Systems, and in the Betica-Rif mountains. It is therefore, a characteristic model of plants, inhabiting areas more Mediterranean in character (e.g. populations LEB 4937, LEB 24229, SEV 104323 and SEV 103776). Another pattern of variation is presented by a great many of the populations studied. in which the individuals present a band ofsclcrenchyma, which is either very thin (from 1-3( -4) cells in thickness) or discontinuous. In many cases, the band fi·agmcnts into strands of greater or lesser extension. This model appears both in the Cantabrian Range and in the Iberian System and the Pyrenees.

Table 2.- Anatomical patterns in Oeschampsia flexuosa. Sizes in ~m. Tableau 2.- Structures anatomiques chez Deschampsia flexuosa. Dimensions en microns.

Thickness Rib Adaxial epidermis Abaxial epidermis Central vascular bundle Mestome cells number

PATTERN A IThick band\

PATTERN B I Intermediate l

PATTERN C IStrands\

(230-) 250-380 (-480) 50-90 ( -120) (4-) 8-14 X 3.9-12 (-14) 15.8-30 X (3.9-) ]8-25

( 160-) 200-450 (-700) 35-80 ( -160) 4-8 X 4-8 ( 12-) 20-25 (-40) X (8-) 25-30 (-40) 40-95 (-120) 14-20 (-24)

220-300 ( -520) 20-80 (-120) 4- I 2 (- 16) X 4- I 2 (- 16) 20-48 X ]6-50

50-80 ( -90) (14-) 17-20(-21)

44-85 15-20 (-28)

288

..

~.


•••• o.l•

• ·~ 00

•

0

0

,.....•••

0 0

•

A

B

c

D

~

~

E'

E

Fig. 2.- Geographical distribution and evolution of anatomical patterns in Deschampsia flexuosa. White pots and A (SEV 111636): pattern A. Black pots and B (LEB 38853): pattern B. Asterisks and C (LEB 9347) pattern C. Modified patterns obtained under cultivation: D' and E'. The same material before cultivation: D and E respectively. (Voucher specimens: D-D', UNEX 21463, Zaragoza, pattern B; E-E', UNEX 21461, Segovia, pattern C). (All sections x 33). Fig. 2.- Distribution geographique et evolution des structures anatomiques dans Deschampsia flexuosa. Points blancs etA (SEV 111636) : structure A. Points noirs et B (LEB 38853) : structure B. Asterisques etC (LEB 9347) : structure C. Structures modifiees obtenues en conditions experimentales D' et E'. Le meme materiel avant culture : D et E. (Materiel temoin: D-D', UNEX 21463, Zaragoza, structure B; E-E', UNEX 21461, Segovia, structure C). (Tous les schemas x 33).


289

The third anatomical model (pattern C; Table 2 and Fig. 2C) is that found in the populations whose individuals present leaves with discontinuous sclcrcnchymatic reinforcements. which appear in the trans vera! section as forming (6-) 8 (-I 0) abaxially localized strands. coinciding with the folds in the lamina and with the apical end of the only existing rib. This model was observed in plants from the high mountain zones of the Pyrenees (e.g. LEB 4934). Cantabrian Range (e.g. LEB 9347). Central System (e.g. SEV I 1585). and the western pcnsinsula with oceanic influence (e.g. Portugal: UNEX I 87 I. UNEX I 7 I 3 and UNEX I 714 ). This model is. also. that mostly presented by plants from Central European populations. Of extraordinary importance in this taxon are the results relative to the reversibility of the anatomical pattern after growing the individuals under experimental conditions. The individuals which presented leaves with a type 8 pattern in situ. either again presented the sclerenchymatic band in the new leaves that grew after one year of cultivation (in three out of the eight cases studied). or the band appeared fragmented into strands of reinforcement. in the transversal section. as occurred in the other five cases (Fig. 2. D and D'). Similar modifications occurred in the case of some of the plants studied with type C anatomical patterns. although now in the contrary sense. since in this case. one observed an increase in the development of the sclerenchyma (Fig. 2. E and E'). Here. the behaviour was similar to that found in Desclwmpsia cespitosa s.l .. in the sense that the plants of the taxa (or the populations) that have their chorological optimum in the MidEuropean ( Euro-Sibcrian) Region develop to a greater degree the sclerenchymatic tissue after cultivation in the experimental garden. C. Desclwmpsia setacea ( H uds.) Hack., Cat. Rais. Gram. Port. 33 ( 1880) Aim setacea Huds .. Fl. Angl. 30 (I 762) Taxon of W and C Europe. from N Spain to S Norway and W Poland. Britain (Clapham ct a! .. I 997). and eastern coasts of South America (Chile: Parodi. I 949: Smythics. I 986). In the Iberian Peninsula it is represented in NW Spain (Galicia. Cantabria). where it inhabits wet heath clearings. on sodden soils (Littorelletm). (2n = 14. Cantabria. Conu1a: Garcia et a/ .. I 997 ). Lea( ana tonn · Desclwmpsia setacm is the only diploid species of the genus. that is present in the Iberian Peninsula. Its most relevant anatomical characteristics (Table I) arc firstly an arrangement of the sclercnchyma reinforcements in strands. as seems to be characteristic of species with Euro-Siberian optimum. and secondly a reduced number of both vascular bundles running the length of the lamina and ribs. which are also very poorly developed. The explanation is that this is the taxon that presents the thinnest leaves. which are setaceous in aspect. It was not possible in this case to study individuals after transplanting conditions. D. Desclwmpsia minor Clayton, Kew Bull. 40 (4): 727 (1984) Holcus cae.1pitosus Boiss .• Bih/. Uni1·. Ge!Uh·e ser. 2. 13:410 ( 1838) Homalachne cespitosa ( Boiss.) Pilg .. Engl. Bot. Jahrh. 74: 227 (I 948) Homoiachne ce.1pitosa ( Boiss.) Pilg .. Engl. Bot. Jahrh. 74: 556 (I 949) Endemic to the peaks of Sierra Nevada (Granada. Spain). inhabiting rock fissures. cliffs. and stony terrain: on schists (Holcion ce.1pitosi, Androsacctalia). (2n = 14: Ki.ipfer. 1969).

290


E. Deschampsia reuteri Pit g., Engl. Bot. Jahrb. 74: 555 ( 1949) Holcus grandijlorus Boiss. in Boiss. & Reut., Pugillus Pl. Nm•. 119 ( 1952) Homolachne grandif! ora ( Boiss. & Reut.) Pilg., Engl. Bot . .Jahrb. 74: 227 ( 1948) Hole us mol/is subsp. reuteri ( Boiss.) Tutin, Bot. J Linn. Soc. 76 (4 ): 363 ( 1978) Endemic to the province of Cadiz (Spain), appearing in shaded thickets within oak woodland (Magnocaricion ). (n = 7: Romero, 1988 ).

Lea( a 1w tom1· In Desclwmpsia minor and D. reuteri neither the section shape of the leaf nor the pattern of leaf anatomy are related to the peninsular taxa studied of the genus Deschampsia, except for the case of Desclwmpsia cespitosa, which also has flat leaves and discontinuous sclerenchyma (Table l ). Unlike Deschampsia ce.1pitosa. however, there is a characteristic presence in Deschampsia reuteri and D. minor of sclerenchy-

Fig. 3.- A, Anatomy of Deschampsia minor (GDA 30660). B, anatomy of D. reuteri (SEV 81512). Fig. 3.- A, anatomie de Deschampsia minor (GDA 30660). B, anatomie de 0. reuteri (SEV 81512).

291

matic columns, connecting the adaxial and abaxial epidermis at the level of vascular bundles (Fig. 3 ), as was pointed out by Paunero and Rivas ( 1968). This anatomic model is frequent, however, in the taxa of Holcus (see Paunero and Rivas, Joe. cit.; Lopez and Devesa, 1992 ).


IV DISCUSSION The genus Deschumpsia in the Iberian Peninsula has been the subject of several taxonomic treatments (see Introduction). There still exists, however, a certain controversy relative to the status of some of them (vide Bayer and Lopez, 1994 ). Similarly, there has been no critical study of the value of the anatomical features in delimiting certain of the recently described taxa (Vivant, 1978; Cervi and Romo, 1981 ), or of the advisability of including under Desclwmpsiu two species of the genus Holcus (Holcus grundiflorus and H. caespitosus), which clearly present features that are close to the genus Deschampsiu (Pilger, 1949).

A. Taxonomic considerations The first treatment of the genus at the peninsular level was due to Willkomm ( 1861 a), who recognized two large groups for the genus on the basis of the features of the awn of the lemma: "Arista recta .whine/usa" (includes Deschampsia ce.1pitosa. D. media (Gouan) Roem. & Schult. and D. refi·acta (Lag.) Roem. & Schult.) and "Arista genicu/ata sa tis Ionge exerta" ( Desclwmpsiaflexuosa stricta Gay in Durieu. D. flexuosa brachyphrl/u Gay in Durieu, D. thuil/ieri Gren. & Gordon and D. haetica (Trin.) Willk. in Willk. & Lange). Later. in 1893, Willkomm gathered new localities of Dcschampsia in his Supplcmentum, and included the D. stricta described by Hackel ( 1880) from the Serra da Sintra (Portugal). a taxon which both authors related to Desclwmpsia jlexuosa and which they segregated above all for its harder. more rigid, and somewhat pointed, leaves. Later. Henriques ( 1905) recognized six taxa for the Portuguese flora. He judged the identity of Dcschampsia stricta to merit no more than the rank of variety within D. jlexuosa ("me parece que mais jus to scria considerar-sc esta planta como variedadc daD. jlexuosa"). In 1956, PaLmero reviewed the genus. recognizing four species for the Iberian Peninsula: Deschampsia cespitosa. D. jlexuosa. D. setacca and D. media. There is special confusion in her concept of Desclwmpsia media, under which she includes plants from Gredos (which she recognizes as fma. congesta Font Quer), that arc very different from the Montpellier typus, and a series of plants from C. N. and E Spain, whose leaves at times present a layer of sclcrenchyma, which is either continuous towards the abaxial surface (the same as in Desclwmpsia cespitosa subsp. media s.s. ). or discontinuous. incorporating two Lagascan binomials -A ira suhtrijlora and A. reji·acta -. with the categories of form and subspecies. respectively. Vivant ( 1978) describes a new subspecies (subsp. hispanica) with the plants identified by PaLmero (loc. cit.) as Deschampsia media, whose presence in Spain he questions. Likewise, he describes as subspecies the plants from Gredos of congested inflorescence ( subsp. gredensis) ignoring the binomial Agrostis gredensis used by Gandoger ( 1905) to designate these plants (vide Castroviejo, 1982). He adscribes A ira rcfi·acta as a subspecies to Deschampsia cespitosa. Recently a contribution from Cervi and Romo ( 1981) has brought to light new combinations and. specially, four new taxa from the genus in the Iberian Peninsula. based fundamentally on the anatomical characteristics of the leaves: Deschampsia media subsp.


292

masclansii, D. hispanica subsp. gallaecica, D. hispanica var. iberica and D. hispanica fma. neilensis. One of the authors (Romo, 1986) later describes the Pyrenees high mountain forms of Deschampsia cespitosa as a new subspecies (pyrenaica) principally on the basis of the size of the individuals, and the more congested nature of the inflorescences, since the anatomical characteristics are wholly coincident, with what is typical for the species. Of great interest is the contribution of Bayer and Lopez ( 1994 ). They adopt an integrative taxonomic approach reuniting under the highly polymorphic Deschampsia cespitosa a good part of the peninsular races that had been described previously, assigning them either to the category of subspecies (subsp. subtriflora, =A ira refracta and probably D. cespitosa subsp. hispanica as well) or variety (var. congesta, =A ira media var. congesta, Agrostis gredensis, Deschampsia cespitosa subsp. gredensis). To a good degree, the criteria that these authors follow is conditioned not only by the great variability that D. cespitosa presents in the rest of its area, with all of the peninsular taxa being possible local variants, but also by the marked variability that has in general been observed in the leaf anatomy, thus calling into question the value of the said character in the taxonomy of the genus. These authors also discard the presence in the Iberian Peninsula of Deschampsia cespitosa subsp.

Table 3.- Taxonomic summary of the genus Deschampsia in the Iberian Peninsula. Taxonomic synonyms described for the Iberian Peninsula are indicated. Those marked • are taxa which delimitations have been mainly based on anatomical characters. Tableau 3.- Resume taxonomique du genre Deschampsia dans Ia Peninsule lberique. On indique aussi les synonymes taxonomiques decrites pour Ia Peninsule. Ceux qui sont marques avec • sont des taxons delimites ssurtout pour leurs caracteres anatomiques.

I. Sect. Deschampsia I. D. cespitosa (L.) Beauv., Agrost. 91, tab. 18. fig. 3 ( 1812) a. subsp. cespitosa b. subsp. subtrijlora (Lag.) Ehr. Bayer & G. Lopez. Anales Jard. Bot. Madrid 52( I): 56 ( 1994) a . var. subtrijlora A ira reji·acta Lag .• Varied. Cienc. 2. 4 ( 19): 39 ( 1805) Deschampsia cespitosa var. angustijiJ/ia Merino. Fl. Gal. 3: 293 ( 1909) D. cespitosa subsp. rejivcta fma. congesta (Font Quer) Rivas Mart., Anal. Ins/. Bot. Cavanilles 21 (I): 296 ( 1963) D. cespitosa subsp. hi.1pw1ica Vivant. Bull. Soc. Bot. France 125 (5-6): 318 (1978)* D. hispanica subsp. hispanica var. hurehana Cervi & Romo. Collect. Bot. 12 ( 4 ): 85 (1981 )* D. hispanica subsp. hi.1panica var. iberica Cervi & Romo, Collect. Bot. 12 (4): 85 (1981)* D. hispanica subsp. gallaecica Cervi & Romo. Collect. Bot. 12 (4 ): 85 ( 1981 )* D. hispanica fma. neilensis Cervi & Romo. Collect. Bot. 12 (4 ): 85 ( 1981 )* 13. var. congesta (Font Quer) Ehr. Bayer & G. Lopez. Anale.1· Jard. Bot. Madrid 52 (I): 64 ( 1994) A ira media var. congesta ("codgesta") Font Quer, Bol. Real Soc. His/. Nat. 25: 266 ( 1925) Agrostis gredensis Gand .• Bull. Soc. Bot. France 52: 460 ( 1905) c. subsp. media (Gouan) Husn .• Gram. 34 ( 1897) Deschampsia elegans Sennen. Bol. Soc. Arag. 15: 253 (1916) D. media subsp. masclansii Cervi & Romo. Collect. Bot. 12 (4 ): 82 ( 1981 )* II. Sect. Avenaria (Rchb.) Holmb .• Bot. Not. 1926: 262 (1926) 2. D. jlexuosa (L.) Trin .• Bull. Sci. A cad. Imp. Sci. Ntersb. I: 66 (1836) A. baetica Trin .• Mem. Acad. St. Petersb. VI Sci. Nat. 2: 17 ( 1836) Deschampsiaflexuosa var. braclnplnHa Gay ex Willk. in Willk. & Lange. Pr()(h: Fl. Hisp. I: 66 (1861) D. stricta Hack .• Catal. Rais. Gram. Port. 18 (1880) D. .flexuosa subsp. iberica Rivas Mart., Trab. Depto. Bot. Fis. Veg. 3: 113 ( 1971) 3. D. setacea (Huds.) Hack .. Cat. Rais. Gram. Port. 33 ( 1880)

293


cespitosa, possibly from not having been able to achieve access to sufficient material of Pyrenees procedence. In summary, Table 3 presents a taxonomic synthesis of the genus Deschampsia in the Iberian Peninsula. Those in which the leaf anatomy data fulfilled a special role in their delimitation are highlighted, particularly the arrangement of the sclerenchymatic reinforcements and the shape of the ribs.

B. Anatomical considerations While the anatomical features are of great importance in the family (see the antecedents in the Introduction), the value of the said character has been questioned in some genera in particular, such as Festuca (Connor, I 960; Aiken and Lefkovich, 1984; Dube and Morisset, 1996) and Deschampsia (Burduja and Tomas, 1971 ). Despite this, the said features are too often used in that sense, even in drawing up dichotomous keys to identification (Auquier, 1973; Kerguelen and Plonka, 1989; Markgraff-Dannenberg, 1980; Stace, 1991, and so on). In relation to Deschampsia there are abundant antecedents concerning leaf anatomy (see Introduction), some carried out with material from the Iberian Peninsula (Paunero, 1956; Vivant, 1978; Cervi and Romo, 1981; Romo, 1986; Lopez and Devesa, 1991 ). None of them, however, approaches the study of the intra- or inter-populational variability. The results of the present study show that the characteristics of the sclerenchyma and its arrangement in the leaf are strongly affected by the environmental conditions, or at least this is so when the individuals are transplanted to standard conditions that are different from those of their places of origin. Particularly noteworthy in this sense are the bidirectional reversions between continuous and discontinuous sclerenchyma. At least in Festuca ovina L., this anatomical feature is known to be strongly influenced by transplantation to new conditions ( Borsos, 1957 ). Thus, in Deschampsia ce~pitosa subsp. cespitosa, which under natural conditions presents the sclerenchyma arranged generally in strands, it may become continuous or subcontinuous towards the abaxial surface after growing the plants in an experimental garden (Fig. I, A and A'). These morpho-anatomical changes of the leaf can only be adscribed to the phenotypical plasticity of the species after variation of the environmental conditions. They basically point to a greater development of the sclerenchyma when these conditions deviate from the optima of humidity and temperature of the Euro-Siberian areas in which this taxon posseses its distribution area. In this case, the new anatomical pattern means that the taxon is coming closer from this point of view to other species with a Mediterranean optimum that exist in the Iberian Peninsula. As in the case of Deschampsia cespitosa subsp. ce.spitosa, in Deschampsia cespitosa subsp. suhtriflora var. suhtriflora the anatomical feature which defines par excellence the taxon (in this case the existence of a continuous band of sclerenchyma towards the abaxial surface) is not genetically fixed, but is a response to the environmental conditions under which the plants are growing. In this case the sclerenchyma, which is generally continuous, may become discontinuous after cultivation (Fig. I, Band 8'). This may be interpreted as a response of this Mediterranean species to the intense irrigation conditions to which the plants are subjected. Here also, then, the variability in the anatomic pattern responds to the phenotypical plasticity of the taxon, thus calling into question the taxonomic value attributed to it by taxonomists who have worked on the group. Also, in the case of Deschampsia ce.spitosa subsp. subtriflora var. congesta, the anatomical model observed in situ hardly changes after cultivation, except in the greater development of the lamina. This


294

phenomenon is perhaps due to the very different conditions of the said plants' natural habitat: the high mountain zones of the Iberian Peninsula. The case of Deschampsia .flexuosa is specially notable. Its anatomical polymorphism had already been noted by Burduja and Tomas ( 1971 ). In the Iberian Peninsula, the taxon presents three patterns of variation in the arrangement of the sclerenchyma under natural conditions (Fig. 2). The sclerenchyma is often discontinuous in high mountain areas with an Euro-Siberian character (Fig. 2, pattern C), while in the plants from the southern Mediterranean mountains it is arranged in a continuous band (Fig. 2, pattern A). Between these two extremes there are intermediate situations (Fig. 2, pattern B), and bidirectional reversal of the character is known after experimental cultivation (sec Fig. 2, D and E). Environmental modifications, therefore, appear in both Deschampsia .flexuosa and D. ce.1pitosa subsp. cespitosa s.l., favouring in both cases the increase or decrease of sclerenchyma. These marked variations in leaf anatomy after transplanting individuals to conditions that are different from natural conditions, are explained as a manifestation of the notable phenotypical plasticity of these species, as already observed Burduja and Tomas ( 1971) and Garcia eta/. ( 1997), and call into question the taxonomic value, that has traditionally been assigned to this character for the species in question, and for the genus in particular. This special response seems to be linked to the variation in environmental conditions, a phenomenon already reported in Festuca by Hackel ( 1882), Kjellqvist ( 1961) and, recently, by Dube and Morisset ( 1996 ). These latter authors found marked changes in the anatomy of Festuca ruhra L. as a function of different environmental conditions: "Le milieu n'est pas un facteur particulierement preponderant dans Ia determination de Ia structure anatomique des feuilles mais certains caractercs y sont plus soumis que d'autres ... Lcs caracteres anatomiques experiment done une variation importante qui n'est surtout pas le seul reflet des conditions ccologiques" (Dube and Morisset, loc.cit.).

C. Supraspecific treatment and the problem of Deschampsia minor and D. reuteri In respect to the supraspecific treatment adopted in the genus, the criteria of Koch ( 1844 ), Rouy ( 1927) and Rozhevits ( 1934) have been followed. Thus, the first of the authors (Koch, loc. cit.) recognized the section Deschampsia in the genus A ira, and included therein the taxa with poorly developed awns ("Aristis paulum tantum inflexa et hasi 1•ix torta") such as Aira cespitosa and A. ll'iheliana Sonder, and the section Avenel/a. He included in this section taxa with markedly exerted awns ("Arista eFidentius infle:w et hasi torta"), such as Airafle:wosa and A. uliginosa Weihe & Rchb. These two sections are recognized for the genus Deschampsia by Rouy ( 1927), but as Campella and Avenaria respectively, the latter using as basonym the section Avenaria originally described by Reichenbach ( 1830) under the genus A ira. The former had already been established by Grise bach ( 1845) taking an alternative name for the genus Deschampsia, that had been proposed by Link in 1827. Rozhevits did the same in 1934, but naming the two sections Campella ("Campelia", =sect. Deschampsia) and Avenaria Rchb. (=sect. Avenella), respectively. Both sections, Desclwmpsia (= sect. Campella) and Avenaria (Rchb.) Holmberg. (= sect. Avenella Koch) are recognized in the present work. Also included in the former is D. setacea, the only diploid species in the territory, and which Albers and Butzin ( 1977) segregated into an independent genus (Aristavena). The said segregation is not supported by either the morpho-anatomical features or even trials with molecular techniques (Garcia eta!., 1997).


295

Finally, in relation to Deschampsia minor and D. reuteri, originally described under Holcus, it should be noted that Willkomm ( 1861 b) had already united them into a group (Fios uterque aristatus) that is well delimited from the rest of the species of this genus (Fios superior aristatus, inferior muticus). This criterion was in essence followed by Bentham and Hooker ( 1883 ), who established the section Homalachne. Since then, some authors have gone still deeper into the said segregation, separating the two off, into an independent genus (Homalachne and Homoiachne, vide Pilger, 1949), or reuniting them under the genus Deschampsia (Clayton, 1984 ). However, at least from an anatomical point of view, the two Iberian taxa have a closer relationship to the genus Holcus than to the genus Deschampsia, and really seem to constitute a group whose character is intermediate between the two genera (Table 4).

Table 4.- Differential characters among Holcus and the sections recognized in Deschampsia. Tableau 4.- Caracteres de differenciation chez Holcus et les sections reconnues de Deschampsia.

Leaves

1/o/cus

1/o/cus

Desclwmpsia

De.\(·/wmpsia

sect. llo/cus

sect. Homahu·hne

sect. Desclwmp.\ia

sect. A\'(:naria

Flat

flat

flat. conduplicated

Comolutc

or convolute Cilumcs

Acuminate

Acuminate

Acute. seldom acuminate

Acute

or acute

Persistent

(D. almfJ/IIjJ/Irea)

Persistent

Persistent

Deciduous

Number of tlm\ crs Lmver tlowcr

llermaphrodite

Hermaphrodite

Hermaphrodite

Hermaphrodite

Upper flower

Male

llermaphrodite

Hcnnaphroditc

Hermaphrodite

Lower lemma

A\\lnless

Av.ned

Awned seldom av.. nless

Straight. geniculate

Geniculate

Straight. helicoid or scarcely

A\\11

or uncinate: subtcrminal:ahsent in

±

Geniculated

lemma

geniculated

>lemma

spikelet; submedian insertion

lemma ± spikelet

>spikelet

subbasal or

subbasal insertion

>

the lo\\-CT lemma

submedian insertion

Girders of sclerenchyma

Present

Present

Absent

296


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297 Kupfer Ph., 1969.- Recherches cytotaxinomique sur Ia !lore des montagnes de Ia peninsula lberique. Bull. Soc. Neuchateloise Sci. Nat., 92, 31-48. Lewton-Brain L., 1904.- On the anatomy of the leaves of British grasses. Trans. Linn. Soc. London, ser. 2, 7, 315-359. Link J.H.F., 1827.- Campella. In: Hortus Regius Botanicus Berolinensis, descriptus. Berolini, 1, 122123. Lopez J. & J .A. Devesa, 1991 .- Contribucion al conocimiento de Ia anatomia foliar de las Aveneae (Poaceae, Pooideae) del centro-oeste de Espana. Anal. Jard. Bot. Madrid, 48, 171-187. Lopez J. & J.A. Devesa, 1992.- Holcus. In: Anatomia foliar y palinologia de las gramineas extremeiias. J.A. Devesa (ed.), publicaciones de Ia Universidad de Extremadura, Monografias Botanicas. Badajoz, 122125. Markgraf-Dannenberg I. (von), 1980.- Festuca. In: Flora Europaea. T.G. Tutin, V.H. Heywood, N.A. Burges et a/., (eds.), Cambridge, 5, 125-153. Martin D.J, 1955.- Features of plant cuticles. An aid to the analysis of the natural diet of grazing animals, with special reference to Scottish hill sheep. Trans. Bot. Soc. Edinburgh, 36, 278-288. Metcalfe C.R., 1960.- Anatomy of the Monocotyledons, I. Gramineae. Oxford, 731p. Nicora E.G. & Z.E. Rugolo, 1987.- Los generos de gramineas de America austral. Argentina, Chile, Uruguay y areas limitrofes de Bolivia, Paraguay y Brasil. Ed. Hemisferio Sur S. A. Buenos Aires, 611p. Parodi L.R., 1949.- Las gramineas sudamericanas del genero Deschampsia. Oarwiniana, 8, 415-475. Paunero E., 1956.- Las Aveneas espaiiolas. I. Anal. /nsf. Bot. Cavanilles, 13, 175-187. Paunero E. & M.A. Rivas, 1968.- Datos sobre Ia anatomia foliar de Holcus gayanus Boiss. y H.setiglumis Boiss. et Reut. Bal. Soc. Argentina Bot., 12, 99-105. Pee-Laby M.E, 1898.- Etude anatomique de Ia feuille des Graminees de Ia France. Ann. Sci. Nat., 8, 227346. Pilger R., 1949.- Additamenta agrostologica. Ill. I. Uber Holcus L. sect. Homalachne Benth. Engl. Bot. Jahrb. Syst., 74, 554-567. Prat H., 1932.- L'epiderme des Graminees. Etude anatomique et systematique. Ann. Sci. Nat. Bot., ser. 10, 14, 117-324. Prat H., 1936.- La systematique des Graminees. Ann. Sci. Nat., 10, 165-258. Prat H., 1960.- Revue d Agrostologie. Vers une classification naturelle des Graminees. Bull. Soc. Bot. France, 107, 32-79. Queiros, M., 1978.- Numeros cromossomicos para a flora portuguesa, 1-15. Bol. Soc. Brot., ser. 2, 52, 69-

77. Reichenbach L., 1830.- Aira. In: Flora Germanica excursoria, Lipsiae, 1, 50.

Renvoize S.A., 1981.- The subfamily Arundinoideae and its position in relation to a general classification of the Gramineae. Kew Bulletin, 36, 85-102. Renvoize S.A., 1982-1987.- A survey of the leaf-blade anatomy in grasses. I. Andropogoneae. Kew Bull., 37, 315-321; II. Arundinelleae, loc. cit., 37, 489-495 (1982); Ill. Garnotieae, loc. cit., 37, 497-500 (1983); IV. Eragrostideae, loc. cit., 37, 469-478 (1983); V. The bamboo allies, loc. cit., 40, 509-535 (1985); VI. Stipeae, loc. cit., 40, 731-736 (1985); VII. Pommereulleae, Orcuttieae& Pappophoreae, loc. cit., 40, 737-744 (1985); VIII. Arundinoideae, loc. cit., 41, 323-338 (1986); IX. Centothecoideae, loc. cit., 41, 339-342 (1986); X. Bambuseae, loc. cit., 42, 201-207 (1987); XI. Paniceae, loc. cit., 42, 739-768 (1987). Romero C., 1988.- Numeros cromosomaticos de plantas occidentales, 472-486. Anales Jard. Bot. Madrid, 45, 273-279. Romo A., 1986.- Oeschampsia cespitosa subsp. pyrenaica Romo, subsp. nova. Folia Bot. Misc., 5, 25-27. Rouy G., 1927.- Oeschampsia. In: Flore de France, Asnieres, Paris & Rochefort, 14, 110-116. Rozhevits R.Y., 1934.- Deschampsia. In: Flora of the U.S.S.R. V.L. Komarov (ed.) (Reprint Bishen Singh Mahendra Pal Singh & 0. Koeltz Sc. Books. India & W Germany), 2, 243-252. Smythies B.E, 1986.- Flora of Spain and the Balearic Islands. Checklist of Vascular Plants. Ill. Sa/icaceaeZygophyllaceae, Agavaceae-Zannichelliaceae. Eng/era, 3, 600-654. Stace C.A., 1991.- New flora of the British Isles. Cambridge, 1226p. Tutin T.G., 1980.- Ho/cus. In: Flora Europaea. T.G. Tutin, V.H. Heywood, N.A. Burges eta/. (eds.), Cambridge, 5, 230-231. Vivant J., 1978.- Sur deux sous-especes iberiques nouvelles de Deschampsia cespitosa (L.) P.B. Bull. Soc. Bot. France, 125, 313-318. Watson L., H.T. Clifford & M.J. Dallwitz, 1985.- The classification of Poaceae: subfamilies and supertribes. Australian J. Bot., 33, 433-484. Watson L. & M.J. Dallwitz, 1988.- Grass genera of the world. Canberra, 45p. Watson L. & M.J. Dallwitz, 1992.- The grass genera of the world. C.A.B. International Cambridge, 1038p. Willkomm M., 1861 a.- Oeschampsia. In: Prodromus Florae Hispanicae. M. Willkomm & J. Lange (eds.), Stuttgart, 1, 65-67. Willkomm M., 1861 b.- Holcus. In: Prodromus Florae Hispanicae. M. Willkomm & J. Lange (eds.), Stuttgart, 1, 73-74. Willkomm E., 1893.- Supplementum Prodromi Florae Hispanicae. Stuttgartiae, 370p. Zemann M., 1906-1907.- Die systematische Bedeutung des Blattbaues der mitteleuropaischen Aira-Arten. 6sterr. Bot. Z., 56, 429-436 (1906); loc. cit., 57, 1-4 (1907).

298 ANNEX I PROVENANCE OF THE STUDIED MATERIAL 1


1 The studied material under experimental conditions is marked •. and that which survived 12 months under those conditions ••. For 0. flexuosa, patterns A, B and C are marked respectively with these letters.

1. D. cespitosa subsp. cespitosa Studied material.- SPAIN. Huesca. Benasque, Pia d'Estan-Pia d'Aiuguallut, CH02, 7.VIII.1986, (JACA 550786). Bielsa, Valley Trigoniero, BH73, 29.VII.1992, (JACA 295092). Bisaurri, BH90, 21.VII.1992, (UNEX 21434 .. ). Gistain, Viados, BH82, 23.VIII.1981, (JACA 256781). Jaca, YN01, 6.VII.1971, (JACA 421171). Jaca, Oroel, Y.N01, 26.VIII.1992, (UNEX 21435.). Laguarta-Cafiardo, YN30, 3.VIII.1984, (JACA 111684). Panticosa, YN23, 12.VIII.1983 (JACA 252483), 24.VII1.1992, (UNEX 21436.). Salient de Gallego, El Petruso, YN14, 21.VII.1982, (JACA 86882). Tramacastilla de Tena, YN18, 27.VIII.1982, (JACA 129382). Villacarli, BG99, 21.VII.1988, (JACA 631788). La Corui'\a. Santiago de Compostela, El Visa, NH34, 24.VI.1956, (MA 178130). Leon. Chozas de Arriba, TN71, 4. Vll.1981, (LEB 35649). Tejeda de Ancares, PH84, 28. Vll.1983, (LEB 22423). Lerida. Lake of San Mauricio, CH 12, 16.VII.1983, (VAL 8695), 19.VIII.1992 (UNEX 21437"), 21.VII.1983, (VAB 831562). Lake Redo, CH12, 9.VIII.1980, (LEB 5842). Valley of Aran, Aiguamoix, CH32, 2.VII.1987, (MA 372939). Viella, Ribagorza river, CH23, 20. Vlll.1992, (UNEX 21438 .. ). Navarra. Goizueta, Artikutza, WN98, 7.V11.1982, (MA 432697). Irati river, Pool of lrubia, XN56, s.d., (JACA 803587). Valley of Roncal, XN64, 9.VII.1955, (MA 175114). lsaba, Zuriza. XN74, 26.VII.1969, (SEV 104317). Vizcaya. Ochandiano, WN26, s.d., (JACA-PV 2720-83). PORTUGAL.- Alto Douro. Barroso, Lameiro Grande, PG03, 23.VI.1943, (MA 7807). 2. D. cespitosa subsp. subtriflora var. subtriflora Studied material.- SPAIN. Albacete. Almenaras, WH46, 21.VII.1992, (UNEX 21439 .. ). Riopar, WH46, 21.VII.1992, (UNEX 21440 .. ). Burgos. Huerta del Rey, UM73, 23.VII.1992, (UNEX 21441 .. ). Castellon de Ia Plana. Bejis, El

Rasinero, XK81, 30.VI.1984, (VAL 7976). El Taro, LaAimarja, XK82, 27.VII.1982, (VAL 6332), 30.VI.1984, (VAL 7969). Villafranca, Els Monllats, YK27, 1.VII.1987, (VAB 870957). Cuenca. Aliaguilla, Los Chicoteros, XK40, Vl.1978, (VAB 780320). Mira, Peak Rebello, XJ39, Vl.1979, (VAB 790570). Port of Cubillo, WK78, 21.VII.1992, (UNEX 21442 .. ). Talayuelas, XK71, Vl.1977, (VAB 770573), Vl.1978, (VAB 780321). Tragacete, Guadalaviar, WK96, 21.VII.1992, (UNEX 21443 .. ). Valdemeca, Jucar, XK05, 21.VII.1992, (UNEX 21444.). Granada. Baza Range, Prados del Rey, WG13, 16.VII.1985, (GDAC 26041). Guadalajara. Terzaga, Chequilla,WK99, 13.V11.1965, (SEV 6261). Terzaga, Peralejos de las Truchas. WK99, 13.VII.1965, (SEV 6263). Huesca. Bisaurri, Coli de Fadas, BH90, 21.VIII.1992, (UNEX 21445 .. ). Jaca, Boalar de Atanis, XN91, 3.VII.1969, (SEV 43170). Vilas de Turbon, Porrodruno, BG99, 12.VIII.1985, (SEV 125684). Jaen. Segura Range, WH21, IX.1954, (MA 199625). El Pozo Range, Peak Cabanas, WG08, 24.VI.1980, (SEV 102173). Leon. Burgo Ranero, UM19, Vlll.1982, (LEB 14036). Camposolillo, UN16, 4.VII.1979, (LEB 14825). Carrasconte, QH25, 9.VIII.1984, (LEB 24583). Castrillino, TT92, 5.VII.1982, (LEB 14606, LEB 14604). Chozas de Arriba, TN70, 25.VI.1982, (LEB 14813), 4.VII.1981, (LEB 36161). Cofifial, Pinzon Valley, UN16, 13.VIII.1972, (LEB 4940), 20.VII.1975 (LEB 26305, LEB 26639). Corporales, QG18, 11.VII.1976, (LEB 4947), 11.VII.1977, (LEB 15132). Geras de Gordon. TN75, 12.VII.1982, (LEB 11869). La Vecilla,UN04, 27.VII.1978, (LEB 14834). La Vega de Los Viejos, QH26, 30.VII.1982, (SEV 125869). Lake of La Baiia, PG98, 8.VIII.1982, (LEB 28196). Lucille, QG29, 2.VIII.1974, (LEB 4941, LEB 8968). Mirantes de Luna, TN65, Vll.1974, (LEB 4946). Pool of Luna, TN65, 2.VII.1992, (UNEX 21446.). Pool of El Parma, UN16, 22.VI.1974, (LEB 1011), 5.VII.1982 (LEB 14607, LEB 14605). Puebla de Lillo, UN16, 17.VI.1972, (LEB 7987), 17.VI.1973, (LEB 7985), 17.VI.1974, (LEB 7986). Port of San Gloria, UN57, 6.VIII.1972, (LEB 4945). Riafio, UN36, 12.VI.1969, (LEB 41243). Teleno Range, QG19, 26.VII.1977. (LEB 9061). Tolibia, UN06, 8.VII1.1977, (LEB 14824). Valdearcos, UNDO, 14.VI.1982, (LEB 30142). Valdepolo, Lake Santiz, UN11, 7.VII.1982, (LEB 35607). Vegacervera, TN90, 11.VII.1970, (LEB 4942). Vegas del Condado, UN02, 3.VIII.1992, (LEB 4944). Villamoratiel de las Matas, UN09, s.d., (LEB 28228). Villargusan, TN56, 2.VII.1969, (LEB 11711). Villaverde de Ia Chiquita, UN21, 10.VII.1982, (LEB 35560). Navarra. Viana. Pool of las Canas, WN40, s.d., (MA364814). Zuriza, lsaba, XN74, 25.VIII.1992, (UNEX 21447'). Palencia. Castrejon de Ia Pefia, UN64, 17.VII.1971, (SEV 24014). Cervera de Pisuerga, UN74, 13.V11.1980, (SEV 102304). Salamanca. Tonobron, UL24, 19.VII.1959, (SEV 30781). Segovia. Somosierra, UL55, 23.VII.1992, (UNEX 21448.). Soria. Agreda, WM93, 23.VII.1992, (UNEX 21449 .. ). Bayubas de Abajo, WL19, 17.VII.1982, (VAB 902361, VAB 800782). Berlanga, Duero river, WL19, Vlll.1978, (VAB 780322). Boos, WMOO, 26.VII.1984, (VAB 843146). Near Soria, WM42, 4.VII.1964, (SEV 6265). Teruel. Abejuela, XK82, 22.VI.1984, (VAB 843144, VAB 902362). Abejuela, El Gallotero, XK82, 4.VIII.1984 (VAL 7971). Bronchales. XK18, Vll.1977, (VAB 770575, VAB 912289). Cantavieja, La Palomita, YK18, Vlll.1981 (VAL 8694, VAB 81 0728). El Tremedal, XK18, 24.VII.1988, (VAB 882800). Griegos, XK17, 21.VII.1992, (UNEX 21450.). Gudar, XK97, Vll.1975, (VAL 2695, VAB 760695). Javalambre, Camarena de Ia Sierra, XK63, 20.VI.1990, (VAB 901193). Lake of Gallocanta. XL23, 1O.VII.1959, (SEV 6260). Nava Torrijas, XK73, 7.VII.1984, (VAL 7973), 22.VI.1984, (VAB 843145). El Pobo Range, XK78, 23.VI.1984, (VAB 843143). Valdelinares, XK07, 18.VI.1983, (VAB 831574). Valencia. Aras de Alpuente, Los Tornajos, XK62, 19.VII.1984, (VAL 7975). El Cabezo de Arroyo Cerezo, XK34, 18.VI.1987, (MA 383173), 15.VI.1988, (VAB 881130). Valladolid. Ataquines, UL46, 24.VI.1968, (LEB 18208). Sardon, UM93, Vll.1983, (LEB 14818), Vlll.1982, (LEB 14821 ). Medina del Campo, UL37, 7.VII1.1975, (SEV 25463).

299

PORTUGAL. Minho. lnsalde, Paredes de Coura, NG34, Vl.1916 (COl).

3. D. cespitosa subsp. subtriflora var. congesta


Studied material.- SPAIN. Avila. El Trampal, Bejar Range, TK66, 28.VIII.1997, (MA 253894). Canchal Negro, TK76, 9. VII. 1987, (MA 489604). Circo de Gredos, UK05, 15. VII. 1982, (MA 406341 ). El Barco de Avila, La Covacha, TK75, 27.VII.1982, (MA 253896). Laguna Grande de Gredos, UK05, 15.VIII.1982, (MA 406369). Central Massif, Barrerones, 15.VIII.1982, (MA 406385). Central Massif, Morezon, UK06, 14.VIII.1982, (MA 406342). Central Massif, Navamediana, UK06, 29.VII.1992, (MA 406370). Peak Almanzar, UK06, 19.VII.1980, (LEB 26476), 17.VII.1992, (UNEX 21451"). Piedrahita, Port of Pena Negra, UK07, 26.VI1.1982, (MA 253895). Salamanca. Candelario, Hoyamoro, TK76, 22.VIII.1983, (MA 286029). El Trampal, TK66, 10.VII.1989, (MA 472461).

4. D. cespitosa subsp. media Studied material.- SPAIN. Burgos. Penahorada, Hentomin, VN41, 6.VIII.1977, (BC 630782). Gerona. Estavar, Val d'Estahuja, DG04, 25.VII.1920, (MA 144883). Lerida. Llivia, Sareja, DG09, 13.VIII.1994, (UNEX 21452"). Teruel. Cantavieja, La Palomita, XK18, Vll.1976, (VAB 902363). Valencia. Ayora Range, Peak Caroche, XJ93, 21.VI.1906, (MA 144874), 4.VII.1915, (SEV 6264). Enguera, Casas de Esnali, XJ91, s.d., (VAL 2810). Enguera Range, XJ91, IX.1979, (VAB 790571 ). FRANCE. Herault. Fondfroide, Montpellier, 1.VII.1853, (G 8081-12, G 8081-85). Hills near Montpellier, 1810, (G 8081-67, G 8081-117, G 8081-116, G 8081-81, G 8081-115). Rettinchiere, 28.1X.1836, (G 8081-113). Roquehaute, Agde, 9.VII.1873, (G 8081-65). Seine et Marne. La Genevraye, Ia Valette, Montpellier, 20.VI.1932, (G 8081-44). Valmargues, 19.VII.1838, (G 8081-46, G 8081-56, G-DC). Sommoinere, Montpellier, s.d., (G 8081-114). 5. D. flexuosa Studied material.- SPAIN. Asturias. Aller, Ladera del Oso, TN88, 21.VII.1982, (LEB 11425, A). Arbas-Leitariegos, QH16, 6.VII.1982, (LEB 13799, B). Arbas -Caldas, TH15, s.d, (SEV 6258, B). Cueto de Arbas, QH06, 8.VIII.1982, (LEB 14875, B). Peak Ferreirua, QH37, 22.VII.1982, (LEB 14874, B). Port Cerredo, QH05, 31.VII.1982, (LEB 13859, C). Avila. Gredos, UK06, 27.VII.1978, (SEV 104326, A). Port of La Lancha, Malagon, UL80, 11.VII.1965, (SEV 6256, A). Port of Menga, Alberche, UK38, 28.VII.1978, (SEV 104327, A). Barcelona. Estany, DG23, 19.VIII.1964, (SEV 104321, A). Burgos. Salas de los lnfantes, UM75, 12.VII.1969, (VAB 920260, B). Caceres. Tornavacas, Peak of Calvitero, TK76, 19.VII.1988, (UNEX 9986, A). Cantabria. Los Carabeos, UN15, 7.VIII.1982, (LEB 14375, B; LEB 14373, B). Port of San Gloria, UN57, 31.VII.1970, (SEV 104324, C). Castellon de Ia Plana. Vistabella, Barranco de Azor, YK26, 19.VII.1987, (VAB 890625, B). Gerona. Ribes de Freser, Nuria, DG39, 16.VIII.1992, (UNEX 21453, A'). Granada. Monachil, UG51, 1O.VII.1981, (SEV 111636, A). Sierra Nevada, VG60, 9.VII.1967, (SEV 104332, A). Sierra Nevada, Penones de San Francisco, VG60, 2.VII.1978, (SEV 103773, A); 6.VII.1979, (SEV 104325, A); 6.VII.1987, (VAL 16664, A).Terreras Azules, VF69, 22.VII.1980, (GDAC 11233, A). Guadalajara. Aldeanueva de Atienza, El Molodron, UL95, 18.VIII.1965, (SEV 19090, A). Peak Ocejon, UL75, Vlll.1965, (SEV 10726, A). Huesca. Aisa, La Magdalena, XN92, 8.VII.1966, (SEV 20293, B). Aneta, CH02, 22.VIII.1992, (UNEX 21454, B'). Astun, Port of Somport, YN04, 24.VIII.1992, (UNEX 21455, B"). Borau, Los Lechines, XN82, 23.VIII.1976, (SEV 101351, B). Bujaruelo, YN33, 25.VII.1982, (VAL 8696, B). Formigal, Sorrio, YN14, 24.VIII.1992, (UNEX 21456, B). Hecho, Selva de Oza, XN83, 25.VIII.1992, (UNEX 21457, B). Port of Cotefablo, YN32, 24.VII.1969, (SEV 104318, B). Valley of Anso, Zuriza, XN74, 6.VIII.1987, (VAB 870636, B). Leon. Abelgas, TN55, 2.VII.1975, (LEB 9347, C); Vll.1975, (LEB 4934, C). Ancares, Pena Cuina, PH74, 25.VII.1970, (LEB 4933, B; LEB 4936, B). Aralia, TN75, s.d., (LEB 26141, C). Catoute, OH14, 26.VII.1973, (LEB 22980, B; LEB 22976, B). Chana de Somoza, QG29, 21.VI.1977, (LEB 11880, B). Cofinal, San Justo, MN16, 12.VII.1976, (LEB 38853, B). Cornion, QH16, 17.VIII.1983, (LEB 19142, B). Fuente del lnfierno, UN37, 25.VII.1981, (LEB 14388, B). La Bana, PG98, 20.VI.1981, (LEB 37384, C). La Majua, QH46, Vlll.1973, (LEB 28421, B; LEB 11743, B). Lazado, QH34, Vlll.1974, (LEB 4938, A). Leitariegos (LEB 14317, B; LEB 13696, B; 12688, C). Leitariegos, Arbas, QH16, 6.VII.1982, (LEB 13800, C). Leitariegos, Pena del Miro, QH16, 17.VII.1984, (LEB 24229, A; LEB 14960, A; LEB 24232, C). Llanaves de Ia Reina, UN46, 18.VIII.1979, (LEB 27514, C). Marana, UN26, 16.VII.1980, (LEB 11225 C); 22.VII.1980, (LEB 12906, C); 25.VII.1980, (LEB 12785, C). Peak Valgrande, 19.VII.1978, (LEB 20957, B). Peak Brana Caballo, TN86, 27.VII.1981, (LEB 26140, A; LEB 14889, A). Pena del Seo, Cadafresnas, PH71, 5.VII.1985, (LEB 37425, C; VAB 910084, C). Pena La Arena, 19.VII.1978, (LEB 20958, A). Pena Trevinca, PG88, 16.VII.1982, (LEB 18990, C). Ponferrada, QG09, 28.VIII.1979, (SEV 123661, C). Pool of Riano, UN36, 28.VI.1982, (VAL 21137, B). Port of La Magdalena, QH25, Vll.1976, (LEB 9932, B). Port of Las Senales, UN17, 29.VII.1974, (LEB 26289, B), s d, (LEB 37832, C). Port of Piedrafita, QH25, 30.VII.1981, (LEB 14500, C). Port of San Gloria, UN57, 28.VI.1982, (VAL 21136, B); 8.VIII.1981, (LEB 14083, B) Port of San Isidro, UNO?, 15.VII.1977, (LEB 20455, B; SEV 11642, C). Port of Vegarada, TN96, 20.VII.1978, (LEB 17101, A). Port of Ponton, UN37, 12.VII.1977, (LEB 27569, C). Puebla de Lillo, UN17, 20.VII.1980, (LEB 12806, C). Riosol, UN17, 16.VII.1980, (LEB 11236, C). La Filera, QG95, 13.VIII.1976, (LEB 11595, C). Teleno, QG19, 18.VII.1978, (LEB 8283, B; LEB 11901, B; LEB 37363, C). San Emiliano, QH46, 3.VIII.1971, (SEV 12318, C).Torrebarrio, TN56, 11.VII.1981, (LEB 14115, C). Valdecesar, UN05, 9.VI.1978, (LEB 10841, B). Vivero, QH24, 27.VI.1977, (LEB 21015, B). Lerida. AigOes Tortes, Lake Ratera, CH31, 19.V.1983, (VAL 6533, C). Bohi, CH21, 20.VIII.1992, (UNEX 21458, B"). Espot, CH31. 19.V.1983, (VAB 902364, C). Espot, AigOes Tortes, CH31, 19.VIII.1992, (UNEX 21459, B"). Seo de Urgel,


300 Guils to Port of Canto, CG59, 22.VII.1969, (SEV 102305, B). Siall, Collado de Faidella, CG36, 18.VIII.1992, (UNEX 21460, B .. ). Logroiio. Port of Piqueras, WM35, 4.VII.1964, (SEV 6253, B). Lugo. Ancares, Pena Rubia, PH73, 21.VII.1952, (SEV 6252, B). Caurel, Oevesa de Rogue ira, PH 51, 24.VI.1979, (SEV 121852, C). Pena Esc rita, PH42, V.1976, (LEB 8545, B). Madrid. Cercedilla, VL11, 23.VII.1968, (SEV 11585, C); 24.VII.1953, (MA 163686, C). Guadarrama, Fuente de los Ge61ogos, VL11, 23.VI.1981, (VAL 8692, A). Guadarrama, Lake of Penalara, VL12, 4.1X.1965, (SEV 10727, A). Navarra. Garralda, XN45, 17.VIII.1988, (VAB 883552, B). Salamanca. La Alberca, Pena de Francia, OE48, 4.VII.1946, (SEV 6254, A). Segovia. Navafria, El Chorro, VL34, 13.VII.1980 (SEV 104323, A). Somosierra, VL45, 23.VII.1992, (UNEX 21461, c··). Soria. Agreda, Port of Piqueras, WM43, 23.VII.1992, (UNEX 21462, c·). Vinuesa, WM24, Vlll.1978, (VAB 780319, B). Teruel. Bronchales, XK28, Vll.1977, (VAB 770572, B). El Tremedal, XK18, 24.VII.1988, (VAB 882917, B). Zamora. Segundera Range, PG76, 19.VII.1973, (SEV 103776, A). Zaragoza. Calatayud, Vicort, XL 17, 24.VI.1950, (SEV 104320, A). Moncayo, WM92, 12.VII.1980, (GOAC 13349, A); 21.VII.1976, (SEV 103380, B); 23.VII.1992 (UNEX 21463, s-·; UNEX 21464, B••); 22.VII.1964, (SEV 6257, B); s.d. (SEV 121851, B). Moncayo, Fountain of Tres Canes, WM92, 11.VI.1983, (VAL 8698, A). PORTUGAL. Algarve. Monchique, B. da Maceira, NB33, 29.VII.1979, (UNEX 1713, C). Beira Alta. Serra da Estrella, Covao das Vacas, PE16, Vll.1982, (UNEX 1871, C). Minho. Peneda, Corga da Matanor< aor< adjacent

7-9 subquadrate

undulated

subnuunded or ovoid with silica cells

303

Tableau 1.- Resume des caracteristiques anatomiques observees dans le materiel etudie. T.S. LAMINA, section transversale du limbe; -, cellules bulliformes non visibles ; (), quelquefois ; (()), rare.


EPIDERMIAL CELLS Adaxial surface Abaxial surface Pubescence Pubescence Size!.!!!!!) Size!.!!!!!)

BULLIFORM CELLS No.!Shape 1\lsitioo

4-20 X 4-20

papillae commonly short stiff hairs present±