Phenomorphology and ecomorphological characters of Maytenus senegalensis L. Shrublands in the Iberian Peninsula: A comparison with other Mediterranean plant communities

This work sets out to establish ecomorphological and phenomorphological patterns in the constituent flora of an Iberian Maytenus senegalensis plant community in order to relate it with or separate it from other types of Mediterranean vegetation. This plant community contains tall shrubs, scrubs, palms and vines of a variety of families. Typical ecomorphological adaptations to the Mediterranean climate were observed in the community studied: an abundance of thorny species or species with tomentose leaves and photosynthetic stems, small leaves and a high presence of species with organs capable of post-fire regeneration. However, the community also showed hybrid characteristics between scrubland (predominantly malacophyllous leaves lasting 9 months on average and low biomass) and tall shrubland communities (predominance of phanerophytes up to 3 m in height, predominantly smooth bark, periodical reposition of leaves, mean lifespan of plants 25 years and mainly fleshy fruit). The community also presents an outstanding quantity of amphiphytes (29%). From a phenological point of view, the formation of buds in winter, flowering in spring, summer–autumn seed dispersal and leaf shedding in summer are typical of other Mediterranean communities, while fructification and growth lasting three seasons, as in the case of Maytenus shrubland, is not. The RVA phenophasic index (balance between reproductive and vegetative phenophases) with values of 0.4–2 shows an intermediate value between typical shrubland (0.3–1) and forest community (1–2) values. The APC index (period of phenophasic activity of the community) differentiates it from others, since it remains at 100% for seven months, three of which are winter months. Phenophasic patterns (overlapping, or not, of flower bud formation, flowering and growth) show how similar M. senegalensis shrubland is to forests because of the absence of phenophase pattern type A (characteristic in open-scrublands); in addition, we have detected a new pattern (type G), still regarded as exclusive to M. senegalensis. As regards the existence of coenomorphs (group of species with similar morpho-functional strategies), the studied community showed much greater diversity than other communities studied to date, with an Strategies Richness index (SRi) of 0.79 (11 groups covering the 14 species existing), pointing to the co-existence of a great variety of eco-phenomorphological strategies, each with a successful response to the same ecological conditions. Finally, some of the species in the community, e.g. Asparagus horridus, Chamaerops humilis and M. senegalensis, stand out because of their unique ecomorphological and phenological characteristics.     

Reproductive biology of twoPrunus species (Rosaceae) in the Northwest Iberian Peninsula

We studied the reproductive biology ofPrunus spinosa andPrunus mahaleb (Prunoideae, Rosaceae) in the northwest Iberian Peninsula. The two species flowered at the same time (peaking on March 9 and 11, respectively in 1990) but differ significantly in their fruit maturation times. Nectar volume peaked in the early morning in both species, and was ten times greater inP. spinosa than inP. mahaleb. Neither species shows apomixis, nor does fruit-set occur if pollinators are excluded. In both species self-pollination resulted in fewer fruits than open pollination. The principal pollinators belong to theApidae family (79% and 63% of visits toP. spinosa andP. mahaleb, respectively). Results are compared with those for other rosaceous plants with fleshy fruits.     

Karyology of theScorzonera (Compositae) species from the Iberian Peninsula

A karyological study of 15 taxa ofScorzonera L. from the Iberian Peninsula has been made. The chromosome numbers found inS. hispanica var.pinnatifida, S. baetica, S. reverchonii, S. angustifolia, S. laciniata var.calcitrapifolia and var.subulata (2n = 14) are new. Diploid cytotypes with 2n = 14 and 2n = 12 prevail, andS. hispanica var.crispatula is the only taxon which exhibits autopolyploidy (2n = 14, 28). x = 7 is considered to be the base chromosome number within the genus, with x = 6 being derived from it by translocation. This and detailed karyotype analyses allow to group the Iberian Peninsula species ofScorzonera into three groups.     

Floral rewards in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The genusScrophularia in the Iberian Peninsula and Balearic Islands comprises two sections,Scrophularia andCanina G. Don. Analyses were carried out on flower production, flower duration and their sexual phases, pollen and nectar production together with observations on their pollinators. Nectar production is correlated with corolla size and pollen production with anther size. The taxa of sect.Scrophularia show greater nectar and pollen production than those of sect.Canina. Also, those of the first section produce more ovules per ovary than those of the second group, production being correlated with the ovary size.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and the Balearic Islands. I.     

Multivariate and cladistic analyses of the purple-flowered species ofErysimum (Cruciferae) from the Iberian Peninsula

A taxonomically difficult purple-flowered group within the genusErysimum, restricted to the Iberian Peninsula, is analyzed by multivariate and cladistic analyses. 51 specimens have been scored for 14 characters. Both principal components and discriminant analyses provide support to the recognition of the five species considered by the author, namelyE. linifolium, E. lagascae, E. baeticum, E. popovii, andE. cazorlense. Cladistic analysis, using 7 characters resulted in a single most parsimonious cladogram containing no homoplasies. The pattern of morphologic divergence follows a clear NW.-SE. trend, which is congruent with the topology of the cladogram. This trend significantly affects growth-form as well as fruit characters, both providing the main grounds for species recognition. The different behavior and significance of several characters in both kinds of analysis is discussed. The co-occurrence of morphologically similar individuals differing in the flower color is discussed, too. Possible explanations for this phenomenon involve hybridization in a wide sense or, alternatively, rejecting the assumption of monophyly for the group.     

Isozyme studies in the genusCheirolophus (Asteraceae:Cardueae-Centaureinae) in the Iberian Peninsula, North Africa and the Canary Islands

The phylogenetic and phenetic analysis of 109 RAPD polymorphisms inS. nebrodensis, a perennial and self-incompatible endemic of four mountain ranges in Spain, andS. viscosus, a self-compatible annual widespread in Europe, as well asS. lividus, S. sylvaticus andS. vulgaris revealed a sister group relationship between the first two species. This result contrasts sharply with the earlier hypothesis based on isozyme variation thatS. viscosus originated from within a paraphyleticS. nebrodensis and that the two species represent a progenitor-derivative pair. After considering possible reasons for the sister group relationship found, including the possibility of rooting artefacts, it is concluded that neither the RAPD data nor the isozyme data allow to draw safe conclusions about the mode of speciation and therefore the relative age of the two species. As a consequence, the limited genetic variation ofS. viscosus in comparison toS. nebrodensis as revealed by both the RAPD and the isozyme data may reflect its population history, geographical distribution, reproductive ecology, or mode of dispersal just as well as its recent origin from a paraphyleticS. nebrodensis. The result of this study calls for a critical reexamination of other taxon pairs postulated to have a progenitor-derivative relationship on the basis of isozyme evidence.     

Taxonomic characterization of populations ofHyacinthoides sect.Somera (Hyacinthaceae) in the northwestern Iberian Peninsula

A classical morphological study ofHyacinthoides sect.Somera in the northwestern Iberian Peninsula revealed the new speciesH. paivae. It differs from the closely related speciesH. italica, H. hispanica, andH. mauritanica by vegetative and generative characters, such as leaf length, raceme shape, tepal length and width. It typically occurs in woodlands in the A Coruña province of NW Spain.     

Survival and long-term maintenance of tertiary trees in the Iberian Peninsula during the Pleistocene: first record of Aesculus L. (Hippocastanaceae) in Spain

The Italian and Balkan peninsulas have been places traditionally highlighted as Pleistocene glacial refuges. The Iberian Peninsula, however, has been a focus of controversy between geobotanists and palaeobotanists as a result of its exclusion from this category on different occasions. In the current paper, we synthesise geological, molecular, palaeobotanical and geobotanical data that show the importance of the Iberian Peninsula in the Western Mediterranean as a refugium area. The presence of Aesculus aff. hippocastanum L. at the Iberian site at Cal Guardiola (Tarrasa, Barcelona, NE Spain) in the Lower–Middle Pleistocene transition helps to consolidate the remarkable role of the Iberian Peninsula in the survival of tertiary species during the Pleistocene. The palaeodistribution of the genus in Europe highlights a model of area abandonment for a widely-distributed species in the Miocene and Pliocene, leading to a diminished and fragmentary presence in the Pleistocene and Holocene on the southern Mediterranean peninsulas. Aesculus fossils are not uncommon within the series of Tertiary taxa. Many appear in the Pliocene and suffer a radical impoverishment in the Lower–Middle Pleistocene transition. Nonetheless some of these tertiary taxa persisted throughout the Pleistocene and Holocene up to the present in the Iberian Peninsula. Locating these refuge areas on the Peninsula is not an easy task, although areas characterised by a sustained level of humidity must have played an predominant role.     

10种杜鹃亚属植物种间杂交的可育性研究

为探索杜鹃花亚属内异种杂交的可育性及其规律,对杜鹃亚属有鳞大花亚组(subsect. Maddenia)、三花杜鹃亚组(subsect. Triflora)、亮鳞杜鹃亚组(subsect. Heliolepida)及腋花杜鹃亚组(subsect. Scabrifolia)等4亚组10个杜鹃花种类的22个杂交组合(其中18个数据完整组合)的可育性进行了研究。结果表明:(1)所涉及的杜鹃亚属不同亚组间及三花杜鹃亚组内杂交均较困难,在18个数据完整组合中高可育与可育组合比率明显偏低,不育比率高(55.6%)。(2)在10个不可育或败育组合中,不能坐果(Cab型)、不能结实(Sab型)和可结实而种子不能发芽(Sng型)的数量分布为6∶1∶3,其不亲和或败育发生的阶段可能涵盖了从前合子期到后合子期的整个阶段。(3)亲本一方为多倍体组合的可育率(41.6%),尤其是母本为多倍体时,比二倍体组合的可育率(50.0%)低且无高可育组合出现,部分印证了倍性是导致该亚属植物不同种类杂交不亲和、不育与育性下降的重要原因,但不是唯一原因,亚组间杂交的可育率(16.7%)明显低于亚组内(三花杜鹃亚组内,58.3%)。(4)与相应的母本自然授粉结果相比,杂交明显导致多数可育组合绿苗率比率和单位可育种子数量比率的大幅度下降,这是双亲遗传差异及多倍体亲本介入后所导致的杂交衰退hybrid weakness现象。(5)在多倍体作母本的情况下,杂交单向不育或非对称遗传渗透现象明显。     

The Middle-to-Upper Palaeolithic transition in Cova Gran (Catalunya, Spain) and the extinction of Neanderthals in the Iberian Peninsula

The excavations carried out in Cova Gran de Santa Linya (Southeastern PrePyrenees, Catalunya, Spain) have unearthed a new archaeological sequence attributable to the Middle Palaeoloithic/Upper Palaeolithic (MP/UP) transition. This article presents data on the stratigraphy, archaeology, and ¹⁴C AMS dates of three Early Upper Palaeolithic and four Late Middle Palaeolithic levels excavated in Cova Gran. All these archaeological levels fall within the 34-32 ka time span, the temporal frame in which major events of Neanderthal extinction took place. The earliest Early Upper Palaeolithic (497D) and the latest Middle Palaeolithic (S1B) levels in Cova Gran are separated by a sterile gap and permit pinpointing the time period in which the Mousterian disappeared from Northeastern Spain. Technological differences between the Early Upper Palaeolithic and Late Middle Palaeolithic industries in Cova Gran support a cultural rupture between the two periods. A series of 12 ¹⁴C AMS dates prompts reflections on the validity of reconstructions based on radiocarbon data. Thus, results from excavations in Cova Gran lead us to discuss the scenarios relating the MP/UP transition in the Iberian Peninsula, a region considered a refuge of late Neanderthal populations.     

Natural hybridization origin of <Emphasis Type="Italic">Rhododendron agastum </Emphasis>(Ericaceae) in Yunnan,China: inferred from morphological and molecular evidence

The natural hybridization that occurs between two sympatric species of Rhododendron subgenus Hymenanthes in Yunnan, China, was investigated. The assumed parents, Rhododendron delavayi Franch. and R. decorum Franch., are morphologically distinct, and the putative hybrid species, R. agastum Balf. f. et W. W. Smith, has an intermediate morphology. We used the main morphological characters, sequences of the nuclear ribosomal DNA ITS region, and the chloroplast DNA trnL-F intronspacer to analyze the three species, and compared these morphological and molecular data with an artificial hybrid between R. decorum (♀) × R. delavayi (♂). From the results, we conclude that R. agastum is a natural hybrid between a female R. delavayi and a male R. decorum.     

Sexual reproduction in someScrophularia species (Scrophulariaceae) from the Iberian Peninsula and the Balearic Islands

The reproductive biology of someScrophularia species in the Iberian Peninsula and Balearic Islands has been studied. Levels of fruit-set in bagged and unbagged inflorescences, the production of seeds/capsule, the effectiveness of seed-set following cross-pollination in the genus and the pollen/ovule ratios have been analyzed.Some observations on floral and reproductive biology in some species ofScrophularia from the Iberian Peninsula and Balearic Islands. II. For first part seeOrtega Olivencia & Devesa 1993.     

Habitat and reproductive phenology of wild boar (Sus scrofa) in the western Iberian Peninsula

We examined the reproductive phenology of wild boar populations in four regions of the western Iberian Peninsula during the 1999/2000 hunting season (October–February). To estimate conception dates and birth distribution frequencies, we used foetal weights. Regions differed significantly, and we detected a relationship between region and birth distribution frequencies. Throughout the year, food availability had a major influence on the distribution of farrowing. Although a short period of high food availability leads to highly synchronous births, even in relatively harsh environmental conditions, adult females that exploit low-quality food items appear to be able to give birth at any time of the year.     

A survey of the response of Rhododendron to in vitro culture

The responses of 7 genotypes of Rhodendron to culture conditions and their establishment as shoot cultures are described. The genotypes represent a broad genetic diversity in the genus. After sterilization and an acclimation period of 3 to 12 months, all the selections were established as shoot cultures on Woody Plant Medium (WPM) supplemented with N⁶(-²-isopenteny) adenine (2iP). Plants with strong episodic growth cycles required the longer acclimation periods. Utilizing shoots from these cultures, the response to a cytokinin series of 0 to 32 M 2iP or BAP (6-Benzylaminopurine) was analyzed. BAP proved toxic to all but the elipidote and lepidote rhododendrons (R. mucronulatum, R. x Boule de Neige, and R. x PJM); however, even with these selections, 2iP stimulated greater shoot multiplication rates. The optimum 2iP level for shoot multiplication varied little with the genotype and levels of 4 to 16 M generally proved optimal, depending on the specific selection. Adventitious shoot production was observed in 3 selections (R. canadense, R. x Boule de Neige and R. x PJM), but only at 2iP levels above 8 M. Shoot multiplication rates of 7 to 21 times were observed, depending on the selection. Using an average utilizable shoot production rate of 40 shoots per culture per 6 week subculture period, some 75,000 shoots can be generated per square meter of culture space per year. The harvested shoots (microcuttings) rooted readily out-of-culture and the resultant plants grew like seedlings.     

Phylogeny of Rhododendron section Vireya (Ericaceae) based on two non-coding regions of cpDNA

Section Vireya (Blume) Copel.f. is one of the most morphologically diverse groups of the genus Rhododendron. Vireyas have a unique distribution for the genus, being predominantly found throughout the Malesian Archipelago. The alpha taxonomy of section Vireya is relatively well understood and taxa are easily distinguished from other rhododendrons by their general appearance. Defining characteristics of the section the possession of seeds with tailed appendages at both ends, the twisting of capsule valves after opening, placentas that separate as thread-like structures from the central axis as the capsule opens are all subject to exceptions. Phylogenetic analyses of two cpDNA regions, psbA-trnH and trnT-trnL intergenic spacers are reported. The results of each analysis were generally congruent, with clades relating strongly to geographic areas. Section Vireya is monophyletic with the inclusion of at least one species of section Rhododendron. Only two of the seven subsections currently circumscribed are monophyletic: Malayovireya Sleumer and Siphonovireya Sleumer. ``Euvireya', all subsections excluding the paraphyletic subsection Pseudovireya, is monophyletic and includes two major clades, one restricted to eastern Malesia and the other to the western and middle Malesia.     

Planktonic chlorophyll a and eutrophication in two Mediterranean littoral systems (Mallorca Island,Spain)

Two Mediterranean littoral systems of Mallorca Island have been studied using planktonic chlorophyll a (total concentration and seasonal variations) as trophic indicator. The studied systems are quite different in geomorphology, physicochemical trends and anthropic incidence. S'Albufera-Alcudia Bay is the drainage outlet of a coastal marsh, with a salt wedge estuarine regime. Andratx Harbour is a narrow bay affected by a high anthropic incidence due to the presence of a small fishing port and mainly a large leisure harbour. Both systems are affected by a typical Mediterranean climate and a negligible tide incidence. The present work is based on several year monitoring of physical, chemical and biological water parameters. In S'Albufera-Alcudia Bay, Chl a concentration ranged from 0.01 to 17.10 mg m^–3, the values dropping from the inland site to the sea. The phytoplankton biomass gradient takes its origin in physical and chemical differences. The variance is mainly related to the inland-sea water axis. Nitrogen and silica enriched inland waters are the main source of eutrophication, hence their relation with Chl a. These nutrients entering the Albufera are mainly derived from leaching from farmed soils. The transition from an inland eutrophic system to an oligotrophic marine one show the filter character of these coastal marshes. In Andratx Harbour, Chl a ranged from a non detectable concentration to 6.36 mg m^–3. The ultraplankton (<5 m) was the most important contributor to total concentration. The ultraplankton in this system is related to influence of more oligotrophic open seawater. A greater proportion of microplankton (>20 m) is related both to higher nutrient concentrations and a higher water column stability. In Andratx Harbour, the enclosure state seems very important for development of phytoplankton blooms.     

Taxonomy and cytogeography of Cardamine raphanifolia and C. gallaecica (Brassicaceae) in the Iberian Peninsula

For a long time it was considered that Cardamine raphanifolia is widely distributed with several infraspecific taxa in the Mediterranean region. Recent studies showed that C. raphanifolia is restricted to the northern Iberian Peninsula. Cardamine gallaecica from NW Spain was regarded to be closely related, but was originally regarded as C. amara, and later described as C. raphanifolia subsp. gallaecica. In this study, we validate the treatment of C. raphanifolia and C. gallaecica as distinct species, and show their clear differentiation based on morphometric and molecular AFLP analyses. Moreover, both species are clearly distinct from C. amara subsp. pyrenaea, which has an overlapping distributional range with C. raphanifolia. Hexaploid (2n = 48) and octoploid cytotypes (2n = 64) of C. raphanifolia, and tetraploid (2n = 32) and hexaploid (2n = 48) cytotypes of C. gallaecica were found. Due to their weak morphological and genetic differentiation, these polyploid races are not classified as separate taxa.     

Presence of Hemitragus aff. cedrensis (Mammalia, Bovidae) in the Iberian Peninsula: Biochronological and biogeographical implications of its discovery at Bolomor Cave (Valencia, Spain)

The discovery of new material in Late Pleistocene levels at Bolomor Cave (Valencia, Spain) raises some questions about the presence of the most ancient record of Hemitragus cedrensis in the peninsula, and its dispersal out of Provence. The morphology and dimensions of some lower teeth confirm the identification of H. aff. cedrensis. Moreover, it presents strong similarities, both morphological and metrical, with the specimens from Caune de l’Arago and bau de l’Aubesier (end of OIS 7 to OIS 5e) rather than with the population from the eponymous locality. The data suggest a dispersal event out of Provence towards the Iberian Peninsula during the Eemian. This dispersal was not stopped by natural barriers such as large rivers, or mountains. The results presented here confirm the biochronological interest of the genus Hemitragus for the Late Pleistocene in Mediterranean Europe.     

Presence of internal photosynthetic cylinder surrounding the stele in stems of the tribe <Emphasis Type="Italic">Salicornieae</Emphasis> (Chenopodiaceae) from SW Iberian Peninsula

Fluorescence microscopy and physiological examination revealed the presence of an inner cylinder of active photosynthetic cells located below the endodermis-like layer in stems of four of the six taxa of the tribe Salicornieae (Chenopodiaceae).     

The relationship between freezing tolerance and thermotropic leaf movement in five Rhododendron species

Summary Leaf movement kinetics in five species of Rhododendron were studied in response to leaf temperature, leaf freezing point, and leaf water deficit. There was a gradient in the degree of leaf curling among species in the following order from the greatest curling to the least curling: Rhododendron catawbiense, R. maximum, R. minus, R. macrophyllum, R. ponticum. Those species found to be tolerant of winter conditions had the most intense leaf movements (both curling and angle) while those species with minimal cold tolerance had limited or no leaf movements. Leaf curling occurred at leaf temperatures above the tissue freezing points in all species. Athough leaf angle was influenced by leaf turgor, general tissue desiccation was not the ultimate cause for thermotropic leaf curling in any species tested. Those species with the greatest leaf curling and angle movements had the highest osmotic potential, the lowest water deficit at the turgor loss point, and the lowest symplastic water fraction. These data suggest that there is a trade off in Rhododendron leaf physiology between cold tolerance (due to leaf movements) and water stress tolerance (due to turgor maintenance mechanisms).