Spatial Distribution of Dorylaimid and Mononchid Nematodes from the Southeast Iberian Peninsula: Environmental Characterization of Chorotypes

The aim of this study was to determine the incidence of 18 environmental variables in the spatial distribution of 30 chorotypes (species groups with significantly similar distribution patterns) of dorylaimid and mononchid nematodes by means of logistic regression in a natural area in the southeastern Iberian Peninsula. Six variables (elevation, color chroma, clay content, nitrogen content, CaCO₃, and plant community associated) were the most important environmental factors that helped explain the distribution of chorotypes. The distribution of most chorotypes was characterized by some (one to three) environmental variables; only two chorotypes were characterized by five or more variables, and four have not been characterized.     

Chorotypes of seaweeds from the western Mediterranean Sea and the Adriatic Sea: An analysis based on the genera Audouinella (Rhodophyta), Cystoseira (Phaeophyceae) and Cladophora (Chlorophyta)

The aim of the present paper is to identify the possible existence of groups of species in the genera Audouinella (Rhodophyta), Cystoseira (Phaeophyceae) and Cladophora (Chloropyta) with significantly similar distribution patterns (chorotypes), in the western Mediterranean Sea and the Adriatic Sea. Of the 98 species studied, 59 were grouped into 11 chorotypes, whereas 39 species remained ungrouped. Thirty‐eight species were included in a generalist chorotype, whereas 6 chorotypes were monospecific. The relationships with the environmental factors that could explain the chorotypes are discussed.     

Modelling chorotypes of invasive vertebrates in mainland Spain

We investigated the existence of chorotypes – assemblages of species with similar geographical ranges – of invasive species in a host territory, and their potential use to advocate similar control or management strategies for species in the same chorotype. We analysed the distribution of 13 exotic terrestrial vertebrate species (six birds, six mammals, and one reptile) with well‐known distributions in mainland Spain. We used the presence/absence data on a grid of 10 km × 10 km UTM cells from the Atlases of terrestrial vertebrates of Spain. These data were aggregated to a grid of 50 km × 50 km UTM cells, because it entailed no loss of meaningful information and allowed dealing with a much lower number of cells. Using cluster analysis and a probabilistic assessment of the classification, we identified seven significant chorotypes: four multispecific and three monospecific. The compound chorotypes grouped together species that tended to share certain characteristics about their introduction, release cause, establishment, and spread. We modelled the chorotypes using a favourability function based on a generalized linear model and 31 variables related to spatial situation, topography, lithology, climatic stability, energy availability, water availability, disturbances, productivity, and human activity. Climatic factors affected the favourability for every chorotype, whereas human variables had a high influence in the distribution of three chorotypes involving eight species. On the basis of these variables, we identified favourable areas for all the chorotypes in mainland Spain. The favourability for a chorotype in an area may be a useful criterion for evaluating the local conservation concern due to the whole set of species. Favourable but unoccupied areas can be used to infer possible colonization areas for each chorotype. We recommend using chorotypes to optimize broad‐scale surveillance of invasive species.     

Common Distribution Patterns of Marsupials Related to Physiographical Diversity in Venezuela

The aim of this study is to identify significant biotic regions (groups of areas with similar biotas) and biotic elements (groups of taxa with similar distributions) for the marsupial fauna in a part of northern South America using physiographical areas as Operational Geographical Units (OGUs). We considered Venezuela a good model to elucidate this issue because of its high diversity in landscapes and the relatively vast amount of information available on the geographical distribution of marsupial species. Based on the presence-absence of 33 species in 15 physiographical sub-regions (OGUs) we identified Operational Biogeographical Units (OBUs) and chorotypes using a quantitative analysis that tested statistical significance of the resulting groups. Altitudinal and/or climatic trends in the OBUs and chorotypes were studied using a redundancy analysis. The classification method revealed four OBUs. Strong biotic boundaries separated: i) the xerophytic zone of the Continental coast (OBU I); ii) the sub-regions north of the Orinoco River (OBU III and IV); and those south to the river (OBU II). Eleven chorotypes were identified, four of which included a single species with a restricted geographic distribution. As for the other chorotypes, three main common distribution patterns have been inferred: i) species from the Llanos and/or distributed south of the Orinoco River; ii) species exclusively from the Andes; and iii) species that either occur exclusively north of the Orinoco River or that show a wide distribution throughout Venezuela. Mean altitude, evapotranspiration and precipitation of the driest month, and temperature range allowed us to characterize environmentally most of the OBUs and chorotypes obtained.     

The distribution patterns of reptiles in the Riff region, northern Morocco

A biogeographical classification of reptiles in the Riff region (northern Morocco, Africa) was carried out to look for shared distribution patterns, here termed chorotypes. Baroni‐Urbani & Buser's similarity index was applied to the presence/absence data of reptiles in 10×10 km UTM (Universal Transverse Mercator) squares, and then UPGMA (Unweighted Pair‐Group Method using arithmetic average) was used to classify the species. A probabilistic method was employed to assess the statistical significance of the groups obtained. An ordination method, the Canonical Correspondence Analysis, was also used to study the distribution of the reptiles within a continuous framework.
A gradual longitudinal replacement of reptile species was found throughout the Riff, with no sharp discontinuities for the distributions of most of the species. This may be due to the biogeographical northward movement of the Saharan boundaries, which have not yet reached biogeographical equilibrium. Thus, Saharan reptiles enter the Riff region from the east, through the lower basin of the River Moulouya.
Seven reptile chorotypes were identified in the Riff, and these comprise Mediterranean species and others endemic to the Maghreb (the region that spans most of North‐western Africa, excluding the Sahara). These chorotypes have a western distribution, and are segregated from one another according to altitude. Historical and ecological processes can account for the distributions shared by these species, which have inhabited the Riff for longer than eastern reptiles.     

Phytogeography of Italian deciduous oak woods based on numerical classification of plant distribution ranges

Abstract. 358 vascular plant species occurring in the deciduous oak woods of Italy were subdivided into 18 phytogeographical groups (chorotypes) through a numerical classification of their total ranges. The distribution patterns of the chorotypes are illustrated by chorograms, i.e. isopleth maps showing the joint distribution of species in Europe and surroundings regions. Except for Mediterranean species, which occur only occasionally in deciduous forests, the oak-wood flora of Italy consists of four main phytogeographical elements, which have a similar incidence (ca. 25 % each): (a) narrow-ranging, southern European species, (b) southeastern European species of submediterranean woodlands, (c) central-European species, (d) widely distributed European species. The oak woods of Italy differ from their central-European counterparts as to the relatively high floristic diversity, and in the presence of several narrow-ranging taxa, centred in different refugial areas for forest vegetation during the glacial period. There is a relation between the distribution patterns of the chorotypes and their relative frequencies in different communities; the latitudinal and longitudinal extensions of the ranges reflect the thermic and hygric conditions of the communities. The oak woods of northern and southern Italy are phytogeographically well-characterized; the former have a maximum of wide-ranging, central European species, the latter host several narrow-ranging taxa. The woodlands of central Italy have intermediate phytogeographical features, which is one of the main reasons for the difficulties in their syntaxonomical arrangement.     

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.     

Distribution patterns of the small mammals (Insectivora and Rodentia) in a transitional zone between the Eurosiberian and the Mediterranean regions

To determine, by means of quantitative analyses, the distribution patterns of the insectivores and rodents in Catalonia using physiographical regions as Operative Geographical Units (OGUs). Catalonia (north‐eastern Iberian Peninsula). Based on the presence/absence of twenty‐five small mammals in thirteen physiographical regions, which were used as OGUs, the following aspects were studied: (a) identification of biotic boundaries; (b) determination of chorological association of species; (c) climatic characterization of the biotic regions and the chorotypes. Groups of biotic regions and species were established, and their statistical significance was tested. The possible effect of several climatic factors on these groupings was studied using discriminant analyses. A significant biotic boundary was found to separate the central and eastern Pyrenees from the remaining physiographical regions. The climatic variables that defined this boundary were related to the severity and the availability of environmental energy. Four chorotypes were identified. One chorotype was constituted by Pyrenean or Pre‐Pyrenean species, an association determined by their mid‐European requirements; another chorotype was formed by Eurosiberian species, but showed a variable degree of tolerance to Mediterranean conditions; a third chorotype included species with a wide distribution that are Mediterranean, anthropic, generalist or have very specific habitats; and finally the fourth chorotype was constituted by a strictly Mediterranean species. The climatic factors that accounted for the distribution of these chorotypes were the mean July temperature and the mean annual precipitation. We conclude that the axial zone of the Pyrenees, except the coastal portion, determines two biotic regions in Catalonia. As for the classification of species, using quantitative techniques for the first time, we offer a new biogeographicalal configuration for the small mammals in this temperate‐Mediterranean transition.     

Quantitative phytogeography of the Italian Beech Forests

This paper presents a numerical analysis of the distribution patterns in Europe and adjoining regions of 242 vascular plant species occurring in the Italian beech forests. The classification of a matrix of species and of 531 Operational Geographic Units (OGUs) led to recognition of 15 different phytogeographic elements (chorotypes). The joint distribution of the species belonging to each chorotype is shown by a chorogram, which is a geographic map obtained by computer processing of the distributional data. Some of the chorograms show the main distributional centers for the beechwood flora in southern Europe: northwestern Balkan Peninsula and eastern Alps, southern Balkan Peninsula, Maritime Alps, northern Apennines, southern Apennines. Only 20% of the species are limited to narrow areas in southern Europe, and were not able to expand considerably their ranges in postglacial times. Migration from the refugia and colonization of vast areas in central and northern Europe led to intensive speciation phenomena, occurring chiefly through the formation of neopolyploid species. Several south European, narrow-ranging species, can be considered as palaeopolyploids with relict character. There is a good relationship between distribution and ecology of the chorotypes.Abbreviations OGU Operational Geographic Unit - OGS Operational Geographic Set - OCS Operational Character Set     

Fuzzy chorotypes as a conceptual tool to improve insight into biogeographic patterns

Chorotypes--statistically significant groups of coincident distribution areas--constitute biogeographic units that are fuzzy by nature. This quality has been referred to in the literature but has not been analyzed in depth or methodologically developed. The present work redefines chorotypes as fuzzy sets from a pragmatic perspective and basically focuses on the methodological and interpretative implications of this approach. The amphibian fauna in the Iberian Peninsula was used as an example to explore the fuzzy nature of chorotypes. The method on which this article is based is a widely used technique to define chorotypes. This method involves the fuzziness that is inherent to the identification between degree of similarity and degree of membership and includes a probabilistic analysis of the classification for the objective delimitation of chorotypes. The main innovation of this paper is a procedure to analyze chorotypes as fuzzy biogeographic units. A set of fuzzy parameters to deal with the biogeographic interpretation of fuzzy chorotypes is also described. A computer program has been developed and is freely available. History may be related to the degree of fuzziness of chorotypes. In our example, with amphibian distributions in Iberia, less fuzzy chorotypes could have a historical explanation, and the internal fuzziness of chorotypes increases with their distance to hypothetical Pleistocene refugia.     

Using chorotypes to deconstruct biogeographical and biodiversity patterns: the case of breeding waterbirds in Europe

Aim To deconstruct the biodiversity pattern of the 152 waterbird species breeding in Europe to better understand its multiple causal processes. Location Continental Europe, Iceland and the British Isles. Methods We considered the orders that are typically comprised by swimming, diving or wading birds, which inhabit marshes, fens, peatlands and fresh, brackish or salt waters, including coastal waters. We used the 55 main river basins of Europe as geographical units, and searched for either chorotypes (groups of similar species distributions) or gradual replacement of species throughout the river basins. Chorotypes were recognized by applying a probabilistic classification method to the distributions. Then we used GLM to characterize the extent and the species richness of each chorotype according to energy availability (higher levels of environmental energy favouring the presence of species), climatic stress due to an excess of energy, availability of water, productivity, seasonality and surface area. Results One hundred and forty species significantly aggregate into nine chorotypes. The other 12 species, most of them marine, are centred on Great Britain, dropping away progressively on coasts further away from there. Differences in either the availability of energy or climatic stress significantly characterized the distribution of seven chorotypes comprising 90.8% of the species. Main conclusions Chorotypes are meaningful and useful to deconstruct biodiversity patterns. Our results suggest that energy is the main factor related to the biogeographical patterns of breeding waterbirds in Europe, and provide an insight into regional trends of species richness previously analysed with a habitat‐scale perspective.     

On identifying common distribution patterns and their causal factors: a probabilistic method applied to pteridophytes in the Iberian Peninsula

Abstract. The aim of this quantitative biogeographical analysis is to identify the existence of specific groups of pteridophytes with significantly similar distribution patterns (chorotypes) in the Iberian Peninsula, as well as to infer the environmental factors that may explain such groupings. We have applied a classification method to the pteridophyte species and have then tested the significance of the resulting groups. This allows the distinction of groups of species that occur together more frequently than expected at random from those which are just a product of the cluster analysis. The majority of the 113 pteridophyte species in the Iberian Peninsula are not distributed independently from each other, but eighteen chorotypes exist that are common to various species and three chorotypes that consist of one species each. Five species are distributed independently of the other species, and so their distribution areas overlap and segregate from the others at random. With the aim of finding a relation between the chorotypes identified and the processes that determine them, we investigated the possible incidence of certain environmental factors on the chorotype distributions by means of logistic regression. Factors that appear most frequently as possible causes of the distribution of the chorotypes are the water availability, climatic stress, availability of environmental energy, and disturbances caused by floods. Resumen . El objetivo de este análisis biogeográfico cuantitativo es identificar la posible existencia de grupos de especies de pteridófitos con patrones de distribución significantivamente similares (corotipos) en la Peninsula Ibérica, así como analizar los factores ambientales que explican tales grupos. Hemos aplicado un método de clasificatión biogeográfica de especies y después hemos comprobado la significación de los grupos resultantes. Esto nos ha permitido distinguir los grupos de especies que se asocian con más frecuencia de lo esperado por azar de aquellos grupos que se deben únicamente al análisis de agrupación. La mayoría de las 113 especies de pteridófitos en la Península Ibérica no se distribuyen independientemente unas de otras, sino que existen 18 corotipos que son comunes a varias especies y tres que están formados por una sola especie. Cinco especies se distribuyen independientemente de las restantes, y por tanto, sus áreas de distribución se solapan y se segregan con las demás al azar. Con objeto de encontrar una relación entre los corotipos identificados en el presente trabajo y los procesos que los determinan, investigamos por medio de regresiones logísticas la posible incidencia de ciertos factores ambientales sobre las distribuciones de los corotipos. Los factores que han aparecido más frecuentemente como posibles causas de las distribuciones de los corotipos son la disponibilidad de agua, el estrés climático, la energía ambiental disponible y los disturbios causados por las inundaciones.     

Biogeographical zonation of African hornbills and their biotic and geographic characterisations

We studied the geographic ranges of the 23 African hornbill species using countries as distribution units, to search for either groups of similar distributions (chorotypes) or gradual replacement of species as two different types of distribution patterns. We used a probabilistic classification method to distinguish between these two types of biogeographic patterns. Then we analysed the influence of climate, area, surface covered by different types of habitat, and human disturbance in each country, as well as biotic features of the species involving the habitat preference, feeding requirements and habits, body size, sexual dimorphism, and metabolic parameters, on the biogeographic patterns detected. We found that 20 species were significantly classified into three chorotypes. The western-central chorotype comprises nine species characterised by their preference for forest habitat and their low sexual body mass dimorphism. These species occur in 25 countries, characterised by their low range of temperatures. The eastern chorotype includes three species characterised by their preference for feeding on the ground and greater dimorphism in mass between the sexes. Seven countries, characterised by their high values of savanna and grassland surface area, constitute the geographic element of this chorotype. The southern chorotype involves the distribution of eight species characterised by their preference for montane and coastal forests or woodlands. Eighteen countries fall into this chorotype and are characterised by their low values of temperature in the coldest month and greater surface area of desert scrub and desert forest. The other three species replace each other mainly following a gradual pattern within the study area.     

Revision of the plant geographical territories of Israel and Sinai

The phytogeographical territories of Israel and Sinai are revised using a large mass of species distribution data. With the help of chorotype frequencies for 25 km² individual squares in Israel, territories were defined on the basis of the first two most frequent chorotypes. The similarity of boundaries in the resulting phytogeographical map with a physiognomic map suggests that the latter may be used in preliminary stages of the preparation of plant geographical maps.     

Scrub communities along a climatic gradient in the southern Balkans: maquis,pseudomaquis and shibljak

The research deals with three scrub communities/formations in the southern Balkans, maquis, pseudomaquis and shibljak, and their changes along the main (macro)ecological gradient. We performed DCA analysis and, since the first axis correlates with climatic data, the projection of relevés on this axis was accepted as proxy for the (macro)ecological gradient, and the species turnover along this gradient was elaborated. Linear regression was used to test the turnover of life forms, chorotypes and the life strategy of species (syntaxonomical affiliation). It was found that life forms, except hemicryptophytes and nanophanerophytes, do not change significantly along the gradient change; most chorotypes change significantly; among species, life strategies (syntaxonomical affiliation), the proportion of species of deciduous and evergreen forests and scrub, as well as species of dry grassland, change significantly. The classification established three clusters, representing three formations and we tested differences among them. It was established that stenomediterranean and Eurasian species and species of perennial grasslands differentiate all three clusters and some groups only two of them. We found that discontinuous formations can be established along a continuous gradient. Communities were also classified within the present synsystematic system.     

Biogeographical analysis of the flea beetle genus Chaetocnema in the Afrotropical Region: distribution patterns and areas of endemism

Aim The areal distributions of Chaetocnema species in the Afrotropical Region have been analysed with the aims of determining the distribution patterns (chorotypes) and identifying the most important areas of endemism for this flea beetle genus in sub‐Saharan Africa. Location Data were collected in sub‐Saharan Africa, including Madagascar. Methods The Afrotropical Region was divided into 103 5° quadrats (operative geographical units, or OGUs). A presence–absence matrix of the Afrotropical Chaetocnema species in the OGUs was analysed by cluster analysis (Baroni Urbani & Buser index and the WPGMA clustering method) to generate distribution pattern data based on similarity of distribution. The most important areas of endemism were identified by parsimony analysis of endemicity. Results The general distribution of Chaetocnema in the Afrotropical Region was found to be associated with moist environments and montane grasslands. Most species exhibit restricted geographical ranges. Cluster analysis revealed 120 spatial distributions that can be grouped into 13 distinct distribution patterns (chorotypes). The most important areas of endemism for Chaetocnema in sub‐Saharan Africa according to the present parsimony analysis of endemicity are: (1) central and eastern Madagascar [endemicity rate (ER) = 61.1%], (2) Western Cape Province (ER = 36.4%), (3) southern Drakensberg (ER = 26.7%), (4) the Shaba Region (ER = 16.7%), and (5) the North‐Kivu Region (ER = 5.0%). Main conclusions There are 123 known species of Chaetocnema in the Afrotropical Region, more than in any other zoogeographical region. About 91% of the species are endemic and they generally exhibit a restricted and often very localized geographical range. The remaining 9% of the species are represented by seven species that also inhabit northern Africa and/or the Arabian peninsula (C. bilunulata Demaison, C. ganganensis Bechyné, C. ljuba Bechyné, C. pulla Chapuis, C. tarsalis Wollaston, and C. wollastoni Baly), three species that widely inhabit the Palaearctic Region (C. conducta (Motschulsky), C. schlaeflini (Stierlin), and C. tibialis (Illiger)), and two species that were introduced (C. confinis Crotch, and C. picipes Stephens).     

On the concept of chorotype

Recent reviews of the meaning of the word ‘chorotype’ in biogeography have led to contrasting definitions and a confusion of concepts. This is because ‘chorotype’ has been used by different authors to express two different concepts: (1) groups of species with overlapping ranges (overall distributions) and (2) groups of species with a similar distribution within a certain area. To avoid confusion, I suggest the term ‘global chorotype’ be used to indicate a group into which species with similar ranges can be classified; and ‘regional chorotype’ be used for a group of species with similar distributions within a certain region. Although the global chorotype represents the world‐wide spatial responses of species to historical and environmental pressures, and does not vary with the area under consideration, a particular species might be classified into different regional chorotypes in different study areas.     

Distribution Patterns of Indigenous Freshwater Fishes in the Tagus River Basin, Spain

Classification and ordination methods used to examine the internal complexity of the Mediterranean Tagus River catchment based on fish distribution revealed that it is not a homogeneous biogeographical unit. The indigenous fishes analyzed in this study are distributed through the basin forming geographical communities (chorotypes), some of which are associated with environmental factors like river morphology, water quality or geographical location. Nevertheless, 40% of the variation in species occurrence remains unexplained by either environmental or geographical variables, suggesting that historical factors may influence the freshwater fish distribution patterns. Three main biogeographical areas, delimited by significant boundaries, were identified. Two of them are identified as the upper and the middle-lower basins of the Tagus River catchment; the third corresponds to the Alagón River and seems to be linked to historical factors of the catchment.     

Photosynthetic pathways,distribution, and ecological characteristics of grass species in egypt

Summary This study is based both on our own results, especially δ¹³C analyses, and on data in the literature. Of the 225 species of Poaceae reported from Egypt, 105 species show the C₃ and 120 the C₄ type of photosynthetic CO₂ fixation. Winter annual and perennial grasses active in winter are mainly C₃ species, while summer annuals and the other perennials are mainly C₄ species. The percentage of C₃ species decreases with decreasing latitude. This seems to be mainly related to increasing temperatures. The C₃ grass species are found preferentially in the Mediterranean, Irano-Turanian, Mediterranean/Irano-Turanian, and Saharo-Arabian chorotypes, while the C₄ species are mainly found in the Sudanian, Saharo-Arabian/Sudanian and Tropical chorotypes. In Egypt, the NADP⁺-ME and NAD⁺-ME subtypes of C₄ photosynthesis are found in about equal numbers of C₄ species, while the PCK subtype is relatively rare.