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.     

Carnivore involvement in bone assemblages based on taphonomic and zooarchaeological analyses of Cova del Coll Verdaguer site (Barcelona,Iberian Peninsula)

Abstract

Carnivores are the main biological agents identified in the Upper Pleistocene assemblage of Cova del Coll Verdaguer (NE Iberian Peninsula). At the site, ungulate bones were primarily accumulated and modified by hyenas, as shown by partly digested bones, coprolites and the pattern of damage on bones. However, other small- to medium-sized carnivores seemed to be involved in the accumulation, owing primarily to the presence of their bones and coprolites. Finally, other carnivores, including the brown bear, used the cave to hibernate and for birthing cubs. The combination of zooarchaeological and taphonomic studies, together with previous coprogenic analyses, at the site indicates that it is only by using these combined approaches that it is possible to identify the majority of the agents involved in an accumulation.     

The Late Holocene extinction of Pinus sylvestris in the western Cantabrian Range (Spain)

Aim To reconstruct the historical biogeography of Pinus sylvestris in the Cantabrian Range (Iberian Peninsula) during the Holocene, and to consider the interactions between vegetation dynamics, climate change and the role of man in the present‐day distribution of the species. Location The study site is a mire (1300 m a.s.l.) at Vega de Viejos, on a south‐facing slope of the western Cantabrian Range, Spain. The region’s present‐day landscape is almost treeless, with the exception of some patches of Quercus pyrenaica and a few copses of Salix and Betula along stream banks. Methods Tree macrofossils from Vega de Viejos were studied by transmitted light and dark‐field reflection microscopy; strobili were subjected to comparative morphological analyses. Two Pinus macrofossils were dated by conventional ¹⁴C methods. Results The taxonomic accuracy achieved in the identification of the macrofossils provided new information regarding the Holocene history of Pinus sylvestris in this territory. Ninety‐five cones of this species were identified; in fact, more than 80% of the 36 identified wood remains were of Pinus gr. sylvestris. Radiocarbon dating revealed that the forest to which the fossils belonged was present until at least 2170 ± 50 yr bp – its disappearance was therefore relatively recent. Main conclusions Pinus sylvestris suffered long‐term isolation, and after the Würm glacial period tended to migrate towards the east. In western Iberia, a temperate climate and autogenic succession favoured broadleaved taxa at the expense of Pinus. Late Holocene human disturbances may have further accelerated the decline of P. sylvestris; in the Cantabrian Range, only a few stands on southern slopes have persisted until the present day. The history of the capercaillie (Tetrao urogallus), a bird characteristic of pure or mixed Palaeartic coniferous forests, was almost certainly affected by the demise of these forests in this area. Cantabrian capercaillies are the only members of this species that live in purely deciduous forests, perhaps a recent adaptation to the regional extinction of pines. Today’s P. sylvestris and capercaillie populations are now highly fragmented and their future, given the predictions of global climate change, is uncertain.     

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.     

First occurrence of the extinct deer Haploidoceros in the Iberian Peninsula in the Upper Pleistocene of the Cova del Rinoceront (Castelldefels,Barcelona)

The Cova del Rinoceront is the first site in the Iberian Peninsula where the genus Haploidoceros has been documented. This discovery of abundant remains is also its first recorded occurrence in Europe during the Upper Pleistocene. The new fossil record strengthens claims that this genus was widely distributed during the Pleistocene, occupying a longer time span at least until MIS 5. Cranial and post-cranial skeletal remains ascribed to this species were recovered from the uppermost layers of the Cova del Rinoceront. The diagnostic features that allow these remains to be unequivocally assigned to H. mediterraneus, include their cranial morphology and antler shape, comprising two sickle-shaped beams, curving backwards and laterally, and a long frontal basal tine. This paper highlights the problems of deer systematics and identification given the high degree of fragmentation of the genus, their relatively homogeneous morphology, a virtual absence of reliable characters and the overlap of measurements between species.     

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.     

The European Far West: Miocene mammal isolation,diversity and turnover in the Iberian Peninsula

Aim To analyse the diversity dynamics of Miocene mammalian faunas in the Iberian Peninsula in order to determine whether the patterns are related to the dispersal of taxa from other areas into this region. Location Mainly the Iberian Peninsula, but two close geographical areas (Central Europe and the Eastern Mediterranean) are also considered in some of our calculations. Methods Genus‐level faunal lists for a total of 299 localities from the Iberian Peninsula, covering 10 successive biochronological units [Mammal Neogene (MN) zones] that span from the latest Early Miocene to the early Pliocene (about 17–4 Ma), were compiled. The dataset was expanded with a further 331 localities in Central Europe and the Eastern Mediterranean for the same time span. Next, a taxonomically standardized database was used to create composite faunal lists of micro‐ and macromammalian genera present during each MN zone. Separate genera‐by‐MN‐zone matrices for both micro‐ and macromammals were built for each region. Mean standing diversity as well as origination and extinction rates were calculated for the Iberian Peninsula, and their correlation with preservation rates is discussed. Simpson’s coefficient of faunal similarity with Central Europe and the Eastern Mediterranean was calculated in order to evaluate whether diversity patterns were related to changes in the affinity of the Iberian mammalian faunas with those of other regions. Results Diversity changes in the Iberian macromammalian faunas coincide with periods of increased faunal similarity with other regions, suggesting a relationship to the expansions and contractions of the geographical ranges of the constituent taxa. This pattern is not recognized for micromammals; that is, their diversity trends are not related to changes in geographical ranges. Main conclusions Climatic shifts result in expansions or contractions in the geographical ranges of macromammals, owing to changes in the distribution of their preferred habitats. The lower dispersal ability of micromammals results in a higher extinction risk when habitat fragmentation confines their populations to relatively small environmental patches. Hence, they are more severely affected by climatic changes. Our results thus emphasize the role of climatic forcing in mammalian biogeography and diversity.     

Short human occupations in the Middle Palaeolithic level I of the Abric Romani rock-shelter (Capellades, Barcelona, Spain)

This paper presents a multidisciplinary study on the size of the occupied surfaces, provisioning strategies and behaviour planning at the Romani rock-shelter, using the Middle Palaeolithic record of the level i. This level is dated around 46.000 BP through U/Th ages. A behavioural interpretation is proposed, which emphasises the activities and the systemic value of the archaeological artefacts and structures. Occupation patterns are identified on the basis of the accumulations formed by human activities. These archaeological accumulations, consisting of artefacts and hearths, are easily defined visually as spatial units. The relationships between these accumulations, established by means of refitted remains, indicate that differences can be established between: 1) small and medium-sized occupation surfaces; 2) restricted and diversified provisioning strategies. This variability suggests that different modes of occupation are represented in the same archaeological level. The human activities reveal the generalization of fire technology. In almost all sizes of the occupation surfaces, the exploitation of vegetal resources near the Abric Romani marks the threshold of the restricted provisioning strategy. Limited use and fragmented knapping activities are recorded in the lithic assemblage. Faunal remains show differential transport. The exploitation of lithic, faunal and vegetal resources characterizes the diversified provisioning strategy. The small occupation surfaces and restricted provisioning strategies suggest short settlements in the Abric Romani. This shorter occupation model complements the longer diversified provisioning strategy recorded in both small and medium-sized occupied surfaces. The selection of precise elements for transport and the possible deferred consumption in the diversified provision strategy suggest an individual supply. In this respect, Neanderthal occupations in the Romani rock-shelter show a direct relation to: 1) hunting strategic resources; 2) high, linear mobility.     

Amphibians and reptiles from the Early Miocene of the Bardenas Reales of Navarre (Ebro Basin, Iberian Peninsula)

The Lower Miocene deposits of the Bardenas Reales of Navarre (NW Ebro Basin, northern Iberian Peninsula) have yielded a diverse vertebrate fauna, including remains of amphibians and reptiles. These remains occur in several localities in the Tudela Formation. The fossiliferous levels belong to the Biozones MN2b-3 (Biozones Z-A of the Ramblian, i.e., Late Aquitanian to Early Burdigalian in age). The amphibians and reptiles represent at least 13 out of 37 vertebrate species. Amphibians consist of a salamandrid urodele and two or three anurans. All the turtles are cryptodirans and consist of the chelydrid Chelydropsis apellanizi, the testudinids Ptychogaster (Temnoclemmys) bardenensis and Ptychogaster ronheimensis, and a Trionychinae indet. Squamates are represented by the anguid lizard Ophisaurus sp., a non-anguid lacertilian, an amphisbaenian, the erycine boid? Eryx sp., and indeterminate colubrids. Crocodilian remains are assigned to the basal alligatoroid Diplocynodon sp. The fossil associations of the Bardenas Reales of Navarre suggest that the vertebrates lived in the centre of an endoreic basin with stretches of water under intertropical to subtropical climatic conditions.     

Palaeoenvironment and palaeoclimate of the Mousterian-Aurignacian transition in northern Iberia: the small-vertebrate assemblage from Cueva del Conde (Santo Adriano, Asturias)

The transition from the Middle Palaeolithic (Mousterian) to the Upper Palaeolithic (Aurignacian) has been one of the prominent themes in the archaeology of the European Palaeolithic for more than 20 years. One of the most controversial questions concerning this period is the extinction of the Neanderthals and their replacement by modern humans. In this context, Cueva del Conde, located in the northern part of the Iberian Peninsula, is an archaeo-palaeontological site that records the Mousterian to Aurignacian transition. It has been excavated since the beginning of the 20th century, first by the Conde de la Vega del Sella and systematically by a team from the University of Oviedo since 2001. Three main zones have been identified: the External Zone, dated to approximately 39 110 ± 520 BP (level N104); the Entrance Platform, dated between 38 250 ± 390 BP and 34 730 ± 500 BP; and Gallery A with a radiocarbon date of approximately 31 540 ± 400 BP (level N2a2). The small-vertebrate assemblages recovered from the water-screening of all sediment from the excavation campaigns represent at least 21 small mammal, amphibian and squamate taxa. The small-vertebrate associations in the three zones suggest a patchy landscape, dominated by humid meadows and woodland areas with the existence of water in the vicinity of the cave. The climate shows a more continental pattern during the Mousterian, though it was milder during the Aurignacian. The small vertebrates of the Cueva del Conde Mousterian and Aurignacian levels suggest a climate that differed from modern day temperatures, between −1.1 and −4.4 °C (mean annual temperature), placing these assemblages during Interstadials 9 to 7 (Is9 to 7).     

Morpho-environmental characterization of the genus Baldellia Parl. (Alismataceae) in the Iberian Peninsula,Balearic islands and North Morocco

Abstract

The genus Baldellia Parl. has always been a complex taxon. Three species and three subspecies have been proposed for the Iberian Peninsula. The morphological variation of this genus in the Iberian Peninsula, Balearic islands and northern Morocco, and its thermo-pluviometric and altitudinal correlation are the focus of this contribution. Twenty-eight morphological variables were examined and subjected to a multivariate statistical analysis. The three morphotypes observed (Form 1, which includes B. ranunculoides subsp. repens and B. alpestris; Form 2, which includes B. ranunculoides and Form 3 – for the new combination B. ranunculoides subsp. ranunculoides var. tangerina (Pau) J. Rocha, A. Crespí, M. García-Barriuso, R. Almeida, J. Honrado, comb. nova proposed here for the first time) seem to represent different reproductive strategies. Morphological variables related with the presence or absence of stolons and the architecture of the inflorescence; the size and number of fruits are the most discriminating variables. Form 1 represents the asexual morphotype; Forms 2 and 3 correspond to morphotypes for which sexual reproduction is preferential. The environmental approach revealed that the asexual form (Form 1) grows in temperate, in more humid conditions, and within a broad altitudinal range. In contrast, the two sexual forms are more common in warmer and drier conditions, and occur over a narrower interval of altitudes.     

Past distribution and ecology of the cork oak (Quercus suber) in the Iberian Peninsula: a pollen-analytical approach

Abstract.  This study presents pollen-analytical data from continental and offshore Iberian Peninsula sites that include pollen curves of Quercus suber , to provide information on the past distribution and ecology of the cork oak ( Q. suber ). Results centre on a new pollen record of Navarrés (Valencia, eastern Spain), which shows that the cork oak survived regionally during the Upper Pleistocene and was important during a mid-Holocene replacement of a local pine forest by Quercus -dominated communities. This phenomenon appears linked to the recurrence of fire and reinforces the value of the cork oak for reforestation programmes in fire-prone areas. In addition to Navarrés, other Late Quaternary pollen sequences (Sobrestany, Casablanca-Almenara, Padul, SU 8103, SU8113, 8057B) suggest last glacial survival of the cork oak in southern and coastal areas of the Peninsula and North Africa. Important developments also occur from the Late Glacial to the middle Holocene, not only in the west but also in the eastern Peninsula. It is suggested that, in the absence of human influence, Q. suber would develop in non-monospecific forests, sharing the arboreal stratum both with other sclerophyllous and deciduous Quercus and Pinus species.     

Biogeographical analysis of species of the tribe Cytiseae (Fabaceae) in the Iberian Peninsula and Balearic Islands

Aim To analyse quantitatively the biogeographical distribution pattern of species of the Cytiseae Bercht. & J. Presl (= Genisteae Benth.) tribe in the Iberian Peninsula and the Balearic Islands, and to identify environmental variables related to the distributional patterns. Location Iberian Peninsula and Balearic Islands, using the 61 administrative provinces as operational geographical units. Methods In order to identify chorotypes (groups of species with similar geographical distribution), we performed a upgma classification based on the similarity index of Baroni‐Urbani & Buser. The method of McCoy et al. [Ecology 67 (1986), 749] enabled us to detect the significant groups and to differentiate them from those groupings that could be generated at random. Logistic regression analyses and environmental gradient analyses (DCA and CCA) were performed in order to find the relationships between the environmental variables and the observed distributional patterns. Results Sixteen chorotypes were obtained between the Cytiseae species of the Iberian Peninsula and the Balearic Islands. The thermal variables showed the greatest influence in species distribution. Specifically, temperatures (maximum, minimum and mean) of the coldest months were associated with the majority of the chorotypes. Main conclusions The species of the Cytiseae tribe were not randomly distributed in space, and can be classified in groups of species with common distributional patterns. The importance of cold tolerance in the distribution of these species, as well as their preference for acidic soils, was demonstrated. Certain general tendencies seem to exist with respect to the distribution of the biota in the Iberian Peninsula, and these seem to be independent of the taxonomic groups involved and to be determined by convergent macroclimatic factors.     

The ammonite genera Fagesia and Neoptychites (family Vascoceratidae) in the Iberian Trough, Spain

The ammonites assigned to the genera Fagesia Pervinquière, and Neoptychites Kossmat, of the Wiedmann (Tübingen, Germany) and Goy, Carretero and Meléndez (Madrid, Spain) collections obtained from the Iberian Trough have been revised. New mainly lower Turonian specimens of the species Fagesia catinus (Mantell), F. tevesthensis (Peron), F. rudra (Stoliczka), F. superstes (Kossmat), F. pachydiscoides Spath, and Neoptychites cephalotus (Courtiller) have also been presented. In addition we have described one new species: F. mortzestus. Studies of the morphologies and the geographical and stratigraphical distributions of all these species have led to the identification of several phylogenetic relationships between them, and to distinguishing one main phase in the evolution of the family Vascoceratidae Douvillé, characterised by the dominance of Fagesia with Neoptychites.     

Taxonomic notes on Trechini and description of a new hypogean species from the Iberian Peninsula (Coleoptera: Carabidae: Trechinae)

In this contribution, a new pubescent Trechus species for the Iberian Peninsula is described, and the use of some morphological traits in the systematic of Trechini are discussed. Micropubescence is common in endogean and troglobite species which are closely related with the “terrestrial-phreatic habitat”. It probably enhances survival in swampy environments through the retention of air bubbles in immersion conditions and promoting flotation. Neither this trait, anophthalmy nor inner sac of aedeagus without sclerotized pieces can be used to support supraspecific taxa. Thus, we propose the formal synonymy of Hydrotrechus with Trechus [Hydrotrechus syn. nov. of Trechus]. To avoid secondary homonymy, we propose the name Trechus triamicorum nom. nov. for Hydrotrechus cantabricus Carabajal, García & Rodríguez 1999 [already occupied by Trechus cantabricus Jeannel 1927, synonym of Trechus distigma Kiesenwetter 1851]. We also propose the synonymization of the subgenus Trichapoduvalius with Apoduvalius [Trichapoduvalius syn. nov. of Apoduvalius]. The new hypogean species with pubescent elytra, Trechus cifrianae sp. nov., is described. The morphology of the species suggests that it probably inhabits the Mesovoid Shallow Substratum (MSS) or the fissure network (mesocaverns and microcaverns) of the cave where the only individual was found. The structure of internal female genitalia suggests a close relationship between T. cifrianae sp. nov. and T. carrilloi Toribio & Rodríguez 1997.     

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.     

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.     

The effect of air temperature on the starting dates of theUlmus, Platanus andOlea pollen seasons in the SE Iberian Peninsula

A study is made of the effect of air temperature on the start of the pollen seasons of three tree species—Ulmus, Platanus andOlea—in the southeastern Iberian Peninsula. These initial results are based on an aerobiological analysis performed over a 4-year period in the city of Granada, Spain. Sampling was carried out with a Burkard spore trap. The main aim of the present study was to use regression analysis to identify the preseasonal date when these species start to accumulate heat in their floral buds, enabling models to be created that indicate the onset of the pollen season for these taxa. ForUlmus there was no significant correlation between the onset of pollination and the mean temperature during the preceding period, whereasPlatanus presented a significant correlation with mean temperature during the month of January, andOlea with mean temperature during the second fortnight in February and the month of March. ForUlmus, the start of the pollen season ranged between 30 January and 8 February; forPlatanus, between 13 and 17 March; and forOlea, between 20 April and 13 May. The accumulated temperature needed to induce the onset of flowering ranged between 121 and 256°C forUlmus, with values of 428–607°C and 656–881°C forPlatanus andOlea, respectively.     

The rise and fall of the mountain hare (Lepus timidus) during Pleistocene glaciations: expansion and retreat with hybridization in the Iberian Peninsula

The climatic fluctuations during glaciations have affected differently arctic and temperate species. In the northern hemisphere, cooling periods induced the expansion of many arctic species to the south, while temperate species were forced to retract in southern refugia. Consequently, in some areas the alternation of these species set the conditions for competition and eventually hybridization. Hares in the Iberian Peninsula appear to illustrate this phenomenon. Populations of Iberian hare (Lepus granatensis), brown hare (Lepus europaeus) and broom hare (Lepus castroviejoi) in Northern Iberia harbour mitochondrial haplotypes from the mountain hare (Lepus timidus), a mainly boreal and arctic species presently absent from the peninsula. To understand the history of this past introgression we analysed sequence variation and geographical distribution of mitochondrial control region and cytochrome b haplotypes of L. timidus origin found in 378 specimens of these four species. Among 124 L. timidus from the Northern Palaearctic and the Alps we found substantial nucleotide diversity (2.3%) but little differentiation between populations. Based on the mismatch distribution of the L. timidus sequences, this could result from an expansion at a time of temperature decrease favourable to this arctic species. The nucleotide diversity of L. timidus mtDNA found in Iberian L. granatensis, L. europaeus and L. castroviejoi (183, 70 and 1 specimens, respectively) was of the same order as that in L. timidus over its range (1.9%), suggesting repeated introgression of multiple lineages. The structure of the coalescent of L. granatensis sequences indicates that hybridization with L. timidus was followed by expansion of the introgressed haplotypes, as expected during a replacement with competition, and occurred when temperatures started to rise, favouring the temperate species. Whether a similar scenario explains the introgression into Iberian L. europaeus remains unclear but it is possible that it hybridized with already introgressed L. granatensis.