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Invasiveness of alien vascular plants in six arid zones of Europe, Africa and America.

INTRODUCTION

Biological invasions currently constitute one of the most important and striking aspects of global environmental change (Thuiller & al., 2007; Vila & al., 2006; Wittemberg & Cock, 2001). The consequences of these invasions are felt in many ways. For example, their effects are seen in the progressive loss of biodiversity, the alteration of biogeochemical cycles, the negative interference with certain human activities and interests, and, in short, in the homogenization of Earth's biota (Atkinson & Cameron, 1993). Future global biodiversity scenarios highlight potentially dramatic increases in biological invasions in ecosystems (SALA & al., 2000). Once an invasive alien species is established in a territory and the process of invasion has been triggered, the measures needed to control or eradicate it are very expensive and not always effective (Pimentel & al., 2001). These measures can also cause collateral damage in the environment. For these reasons, preventative action is more useful. Consequently, those studies which are designed to discover the complex mechanisms and patterns which bring about and favour invasion processes can produce predicative models. These models then enable us to identify and evaluate situations where there is a risk.

Two approaches to the study of the phenomenon are (1) the identification of the attributes of the species which enable their success in the process of invasion (invasiveness), and (2) the analysis of the conditions of the habitats that make them more vulnerable to invasion by alien plants (invasibility). Environments and regions with extreme climatic conditions and little human influence seem to be those which have less capacity to receive alien species (Sanz Elorza & al., 2006; GASSO & al, 2009). Mountainous areas, the Polar regions, the cold continental steppes and desert and arid areas are examples of these cases. In contrast, those coastal regions which are in mid-latitude countries and which have a high population density are those with the highest numbers of alien flora, both total and invasory (Sobrino & al., 2002). In the case of the arid regions, the study of the phenomenon is especially interesting for various reasons. Although these regions do not have lush vegetation, they do amass a rich flora with numerous endemic species adapted to survive in very concrete and hostile environments. Examples of such areas would be places with scarce and unevenly spread rainfall, areas with high temperatures or wide fluctuations in temperature, or regions with very low relative humidity or drying winds. Other examples would be regions with intense rates of sunshine, severe night radiation frosts, high evapotranspiration values or saline ground. These factors cause species to develop highly specialized adaptations. Such mechanisms could include a reduction in the transpiration surface, a shortening of life cycles, the development of tissues to store water or CAM and C-4 metabolisms. The phenomenon of biological invasions in arid zones has not been widely studied, at least on a global scale, unlike other in habitats and countries where information on the presence and distribution of alien plant species can be found in a variety of published and unpublished accounts and databases (Meyerson & Mooney, 2007). However, if the predictions for the future of the climate come true, the extension and distribution of arid zones in the world are going to undergo changes which will require us to take into account the study of all the phenomena and biological processes which are related to them.

The objectives of this study are (1) to determine which life-history traits are related to the invasiveness of alien plants in arid zones; (2) to clarify the differences in the characteristics of the alien flora in several representative arid regions of the world; and (3) to help develop explicit invasion risk protocols for these zones.

MATERIAL AND METHODS

In order to carry out the study, we have chosen six representative arid zones from around the world. These are the Mojave and Sonoran Deserts in North America, the Atacama and Peruvian Deserts in South America, the Sahara in the north of Africa and the regions of Tabernas y Monegros (Spain) in southeast Europe (Figure 1). The Mojave Desert stretches through southwest California. Its climate is characterised by seasonal and daily temperature fluctuations and, on occasions, it suffers from long periods of frost. It includes extremely arid areas, like Death Valley, with average annual mean precipitation of less than 50 mm. The Sonoran Desert extends from the south of California to the south of the Mojave Desert, continuing through southwest Arizona and the northwest of Mexico. Its pattern of temperatures is less cold, although records show an average absolute minimum temperature of--9.5[degrees]C (Aguajita/Quitobaquito) and an average absolute maximum of 48.9 [degrees]C (Sierra del Rosario). The annual mean precipitation ranges from 40.22 mm in El Riito to 267.0 mm in Aguajita/Quitobaquito (Felger, 2000). The Atacama and Peruvian Deserts include some of the driest places on earth, with practically no rainfall, (less than 1 mm of average annual rainfall in Arica and Iquique), although at their southern limit (Copiapo) there was 64 mm of annual mean precipitation (Mcginnies, 1977). The Sahara is the largest desert in the world. It extends from the Atlantic coast of Africa to the Red Sea. It also includes some of the warmest and driest areas in the world. For example, the Great Tanezrouft has an annual mean precipitation of less than 25 mm and in El Azizia, in Libya, the highest temperature in the world was recorded (58[degrees]C in the shade). The Tabernas Desert extends through southeast Spain and covers an area of approximately 360 [km.sup.2]. The annual mean precipitation of most of this area ranges from 180 to 250 mm. The maximum temperatures in the hottest months are above 34[degrees]C and the minimum temperatures in the coldest months are rather mild with only light frosts. The Monegros Desert is a wide continental steppe area that stretches through the north-eastern part of the Iberian Peninsula in Spain, and covers an area of approximately 2,760 [km.sup.2]. Its climate has some extreme temperatures, with maximum summer temperatures that can reach over 40[degrees]C, and minimum winter temperatures that can be as low as --12[degrees]C. The annual mean precipitation ranges from 300 to 420 mm, although it is very irregularly spread out over the year.

[FIGURE 1 OMITTED]

For each area, we have catalogued the alien vascular flora from the following sources: Mojave (Baldwin & al., 2002); Sonoran Desert (Felger, 2000; Nabhan & al., 2002); Sahara (Ozenda, 1991; Quezel & Santa, 1962-1963; Maire, 1952-1980); Atacama and Peruvian Deserts (Dillon, 2005); Tabernas (Kunkel, 1988; Sagredo, 1987; Dana & al., 2001); Monegros (Molero & Blanche, 1998; Sanz Elorza, 2006). We have only taken into account naturalized species, according to the criteria of Richardson & al. (2000), and have not included casuals or sub-spontaneous species. From the information obtained from the cited sources, we considered that a species shows an invasive characteristic when in at least one of the regions studied there are reasons to think that it fulfils the criteria established by Richardson & al. (2000) and Pysek & al. (2004) relating to the term invader when applied to vascular plants. In short, the aforementioned authors define an invasive plant as an alien species with self-sustainable populations without direct human intervention that produces offspring at considerable distances from the parent plants, and thus has the potential to spread over large areas. Invasiveness is quantified as a continuous numeric variable (number of regions where it shows an invasive characteristic).

Once the list of alien naturalized plants in the six arid zones studied had been compiled, a database was created with eight biological and ecological plant traits, according to the scientific literature (Pysek & al., 1995; Goodwin & al., 1999; Lloret & al., 2003; Sanz Elorza & al., 2006; Gasso & al., 2009). These were the traits that could potentially be related to the invasion success of alien species. These variables are clonality (yes, no), pollination model (wind, insects, water), dispersal syndrome (anemochorous, hydrochorous, zoochorous and others that include barochorous, balistochorous, etc.), linking to moist soils (yes, no), seed size (very small if < 1 mm, small between 1-3 mm, medium between 3-5 mm, large between 5-10 mm and very large if > 10 mm), linking to disturbed habitats (yes, no), salinity tolerance (yes, no) and longevity (annual, perennial). We used a test [chi square] in order to determine if significant differences exist between the results obtained in each of the areas.

For statistical analysis, we used Generalized Linear Mixed Models (GLMM) to model the species invasion success (invasiveness) according to the eight predictors previously described. Typical GLM can estimate and test any univariate or multivariate general linear model, including those for multiple regression, analysis of variance or covariance, and other procedures such as discriminant analysis and principal components. The model is:

Y = XB + e

where y is a vector or matrix of dependent variables (they should be continuous numeric variables), x is a vector o matrix of independent variables (they may be either continuous or categorical dummy variables, according to the type of model), b is a vector or matrix of regression coefficients, and e is a vector or matrix of random errors (Searle, 1971; Winer, 1971; Neter & al., 1985; Cohen & Cohen, 1983).

After the parameters of the model have been estimated, they can be tested by any general linear hypothesis (in our model, life-history traits are related to the invasiveness of alien plants in arid zones) in the following form:

ABC' = D

where a is a matrix of linear weights on coefficients across the independent variables (the rows of b), C is a matrix of linear weights on the coefficients across dependent variables (the columns of b). In our model, c is an identity matrix. b is the matrix of regression coefficients of effects, and d is a null hypothesis matrix (null matrix). However, there is a clear bias as far as both the invasive characteristics of the species and their biological and ecological attributes are concerned. This is due to the phylogenetic relationship, and is defined as taxonomic bias. This causes an erroneous increase in the number of degrees of freedom as each species is used as an independent piece of data (Brandle & al., 2003), with the resulting increase of type-I errors. GLMM enables the phylogenetic effect to be mitigated by means of the incorporation of the taxonomic categories, in our case the genus, as nested random factors. This partly controls the probable lack of independence of the values assumed by the variables in species that are phylogenetically close (Gasso & al., 2009).

RESULTS

Species approach

The inventory of the alien flora of the areas studied, extracted from bibliographical sources, lists a total of 347 species although there are notable qualitative and quantitative differences as far as the location of each area is concerned (Table 1).

Accounting for taxonomic relationships, of the eight selected predictors, only two are significantly related to the invasion characteristic (Table 2). These are dispersal syndrome and linking to disturbed sites. If taxonomic bias is not considered, the salinity tolerance (F-ratio = 9.097; p-value = 0.0027) also appears to be significantly related to the invasion characteristic, as halophyte species are concentrated in a few families and genera. With regard to the dispersal syndrome, the wind and rain can be seen to be the main vectors of invasive alien species (Figure 2). As far as linking to disturbed sites is concerned the invasive species are largely ruderal and nitrophilous (Figure 2).

Alien floras of arid zones

The analysis of the differences in the percentages of alien species obtained in each area for each of the predictors, with relation to the overall arid areas shows that, in general, the dry areas of the Old World are more distinct than those of the American continent (Table 3).

[FIGURE 2 OMITTED]

DISCUSSION AND CONCLUSIONS

Our approach enables us to identify the factors that favour the invasiveness of introduced species in arid zones. Until now, only a few species attributes have been shown to be significantly related to the invasiveness of alien plants (Thompson & al., 1995; Sobrino & al., 2002; Lloret & al., 2003; Sanz Elorza & al., 2006). The two predictors that we have found to be related to the phenomenon in dry areas had already been related to the invasiveness of alien species in other parts of the world, and in other ecological environments. The relation between dispersal syndrome and invasion success of the species dispersed by wind (anemochorous) in the western Mediterranean (Gasso & al., 2009) has been proved and coincides with what has been observed in dry areas. Nevertheless, in these dry areas, species dispersed by water (hydrochorous) have also shown themselves to be capable of becoming successful invaders. Obviously, in dry areas the environments where hydrochorous species can trigger invasion processes are very limited, therefore resulting in surface extension areas which are not very significant. Moreover, in specific concrete and delimited wetland habitats the application of prevention and control strategies is much more straightforward. On the other hand, species linked to moist soils have not shown themselves to be more efficient invaders in any significant way, although in areas where agriculture with irrigation is important the opposite may be true. This situation occurs in Monegros (Table 3), and is the explanation for the different behaviour of this area with respect to the general pattern. Many of the invasive alien species present are weeds of irrigated crops that come from tropical areas (Sanz Elorza & al., 2006).

Apart from dispersal syndrome and linking to disturbed sites, the rest of the traits studied have not been found to be related to invasion success. It is striking that characteristics such as seed size, capacity for clonal reproduction, linking to moist soils or longevity do not explain the phenomenon of invasiveness of alien plants in dry areas, as could have at first been assumed. This situation reinforces the theory of the absence of a general pattern of the attributes of invasive species, as these are largely dependent on the type of habitat and the geographical environment in which they are found (Thompson & al., 1995).

In the comparison of the alien flora of the six areas studied (Table 3), it can be seen that the Tabernas Desert differs significantly in all the attributes with respect to the global pattern of the arid areas. Monegros is only 700 km away, and is only significantly different with respect to linking to moist soils due to the importance of irrigated agriculture. The attributes of the alien flora of Monegros seem to bear a closer resemblance to the alien flora of the deserts of North America, where the arid conditions are more similar and not as extreme as those of the Sahara or the Atacama and peruvian Deserts. These last two, which are the most genuine deserts, show significant differences in biological types, with a greater presence of therophytes and an almost total absence of phanerophytes and chamaephytes. This is easily explained if we consider the conditions of extreme dryness of these deserts and the absence of wetlands and fluvial courses that could mitigate the situation on a local scale. In these conditions, the therophytes are the best adapted to survive, as they take advantage of any sign of temporary humidity that allows them to complete their cycle quickly. In the alien flora of the Sahara, significant differences can be observed in the global pattern of seed size, as the species with small seeds acquire greater quantitative importance. Many of the alien species in the Sahara are, in fact, therophytes of Mediterranean origin with small seed size. The Sonoran Desert does not significantly differ from the general pattern in any of the traits we considered. This is partly because this zone had the greatest number of alien species of all the arid zones in the study. This could be because there are more environments which are altered by human influence, as there are several cities with large populations in this desert, unlike in the other five areas. In the Mojave Desert, the significant difference which can be seen with respect to dispersal syndrome is worth noting. This is caused by the greater frequency of alien species with anemochorous dispersion.

We can conclude that in dry areas, dispersal syndrome is a determining factor of invasiveness. This agrees with the descriptions in the classic literature about the ecology of invasive alien plants (Richardson & al., 2000; Kolar & Lodge, 2001; Prinzing & al, 2002). Moreover, the harsh environmental conditions of the arid zones create an ecological barrier that is difficult for alien plants to overcome, as they tend to be concentrated in environments which are altered by human activity. For this reason, the best preventative measure against invasive alien plants has been shown to be making the preservation of the habitats of arid zones a priority in nature conservation policies. The differences in the attributes of the alien species in the six areas studied show that, in spite of the fact that they all have an arid climate in common, there are more subtle environmental factors in play which cause deviations with respect to a hypothetical general situation. Therefore, the theory of the absence of a general pattern in the attributes of alien species is proved to be true. These species are largely dependent on local conditions, even in the case of regions with major climatic analogies.

doi:10.5209/rev_LAZA.2010.v31.7

Recibido: 22 febrero 2009

Aceptado: 25 noviembre 2010

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Mario Sanz Elorza (*), Francisco Gonzalez Bernardo (**), Alfredo Serreta Olivan (***) & Laura P. Gavilan Iglesias (****)

* Gerencia Territorial del Catastro. C/ Ildefonso Rodriguez 1. E-40001 Segovia. E-mail: mario.sanz@segovia.catastro.meh.es ** Area de Ingenieria Cartografica, Geodesica y Fotogrametria. Escuela Superior y Tecnica de Ingenieria Agraria. Universidad de Leon. Avda. Portugal 41. E-24071 Leon. E-mail: francisco@agrotecnos.es *** Departamento de Ingenieria de Diseno y Fabricacion. Escuela Politecnica Superior. Universidad de Zaragoza. Carretera de Cuarte s/n. E-22071 Huesca. E-mail: serreta@unizar.es **** Departamento de Botanica. Universidad de Salamanca. Avda. Licenciado Mendez Nieto s/n, E-37007 Salamanca. E-mail: lpgavilan@usal.es
Table 1

Alien flora of the six arid areas studied (+ presence, --absence)

Family             Species

Agavaceae          Agave americana
Agavaceae          Agave fourcroydes
Agavaceae          Agave sisalana
Aizoaceae          Carpobrotus chilensis
Aizoaceae          Mesembryanthemum crystallinum
Aizoaceae          Mesembryanthemum nodiflorum
Alismataceae       Sagittaria montevidensis
Amaranthaceae      Amaranthus albus
Amaranthaceae      Amaranthus blitoides
Amaranthaceae      Amaranthus blitum
Amaranthaceae      Amaranthus deflexus
Amaranthaceae      Amaranthus graecizans
Amaranthaceae      Amaranthus muricatus
Amaranthaceae      Amaranthus retroflexus
Amaranthaceae      Amaranthus spinosus
Amaranthaceae      Amaranthus viridis
Anacardiaceae      Schinus molle
Anacardiaceae      Rhus lancea
Apocynaceae        Catharanthus roseus
Apocynaceae        Nerium oleander
Asclepiadaceae     Cryptostegia grandiflora
Araceae            Pistia stratiotes
Cactaceae          Cylindropuntia spinosior
Cactaceae          Cylindropuntia tunicata
Cactaceae          Opuntia ficus-indica
Cactaceae          Opuntia lindheimeri var. linguiformis
Cactaceae          Opuntia monacantha
Capparaceae        Cleome viscosa
Caryophyllaceae    Agrostemma githago
Caryophyllaceae    Cerastium glomeratum
Caryophyllaceae    Gypsophila paniculata
Caryophyllaceae    Herniaria cinerea
Caryophyllaceae    Sagina apetala
Caryophyllaceae    Silene conoidea
Caryophyllaceae    Silene nocturna
Caryophyllaceae    Silene rubella
Caryophyllaceae    Silene succulenta
Caryophyllaceae    Spergularia marina
Caryophyllaceae    Stellaria media
Caryophyllaceae    Vaccaria hispanica
Compositae         Artemisia biennis
Compositae         Artemisia verlotiorum
Compositae         Aster squamatus
Compositae         Bidens bipinnata
Compositae         Bidens frondosa
Compositae         Carduus nutans
Compositae         Carthamus lanatus
Compositae         Carthamus tinctorius
Compositae         Acroptilon repens
Compositae         Centaurea calcitrapa
Compositae         Centaurea melitensis
Compositae         Centaurea solstitialis
Compositae         Cichorium intybus
Compositae         Cnicus benedictus
Compositae         Conyza bonariensis
Compositae         Conyza canadensis
Compositae         Conyza sumatrensis
Compositae         Cotula australis
Compositae         Cotula coronopifolia
Compositae         Chrysanthemum coronarium
Compositae         Dimorphoteca sinuata
Compositae         Eclipta prostrata
Compositae         Euryops subcarnosus
Compositae         Flaveria trinervia
Compositae         Galinsoga ciliata
Compositae         Gnaphalium luteo-album
Compositae         Grindelia squarrosa
Compositae         Hedypnois cretica
Compositae         Helianthus annuus
Compositae         Helianthus ciliaris
Compositae         Lactuca serriola
Compositae         Chamomilla suaveolens
Compositae         Pentzia incana
Compositae         Pulicaria paludosa
Compositae         Senecio vulgaris
Compositae         Sonchus asper
Compositae         Sonchus oleraceus
Compositae         Sonchus tenerrimus
Compositae         Silybum marianum
Compositae         Taraxacum officinale
Compositae         Tragopogon porrifolius
Compositae         Verbesina encelioides
Compositae         Xanthium spinosum
Compositae         Xanthium strumarium
Convolvulaceae     Convolvulus arvensis
Convolvulaceae     Ipomoea x leucantha
Convolvulaceae     Ipomoea triloba
Cruciferae         Brassica campestris
Cruciferae         Brassica napus
Cruciferae         Brassica nigra
Cruciferae         Brassica tournefortii
Cruciferae         Cakile maritima
Cruciferae         Camelina microcarpa
Cruciferae         Cardaria draba
Cruciferae         Chorispora tenella
Cruciferae         Conringia orientalis
Cruciferae         Coronopus didymus
Cruciferae         Coronopus squamatus
Cruciferae         Descurainia sophia
Cruciferae         Diplotaxis erucoides
Cruciferae         Diplotaxis virgata
Cruciferae         Draba reptans
Cruciferae         Eruca sativa
Cruciferae         Erysimum cheiri
Cruciferae         Hirschfeldia incana
Cruciferae         Lepidium campestre
Cruciferae         Lepidium perfoliatum
Cruciferae         Malcolmia africana
Cruciferae         Raphanus raphanistrum
Cruciferae         Sisymbrium altissimum
Cruciferae         Sisymbrium erysimoides
Cruciferae         Sisymbrium irio
Cruciferae         Sisymbrium orientale
Cucurbitaceae      Citrullus colocynthis
Cucurbitaceae      Citrullus lanatus
Cucurbitaceae      Cucumis melo
Cucurbitaceae      Cucumis myriocarpus
Cymodoceaceae      Halodule wrightii
Cyperaceae         Cyperus involucratus
Cyperaceae         Cyperus iria
Cyperaceae         Cyperus rotundus
Cyperaceae         Kyllinga brevifolia
Chenopodiaceae     Atriplex rosea
Chenopodiaceae     Atriplex semibaccata
Chenopodiaceae     Atriplex suberecta
Chenopodiaceae     Atriplex wrightii
Chenopodiaceae     Bassia hyssopifolia
Chenopodiaceae     Bassia scoparia
Chenopodiaceae     Chenopodium album
Chenopodiaceae     Chenopodium ambrosioides
Chenopodiaceae     Chenopodium berlandieri
Chenopodiaceae     Chenopodium foliosum
Chenopodiaceae     Chenopodium murale
Chenopodiaceae     Chenopodium vulvaria
Chenopodiaceae     Cycloloma atriplicifolium
Chenopodiaceae     Halogeton sativus
Chenopodiaceae     Salsola tragus
Chenopodiaceae     Salsola paulsenii
Elaeagnaceae       Elaeagnus angustifolia
Elatinaceae        Elatine chilensis
Euphorbiaceae      Chamaesyce nutans
Euphorbiaceae      Chamaesyce maculata
Euphorbiaceae      Chamaesyce prostrata
Euphorbiaceae      Chamaesyce serpens
Euphorbiaceae      Euphorbia hyssopifolia
Euphorbiaceae      Euphorbia peplus
Euphorbiaceae      Ricinus communis
Geraniaceae        Erodium cicutarium
Geraniaceae        Erodium malacoides
Geraniaceae        Erodium moschatum
Gramineae          Agrostis stolonifera
Gramineae          Apera interrupta
Gramineae          Arundo donax
Gramineae          Avena barbata
Gramineae          Avena fatua
Gramineae          Avena sterilis
Gramineae          Briza minor
Gramineae          Bromus arenarius
Gramineae          Bromus diandrus
Gramineae          Bromus hordeaceus
Gramineae          Bromus rubens
Gramineae          Bromus madritensis
Gramineae          Bromus tectorum
Gramineae          Bromus trinii
Gramineae          Bromus willdenowii
Gramineae          Cenchrus echinatus
Gramineae          Cenchrus incertus
Gramineae          Cynodon dactylon
Gramineae          Cynodon transvalensis
Gramineae          Chloris gayana
Gramineae          Chloris virgata
Gramineae          Dactyloctenium aegyptium
Gramineae          Digitaria bicornis
Gramineae          Digitaria ciliaris
Gramineae          Digitaria sanguinalis
Gramineae          Echinochloa colona
Gramineae          Echinochloa crus-galli
Gramineae          Echinochloa hispidula
Gramineae          Echinochloa oryzicola
Gramineae          Echinochloa oryzoides
Gramineae          Eleusine indica
Gramineae          Elymus hispidus
Gramineae          Eragrostis barrelieri
Gramineae          Eragrostis cilianensis
Gramineae          Eragrostis curvula
Gramineae          Eragrostis lehmanniana
Gramineae          Eragrostis minor
Gramineae          Eragrostis viscosa
Gramineae          Eriochloa acuminata
Gramineae          Eriochloa contracta
Gramineae          Gastridium ventricosum
Gramineae          Holcus lanatus
Gramineae          Hordeum distichon
Gramineae          Hordeum murinum
Gramineae          Lamarckia aurea
Gramineae          Lolium perenne
Gramineae          Lolium temulentum
Gramineae          Oryza sativa
Gramineae          Panicum antidotale
Gramineae          Panicum capillare
Gramineae          Panicum dichotomiflorum
Gramineae          Panicum miliaceum
Gramineae          Panicum repens
Gramineae          Paspalum dilatatum
Gramineae          Paspalum paspalodes
Gramineae          Pennisetum ciliare
Gramineae          Pennisetum setaceum
Gramineae          Pennisetum villosum
Gramineae          Phalaris canariensis
Gramineae          Phalaris caroliniana
Gramineae          Phalaris minor
Gramineae          Phalaris paradoxa
Gramineae          Poa annua
Gramineae          Polypogon monspeliensis
Gramineae          Polypogon viridis
Gramineae          Saccharum ravennae
Gramineae          Schismus arabicus
Gramineae          Schismus barbatus
Gramineae          Secale cereale
Gramineae          Setaria pumila
Gramineae          Setaria verticillata
Gramineae          Setaria viridis
Gramineae          Sorghum bicolor
Gramineae          Sorghum halepense
Gramineae          Tragus racemosus
Gramineae          Triticum aestivum
Gramineae          Vulpia bromoides
Gramineae          Vulpia myuros
Hydrocharitaceae   Hydrilla verticillata
Hydrocharitaceae   Egeria densa
Iridaceae          Iris germanica
Juncaceae          Juncus acutus
Labiatae           Lamium amplexicaule
Labiatae           Marrubium vulgare
Labiatae           Mentha spicata
Labiatae           Salvia sclarea
Labiatae           Stachys arvensis
Leguminosae        Acacia farnesiana
Leguminosae        Acacia saligna
Leguminosae        Alhagi pseudalhagi
Leguminosae        Caesalpinia gilliesii
Leguminosae        Ceratonia siliqua
Leguminosae        Indigofera suffruticosa
Leguminosae        Leucaena leucocephala
Leguminosae        Medicago lupulina
Leguminosae        Medicago polymorpha
Leguminosae        Medicago sativa
Leguminosae        Melilotus alba
Leguminosae        Melilotus indicus
Leguminosae        Parkinsonia aculeata
Leguminosae        Pediomelum rhombifolium
Leguminosae        Robinia pseudoacacia
Leguminosae        Trifolium repens
Leguminosae        Vicia villosa
Liliaceae          Gasteraloe x beguinii
Liliaceae          Aloe vera
Liliaceae          Asphodelus fistulosus
Lythraceae         Ammannia coccinea
Lythraceae         Ammannia robusta
Malvaceae          Malva neglecta
Malvaceae          Malva parviflora
Malvaceae          Malva sylvestris
Malvaceae          Sida spinosa
Meliaceae          Melia azedarach
Molluginaceae      Glinus lotoides
Molluginaceae      Glinus radiatus
Molluginaceae      Mollugo cerviana
Moraceae           Ficus carica
Moraceae           Maclura pomifera
Myrtaceae          Eucalyptus camaldulensis
Nyctaginaceae      Boerhavia diffusa
Nyctaginaceae      Mirabilis jalapa
Onagraceae         Oenothera drummondii
Onagraceae         Oenothera laciniata
Oxalidaceae        Oxalis corniculata
Oxalidaceae        Oxalis pes-caprae
Palmae             Phoenix dactylifera
Papaveraceae       Argemone ochroleuca
Papaveraceae       Glaucium corniculatum
Papaveraceae       Fumaria capreolata
Papaveraceae       Fumaria officinalis
Papaveraceae       Fumaria parviflora
Papaveraceae       Papaver dubium
Papaveraceae       Papaver somniferum
Pedaliaceae        Sesamum orientale
Pinaceae           Pinus halepensis
Plantaginaceae     Plantago lanceolata
Plantaginaceae     Plantago major
Polygonaceae       Polygonum argyrocoleon
Polygonaceae       Polygonum arenastrum
Polygonaceae       Polygonum aviculare
Polygonaceae       Polygonum persicaria
Polygonaceae       Rumex conglomeratus
Polygonaceae       Rumex obtusifolius
Polygonaceae       Rumex pulcher
Polygonaceae       Rumex crispus
Pontederiaceae     Eichhornia crassipes
Pontederiaceae     Heteranthera limosa
Pontederiaceae     Heteranthera reniformis
Pontederiaceae     Heteranthera rotundifolia
Portulacaceae      Portulaca oleracea
Primulaceae        Anagallis arvensis
Punicaceae         Punica granatum
Ranunculaceae      Nigella sativa
Rosaceae           Prunus dulcis
Rubiaceae          Galium aparine
Rubiaceae          Sherardia arvensis
Rutaceae           Ruta chalepensis
Salicaceae         Populus euphratica
Salviniaceae       Salvinia molesta
Scrophulariaceae   Bacopa monnieri
Scrophulariaceae   Lindernia dubia
Scrophulariaceae   Veronica anagallis-aquatica
Scrophulariaceae   Veronica arvensis
Scrophulariaceae   Veronica persica
Simaroubaceae      Ailanthus altissima
Solanaceae         Calibrachoa parviflora
Solanaceae         Datura innoxia
Solanaceae         Datura metel
Solanaceae         Datura stramonium
Solanaceae         Lycopersicon esculentum
Solanaceae         Nicotiana glauca
Solanaceae         Physalis pubescens
Solanaceae         Solanum americanum
Solanaceae         Solanum elaeagnifolium
Solanaceae         Solanum nigrum
Solanaceae         Solanum triflorum
Tamaricaceae       Tamarix aphylla
Tamaricaceae       Tamarix chinensis
Tamaricaceae       Tamarix gallica
Tamaricaceae       Tamarix parviflora
Tamaricaceae       Tamarix ramosissima
Ulmaceae           Celtis australis
Ulmaceae           Ulmus parviflora
Ulmaceae           Ulmus pumila
Umbelliferae       Ammi visnaga
Umbelliferae       Anethum graveolens
Umbelliferae       Apium graveolens
Umbelliferae       Coriandrum sativum
Umbelliferae       Cyclospermum leptophyllum
Umbelliferae       Torilis nodosa
Urticaceae         Urtica urens
Verbenaceae        Lippia nodiflora
Zygophyllaceae     Kallstroemia parviflora
Zygophyllaceae     Tribulus terrestris
Zygophyllaceae     Zygophyllum fabago

Family             Original geographic range

Agavaceae          Mexico
Agavaceae          Neotropical
Agavaceae          Neotropical
Aizoaceae          America del Sur
Aizoaceae          South Africa
Aizoaceae          Mediterranean-saharian
Alismataceae       South America
Amaranthaceae      North America
Amaranthaceae      North America
Amaranthaceae      Tropical s.l.
Amaranthaceae      South America
Amaranthaceae      Tropical s.l.
Amaranthaceae      South America
Amaranthaceae      North America
Amaranthaceae      Neotropical
Amaranthaceae      Tropical s.l.
Anacardiaceae      South America
Anacardiaceae      Africa
Apocynaceae        Madagascar
Apocynaceae        Mediterranean-saharian
Asclepiadaceae     Madagascar
Araceae            Tropical s.l.
Cactaceae          North America
Cactaceae          North America (Chihuahua, Texas)
Cactaceae          Neotropical (from Mexico to Colombia)
Cactaceae          North America (Texas)
Cactaceae          South America
Capparaceae        Sahara, Arabia and Central Africa
Caryophyllaceae    Irano-turanian
Caryophyllaceae    Eurasian
Caryophyllaceae    Eastern Europe and western Asia
Caryophyllaceae    SE Europe, N Africa and W Asia
Caryophyllaceae    Holartic
Caryophyllaceae    Mediterranean-irano-turanian
Caryophyllaceae    Mediterranean
Caryophyllaceae    Mediterranean
Caryophyllaceae    Central Mediterranean basin
Caryophyllaceae    Eurasian
Caryophyllaceae    Europe and Mediterranean
Caryophyllaceae    Irano-turanian
Compositae         Eurasian
Compositae         China
Compositae         America s.l.
Compositae         Neotropical
Compositae         North America
Compositae         Eurasian and Mediterranean
Compositae         Mediterranean basin
Compositae         Paleotropical
Compositae         C and W Asia
Compositae         Mediterranean basin
Compositae         Mediterranean basin
Compositae         Mediterranean-irano-turanian
Compositae         Eurasian and Mediterranean
Compositae         Mediterranean basin
Compositae         Neotropical
Compositae         North America
Compositae         South America
Compositae         Australia and New Zealand
Compositae         South Africa
Compositae         Southern Mediterranean basin
Compositae         Southern Africa
Compositae         Neotropical
Compositae         Southern Africa
Compositae         America s.l.
Compositae         Central and South America
Compositae         Eurasian
Compositae         From Wyoming to New Mexico
Compositae         Mediterranean Basin
Compositae         North America
Compositae         Southeastern USA and Mexico
Compositae         Europe and Mediterranean
Compositae         North America
Compositae         Southern Africa
Compositae         Europe
Compositae         Cosmopolitan
Compositae         Europe and Mediterranean
Compositae         Europe and Mediterranean
Compositae         Mediterranean
Compositae         Europe and Mediterranean
Compositae         Europe and Mediterranean
Compositae         Eastern Mediterranean basin
Compositae         North America
Compositae         South America
Compositae         North America
Convolvulaceae     Mediterranean-irano-turanian
Convolvulaceae     Neotropical
Convolvulaceae     Neotropical
Cruciferae         Eurasian
Cruciferae         Unknown
Cruciferae         Eurasian
Cruciferae         Mediterranean-irano-turanian
Cruciferae         Eurasia and Northern Africa
Cruciferae         Eurasian
Cruciferae         Europe and SW Asia
Cruciferae         Eurasian
Cruciferae         Irano-turanian
Cruciferae         Neotropical
Cruciferae         Eurasia and northern Africa
Cruciferae         Europe and Mediterranean
Cruciferae         Mediterranean
Cruciferae         Western Mediterranean basin
Cruciferae         North America
Cruciferae         Mediterranean-irano-turanian
Cruciferae         Eastern Mediterranean basin
Cruciferae         Mediterranean-irano-turanian
Cruciferae         Europe and W Asia
Cruciferae         Eastern Europe and W Asia
Cruciferae         S Europe and NW Africa
Cruciferae         S Europe, SW Asia and N Africa
Cruciferae         Eastern Europe and W Asia
Cruciferae         Southern Mediterranean basin
Cruciferae         Mediterranean-irano-turanian
Cruciferae         Mediterranean-irano-turanian
Cucurbitaceae      Saharian-arabian
Cucurbitaceae      South Africa
Cucurbitaceae      South Africa
Cucurbitaceae      Central and South Africa
Cymodoceaceae      Coastal Mexico and Caribbean
Cyperaceae         Madagascar
Cyperaceae         E Africa, W and C Asia
Cyperaceae         Eurasian
Cyperaceae         Neotropical
Chenopodiaceae     Europe, C. and SW Asia
Chenopodiaceae     Australia
Chenopodiaceae     Australia
Chenopodiaceae     North America
Chenopodiaceae     Mediterranean-irano-turanian
Chenopodiaceae     Mediterranean-irano-turanian
Chenopodiaceae     Cosmopolitan
Chenopodiaceae     Neotropical
Chenopodiaceae     North America
Chenopodiaceae     Eurasian
Chenopodiaceae     Mediterranean s.l.
Chenopodiaceae     Mediterranean s.l.
Chenopodiaceae     Central North America
Chenopodiaceae     SW Europe and Northern Africa
Chenopodiaceae     Eurasia and northern Africa
Chenopodiaceae     Eastern Europe and western Asia
Elaeagnaceae       Temperate Asia
Elatinaceae        South America
Euphorbiaceae      Caribbean
Euphorbiaceae      Eastern North America
Euphorbiaceae      Neotropical
Euphorbiaceae      Neotropical
Euphorbiaceae      Neotropical
Euphorbiaceae      Eurasia and northern Africa
Euphorbiaceae      Eastern Africa
Geraniaceae        Eurasia and northern Africa
Geraniaceae        Mediterranean-irano-turanian
Geraniaceae        Mediterranean
Gramineae          Eurasia and northern Africa
Gramineae          Eurasian
Gramineae          South and central Asia
Gramineae          Mediterranean
Gramineae          Europe
Gramineae          Mediterranean
Gramineae          Southern and western Europe
Gramineae          Australia
Gramineae          Mediterranean Basin
Gramineae          Eurasian
Gramineae          Europe and Mediterranean basin
Gramineae          Europe and Mediterranean basin
Gramineae          Eurasia and Mediterranean basin
Gramineae          Western South America
Gramineae          South America
Gramineae          America s.l.
Gramineae          Neotropical
Gramineae          Warm temp and tropical Old World
Gramineae          South Africa
Gramineae          Central and southern Africa
Gramineae          Warm temp regions worldwide
Gramineae          Mediterranean and tropical
Gramineae          Asia
Gramineae          Eurasia and N Africa
Gramineae          Eurasia and Mediterranean basin
Gramineae          Paleotropical
Gramineae          Warm temp and tropical Old World
Gramineae          Eastern Asia
Gramineae          Eastern Asia
Gramineae          Eastern Asia
Gramineae          Paleotropical
Gramineae          Eurasia and Mediterranean basin
Gramineae          Mediterranean
Gramineae          Southern Africa
Gramineae          Southern Africa
Gramineae          Southern Africa
Gramineae          Eurasia and northern Africa
Gramineae          Neotropical
Gramineae          North America
Gramineae          Central and South United States
Gramineae          Mediterranean basin s.l.
Gramineae          Eurasia and Mediterranean basin
Gramineae          Irano-turanian
Gramineae          Eurasia and Mediterranean basin
Gramineae          Mediterranean
Gramineae          Eurasia and Mediterranean basin
Gramineae          Mediterranean
Gramineae          Eastern Asia
Gramineae          India
Gramineae          North America
Gramineae          North America
Gramineae          Temperate Asia
Gramineae          Paleotropical
Gramineae          South America
Gramineae          Neotropical
Gramineae          Mediterranean and tropical
Gramineae          Eastern Africa
Gramineae          Eastern Africa
Gramineae          NW Africa and Canary islands
Gramineae          Eastern United States
Gramineae          Mediterranean-irano-turanian
Gramineae          Europe and Mediterranean basin
Gramineae          Eurasian
Gramineae          Mediterranean and paleotropical
Gramineae          Mediterranean and paleotropical
Gramineae          Eurasia and northern Africa
Gramineae          Saharian-arabian
Gramineae          Mediterranean-saharian-arabian
Gramineae          Southwestern Asia
Gramineae          Eurasia and Mediterranean basin
Gramineae          Europe and Mediterranean basin
Gramineae          Eurasian
Gramineae          Tropical Africa
Gramineae          Paleotropical
Gramineae          Paleotropical
Gramineae          Irano-turanian
Gramineae          Europe and Mediterranean basin
Gramineae          Europe and Mediterranean basin
Hydrocharitaceae   Eurasian
Hydrocharitaceae   North America
Iridaceae          Unknown
Juncaceae          Cosmopolitan
Labiatae           Eurasian
Labiatae           Europe and Mediterranean
Labiatae           Unknown
Labiatae           Irano-turanian
Labiatae           Europe and Mediterranean
Leguminosae        Neotropical
Leguminosae        Australia
Leguminosae        Western Asia
Leguminosae        South America
Leguminosae        Eastern Mediterranean basin
Leguminosae        Neotropical
Leguminosae        Neotropical
Leguminosae        Europe and Mediterranean basin
Leguminosae        Mediterranean s.l.
Leguminosae        Irano-turanian
Leguminosae        Eurasian
Leguminosae        Mediterranean-irano-turanian
Leguminosae        Neotropical
Leguminosae        North America
Leguminosae        North America
Leguminosae        Europe and Mediterranean basin
Leguminosae        Eastern Europe and western Asia
Liliaceae          South Africa
Liliaceae          Neotropical
Liliaceae          Mediterranean
Lythraceae         Neotropical
Lythraceae         Neotropical
Malvaceae          Europe, NW Africa and SW Asia
Malvaceae          Mediterranean-irano-turanian
Malvaceae          Eurasian and Mediterranean
Malvaceae          Paleotropical
Meliaceae          Eastern and southern Asia
Molluginaceae      Mediterranean and tropical
Molluginaceae      Neotropical
Molluginaceae      Mediterranean and tropical
Moraceae           Eastern Mediterranean basin
Moraceae           North America
Myrtaceae          Australia
Nyctaginaceae      Pantropical
Nyctaginaceae      Neotropical
Onagraceae         North America
Onagraceae         Eastern United States
Oxalidaceae        Old World
Oxalidaceae        South Africa
Palmae             Saharo-arabian
Papaveraceae       Western and central Mexico
Papaveraceae       Mediterranean-irano-turanian
Papaveraceae       Mediterranean s.l.
Papaveraceae       Mediterranean s.l.
Papaveraceae       Mediterranean s.l.
Papaveraceae       Eurasia and northern Africa
Papaveraceae       Eastern Mediterranean basin
Pedaliaceae        Paleotropical
Pinaceae           Mediterranean
Plantaginaceae     Eurasia and Mediterranean bassin
Plantaginaceae     Europe and Mediterranean basin
Polygonaceae       Saharian-arabian
Polygonaceae       Cosmopolitan
Polygonaceae       Cosmopolitan
Polygonaceae       Cosmopolitan
Polygonaceae       Europe, N Africa, C and W Asia
Polygonaceae       Europe and W Asia
Polygonaceae       Eurasian
Polygonaceae       Cosmopolitan
Pontederiaceae     Neotropical
Pontederiaceae     Neotropical
Pontederiaceae     Neotropical
Pontederiaceae     Neotropical
Portulacaceae      Cosmopolitan
Primulaceae        Europe, W. Asia and N. Africa
Punicaceae         Irano-turanian
Ranunculaceae      Eastern Asia
Rosaceae           Irano-turanian
Rubiaceae          Europe and Mediterranean
Rubiaceae          Eurasian and Mediterranean
Rutaceae           Mediterranean basin
Salicaceae         Central and southwesterm Asia
Salviniaceae       Brazil
Scrophulariaceae   Tropical s.l.
Scrophulariaceae   North America
Scrophulariaceae   Eurasia and northern Africa
Scrophulariaceae   Eurasian and Mediterranean
Scrophulariaceae   Southwestern Asia
Simaroubaceae      China
Solanaceae         South America
Solanaceae         Central America and Mexico
Solanaceae         Neotropical
Solanaceae         America s.l.
Solanaceae         Central and South America
Solanaceae         South America
Solanaceae         Eastern United States
Solanaceae         South America
Solanaceae         South America
Solanaceae         Eurasian and Mediterranean
Solanaceae         South America
Tamaricaceae       From Africa to India
Tamaricaceae       Eastern Asia
Tamaricaceae       Mediterranean-saharian-arabian
Tamaricaceae       Eastern Mediterranean basin
Tamaricaceae       Eastern Asia
Ulmaceae           Eastern Mediterranean basin
Ulmaceae           China and Japan
Ulmaceae           Central Asia
Umbelliferae       Mediterranean-irano-turanian
Umbelliferae       Eastern Mediterranean basin
Umbelliferae       Eurasian
Umbelliferae       North Africa and west Asia
Umbelliferae       Brazil
Umbelliferae       Mediterranean
Urticaceae         Cosmopolitan
Verbenaceae        Tropical s.l.
Zygophyllaceae     Southern USA and Mexico
Zygophyllaceae     Eurasia and northern Africa
Zygophyllaceae     Irano-turanian

                            Sonoran
Family             Mojave   Desert    Atacama

Agavaceae            --        --        -
Agavaceae            --        --        -
Agavaceae            --        --        -
Aizoaceae            --        +         -
Aizoaceae            --        +         +
Aizoaceae            --        +         -
Alismataceae         --        --        -
Amaranthaceae        --        +         -
Amaranthaceae      native   native       -
Amaranthaceae        --        --        -
Amaranthaceae        --        --        -
Amaranthaceae        --        --        -
Amaranthaceae        --        --        -
Amaranthaceae        --        --        -
Amaranthaceae        +         --        -
Amaranthaceae        --        --        +
Anacardiaceae        --        --     native
Anacardiaceae        --        +         -
Apocynaceae          --        +         -
Apocynaceae          --        +         -
Asclepiadaceae       --        +         -
Araceae              --        +         -
Cactaceae            --     native       -
Cactaceae            --        --        +
Cactaceae            +         --        -
Cactaceae            --        +         -
Cactaceae            --        --        -
Capparaceae          --        +         -
Caryophyllaceae      --        --        -
Caryophyllaceae      --        +         -
Caryophyllaceae      +         --        -
Caryophyllaceae      --        +         -
Caryophyllaceae      --        --        +
Caryophyllaceae      --        +         -
Caryophyllaceae      --        --        -
Caryophyllaceae      --        --        -
Caryophyllaceae      --        --        -
Caryophyllaceae      --        +         -
Caryophyllaceae      --        +         +
Caryophyllaceae      --        --        -
Compositae           +         --        -
Compositae           --        --        -
Compositae           --        --        -
Compositae           --        --        -
Compositae           --        --        -
Compositae           +         --        -
Compositae           +         --        -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           +         +         +
Compositae           +         +         -
Compositae           --        +         -
Compositae           +         --        -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        --        -
Compositae           --        +         +
Compositae           --        --        +
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        --     native
Compositae           +         +         -
Compositae           +         --        -
Compositae           --        +         -
Compositae           --        +         -
Compositae           +         +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         +
Compositae           --        +         -
Compositae           --        +         +
Compositae           --        +         +
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Compositae           --        +         -
Convolvulaceae       --        +         +
Convolvulaceae       --        +         -
Convolvulaceae       --        +         -
Cruciferae           --        --        +
Cruciferae           --        --        -
Cruciferae           --        +         -
Cruciferae           +         +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        --        -
Cruciferae           --        +         -
Cruciferae           +         --        -
Cruciferae           --        +         -
Cruciferae           --        --        -
Cruciferae           +         +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        +         -
Cruciferae           --        --        -
Cruciferae           +         +         +
Cruciferae           --        +         -
Cucurbitaceae        --        --        -
Cucurbitaceae        --        +         +
Cucurbitaceae        --        +         -
Cucurbitaceae        --        --        -
Cymodoceaceae        --        +         -
Cyperaceae           --        +         -
Cyperaceae           --        +         -
Cyperaceae           --        +         +
Cyperaceae           +         +         -
Chenopodiaceae       --        +         -
Chenopodiaceae       +         +         -
Chenopodiaceae       --        +         -
Chenopodiaceae       --        +         -
Chenopodiaceae       --        +         -
Chenopodiaceae       +         +         -
Chenopodiaceae       +         +         -
Chenopodiaceae       --        --        -
Chenopodiaceae     native      +         -
Chenopodiaceae       +         --        -
Chenopodiaceae       +         +         +
Chenopodiaceae       --        --        -
Chenopodiaceae       +         --        -
Chenopodiaceae       +         --        -
Chenopodiaceae       --        +         -
Chenopodiaceae       +         --        -
Elaeagnaceae         +         --        -
Elatinaceae          --        +         -
Euphorbiaceae        --        --        -
Euphorbiaceae        --        +         -
Euphorbiaceae        --        +         -
Euphorbiaceae        --        +      native
Euphorbiaceae        --        +         -
Euphorbiaceae        --        +         -
Euphorbiaceae        --        +         -
Geraniaceae          +         +         +
Geraniaceae          --        --        +
Geraniaceae          --        +         +
Gramineae            +         --        -
Gramineae            +         --        -
Gramineae            +         +         -
Gramineae            +         --        +
Gramineae            +         +         +
Gramineae            --        --        +
Gramineae            --        +         +
Gramineae            +         +         -
Gramineae            --        +         -
Gramineae            +         +         -
Gramineae            --        +         -
Gramineae            +         +         -
Gramineae            --        +         -
Gramineae            +         +         -
Gramineae            --        +      native
Gramineae            --        +      native
Gramineae            --        +         -
Gramineae            +         +         +
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            +         +      native
Gramineae            --        +         +
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            +         --        -
Gramineae            --        +         +
Gramineae            --        +         -
Gramineae            --        --        -
Gramineae            --        --        -
Gramineae            --        --        -
Gramineae            --        +         -
Gramineae            +         --        -
Gramineae            --        +         -
Gramineae            --        +         +
Gramineae            +         +         -
Gramineae            +         +         -
Gramineae            +         --        -
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            +         --        -
Gramineae            --        --        -
Gramineae            --        +         +
Gramineae            --        +         +
Gramineae            +         +         -
Gramineae            +         +         -
Gramineae            --        --        -
Gramineae            +         +         -
Gramineae            --        --        -
Gramineae            --        --        -
Gramineae            --        +         -
Gramineae            --        --        -
Gramineae            +         +         -
Gramineae            --        --        -
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            +         --        +
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            --        +         -
Gramineae            --        +         +
Gramineae            --        +         +
Gramineae            --        +         +
Gramineae            --        +         -
Gramineae            +         +         -
Gramineae            +         +         -
Gramineae            +         --        -
Gramineae            +         +         -
Gramineae            --        +         +
Gramineae            --        +         -
Gramineae            +         +         +
Gramineae            --        +         -
Gramineae            --        +         +
Gramineae            --        +         -
Gramineae            +         +         -
Gramineae            +         +         +
Hydrocharitaceae     +         +         -
Hydrocharitaceae     --        +         -
Iridaceae            --        --        -
Juncaceae            --     native       +
Labiatae             --        +         -
Labiatae             +         +         +
Labiatae             +         --        -
Labiatae             --        --        -
Labiatae             --        --        +
Leguminosae          --        --        -
Leguminosae          --        --        -
Leguminosae          +         +         -
Leguminosae          --        +         -
Leguminosae          --        +         -
Leguminosae          --        +         -
Leguminosae          --        +         -
Leguminosae          --        +         -
Leguminosae          --        +         +
Leguminosae          --        --        -
Leguminosae          --        +         -
Leguminosae          +         +         +
Leguminosae          --        +      native
Leguminosae          --        +         -
Leguminosae          --        --        -
Leguminosae          --        +         -
Leguminosae          --        --        -
Liliaceae            --        --        -
Liliaceae            --        +         -
Liliaceae            --        +         -
Lythraceae           --        --     native
Lythraceae           --     native       -
Malvaceae            --        +         -
Malvaceae            +         +         +
Malvaceae            --        --        +
Malvaceae            --        --        -
Meliaceae            --        +         -
Molluginaceae        --        +         -
Molluginaceae        --        +         -
Molluginaceae        +         +         -
Moraceae             --        --        -
Moraceae             --        +         -
Myrtaceae            --        +         -
Nyctaginaceae        --        +         +
Nyctaginaceae        --        +         -
Onagraceae           --        --        +
Onagraceae           +         --        -
Oxalidaceae          +         +         +
Oxalidaceae          --        --        +
Palmae               +         --        -
Papaveraceae         --        +         -
Papaveraceae         --        +         -
Papaveraceae         --        --        +
Papaveraceae         --        +         -
Papaveraceae         --        --        +
Papaveraceae         --        --        -
Papaveraceae         --        --        -
Pedaliaceae          --        +         -
Pinaceae             --        --        -
Plantaginaceae       --        +         +
Plantaginaceae       +         +         +
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Polygonaceae         --        +         -
Pontederiaceae       --        +         -
Pontederiaceae       --        --        -
Pontederiaceae       --        --        -
Pontederiaceae       --        --        -
Portulacaceae        --        +         +
Primulaceae          --        +         +
Punicaceae           --        --        -
Ranunculaceae        --        --        -
Rosaceae             --        --        -
Rubiaceae            --        +         +
Rubiaceae            --        +         -
Rutaceae             --        +         -
Salicaceae           --        --        -
Salviniaceae         --        +         -
Scrophulariaceae     --        +         -
Scrophulariaceae     --        --        -
Scrophulariaceae     +         +         -
Scrophulariaceae     --        +         -
Scrophulariaceae     --        +         +
Simaroubaceae        --        --        -
Solanaceae           --        +         -
Solanaceae           +         +         +
Solanaceae           --        --        -
Solanaceae           --        +         -
Solanaceae           --        --        -
Solanaceae           +         +      native
Solanaceae           +         +         -
Solanaceae           --        +         -
Solanaceae           --        +         -
Solanaceae           --        --        -
Solanaceae           +         --        -
Tamaricaceae         +         +         -
Tamaricaceae         +         +         -
Tamaricaceae         +         --        -
Tamaricaceae         +         +         -
Tamaricaceae         +         +         -
Ulmaceae             --        --        -
Ulmaceae             --        +         -
Ulmaceae             +         --        -
Umbelliferae         --        --        +
Umbelliferae         --        +         -
Umbelliferae         --        --        +
Umbelliferae         --        +         -
Umbelliferae         --        +         -
Umbelliferae         --        +         -
Urticaceae           --        +         +
Verbenaceae          --     native    native
Zygophyllaceae       +         --        -
Zygophyllaceae       --        +         -
Zygophyllaceae       +         +         -

Family             Sahara   Tabernas   Monegros

Agavaceae            --        +          -
Agavaceae            --        +          -
Agavaceae            --        +          -
Aizoaceae            --        --         -
Aizoaceae            +         +          -
Aizoaceae          native    native       -
Alismataceae         --        --         +
Amaranthaceae        --        --         +
Amaranthaceae        --        --         +
Amaranthaceae        --        --         +
Amaranthaceae        --        --         +
Amaranthaceae        --        +          +
Amaranthaceae        +         +          +
Amaranthaceae        +         --         +
Amaranthaceae        --        --         -
Amaranthaceae        +         +          +
Anacardiaceae        --        +          -
Anacardiaceae        --        --         -
Apocynaceae          --        --         -
Apocynaceae        native    native       -
Asclepiadaceae       --        --         -
Araceae              --        --         -
Cactaceae            --        --         +
Cactaceae            --        --         -
Cactaceae            +         +          -
Cactaceae            --        --         +
Cactaceae            --        --         +
Capparaceae        native      --         -
Caryophyllaceae      --        --         +
Caryophyllaceae      --      native     native
Caryophyllaceae      --        --         -
Caryophyllaceae      --      native     native
Caryophyllaceae      --        --         -
Caryophyllaceae      --        --       native
Caryophyllaceae      +       native     native
Caryophyllaceae      +       native     native
Caryophyllaceae      +         --         -
Caryophyllaceae      --      native     native
Caryophyllaceae      --        --       native
Caryophyllaceae      --        --         +
Compositae           --        --         -
Compositae           --        --         +
Compositae           --        +          +
Compositae           --        --         +
Compositae           --        --         +
Compositae           --        --       native
Compositae           --        --       native
Compositae           --        --         -
Compositae           --        --         -
Compositae           --      native     native
Compositae           --      native     native
Compositae           --        --       native
Compositae           --        --       native
Compositae           --        --       native
Compositae           --        +          +
Compositae           --        --         +
Compositae           --        --         +
Compositae           --        --         -
Compositae           --        --         -
Compositae           +       native     native
Compositae           --        --         -
Compositae           --        --         +
Compositae           --        --         -
Compositae           --        --         -
Compositae           --        --         +
Compositae           +         --         -
Compositae           --        --         -
Compositae           --      native     native
Compositae           --        --         +
Compositae           --        --         -
Compositae           --      native     native
Compositae           --        --         -
Compositae           --        --         -
Compositae           --      native       -
Compositae           --        --       native
Compositae           +         --       native
Compositae           +       native     native
Compositae           --      native     native
Compositae           --      native     native
Compositae           --        --       native
Compositae           --        --         -
Compositae           --        --         -
Compositae           --        --         +
Compositae           --        --         +
Convolvulaceae       +         --       native
Convolvulaceae       --        --         -
Convolvulaceae       --        --         -
Cruciferae           --        --         -
Cruciferae           --        --         +
Cruciferae           --        --         -
Cruciferae           +         --         -
Cruciferae           --        --         -
Cruciferae           --        --       native
Cruciferae           --      native     native
Cruciferae           --        --         -
Cruciferae           --        --         +
Cruciferae           --        --         -
Cruciferae         native      --       native
Cruciferae           --      native     native
Cruciferae           +         --       native
Cruciferae           +       native     native
Cruciferae           --        --         -
Cruciferae         native    native     native
Cruciferae           --        --         +
Cruciferae           --        --       native
Cruciferae           --        --         -
Cruciferae           --        --         -
Cruciferae         native    native     native
Cruciferae           --        --         -
Cruciferae           --        --         -
Cruciferae           +       native       -
Cruciferae           +       native     native
Cruciferae           --      native     native
Cucurbitaceae      native      +          -
Cucurbitaceae        --        --         -
Cucurbitaceae        --        --         -
Cucurbitaceae        --        --         +
Cymodoceaceae        --        --         -
Cyperaceae           --        --         -
Cyperaceae           --        --         -
Cyperaceae           +       native       -
Cyperaceae           --        --         -
Chenopodiaceae       --      native     native
Chenopodiaceae       --        +          -
Chenopodiaceae       --        --         -
Chenopodiaceae       --        --         -
Chenopodiaceae       --      native     native
Chenopodiaceae       --        --       native
Chenopodiaceae       --      native     native
Chenopodiaceae       --        +          +
Chenopodiaceae       --        --         -
Chenopodiaceae       --        --         -
Chenopodiaceae       +       native     native
Chenopodiaceae       +         --       native
Chenopodiaceae       --        --         -
Chenopodiaceae     native    native       -
Chenopodiaceae       --      native     native
Chenopodiaceae       --        --         -
Elaeagnaceae         --        --         -
Elatinaceae          --        --         -
Euphorbiaceae        --        --         +
Euphorbiaceae        --        --         -
Euphorbiaceae        --        --         +
Euphorbiaceae        --        --         -
Euphorbiaceae        --        --         -
Euphorbiaceae      native    native     native
Euphorbiaceae        +         +          -
Geraniaceae        native    native     native
Geraniaceae          --      native     native
Geraniaceae          --        --         -
Gramineae          native      --       native
Gramineae            --        --         -
Gramineae            --        +          +
Gramineae          native      --       native
Gramineae            +         --       native
Gramineae          native      --       native
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            --        --       native
Gramineae            --        --         -
Gramineae            --      native     native
Gramineae            --      native     native
Gramineae            --        --       native
Gramineae            --        --         -
Gramineae            --        --         +
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            +       native     native
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae          native      --         -
Gramineae          native      --         -
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            --        --       native
Gramineae            +         --         +
Gramineae            --        --       native
Gramineae            --        --         +
Gramineae            --        --         +
Gramineae            --        --         +
Gramineae            --        --         +
Gramineae            --        --       native
Gramineae          native      --       native
Gramineae            +         --         +
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae          native      --       native
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            --        --       native
Gramineae            --        --         +
Gramineae          native    native     native
Gramineae          native    native     native
Gramineae            --        --         -
Gramineae            --        --         -
Gramineae            --        --         +
Gramineae            --        --         -
Gramineae            --        --         +
Gramineae            --        --         +
Gramineae            --        --         +
Gramineae            +         --         -
Gramineae            --        --         -
Gramineae            --        --         +
Gramineae            --        --         -
Gramineae            +         --         -
Gramineae            --        --         -
Gramineae          native      --         +
Gramineae            --        --         -
Gramineae          native    native     native
Gramineae            --        --       native
Gramineae            --        --       native
Gramineae          native    native     native
Gramineae            --      native     native
Gramineae          native    native     native
Gramineae          native      --         -
Gramineae          native    native     native
Gramineae            --        --         -
Gramineae            --        --       native
Gramineae          native      --       native
Gramineae            --        --       native
Gramineae            --        --         -
Gramineae            --        +          +
Gramineae          native      --         -
Gramineae            --        --         +
Gramineae            --        --         -
Gramineae            --        --         -
Hydrocharitaceae     --        --         -
Hydrocharitaceae     --        --         -
Iridaceae            --        --         +
Juncaceae          native    native     native
Labiatae             --      native     native
Labiatae             --        --       native
Labiatae             --        --         -
Labiatae             --        --         +
Labiatae             --        --         -
Leguminosae          --        +          -
Leguminosae          --        +          -
Leguminosae          --        --         -
Leguminosae          --        --         -
Leguminosae          --        +          -
Leguminosae          --        --         -
Leguminosae          --        --         -
Leguminosae          --      native     native
Leguminosae          --      native       -
Leguminosae          --        --         +
Leguminosae          --        --       native
Leguminosae          +       native     native
Leguminosae          +         +          -
Leguminosae          --        --         -
Leguminosae          --        --         +
Leguminosae          --        --       native
Leguminosae          --        --         +
Liliaceae            --        --         +
Liliaceae            --        --         -
Liliaceae            --      native     native
Lythraceae           --        --         +
Lythraceae           --        --         +
Malvaceae            --      native     native
Malvaceae          native    native     native
Malvaceae            --      native     native
Malvaceae            --        --         +
Meliaceae            --        --         -
Molluginaceae        --        --         -
Molluginaceae        --        --         -
Molluginaceae      native      --         -
Moraceae             +         +          +
Moraceae             --        --         -
Myrtaceae            --        +          -
Nyctaginaceae        --        --         -
Nyctaginaceae        --        --         -
Onagraceae           --        --         -
Onagraceae           --        --         -
Oxalidaceae          --      native       -
Oxalidaceae          --        +          -
Palmae             native      +          -
Papaveraceae         --        --         -
Papaveraceae         --        --       native
Papaveraceae         --      native       -
Papaveraceae         --      native     native
Papaveraceae         --      native     native
Papaveraceae         +       native     native
Papaveraceae         --        --         +
Pedaliaceae          --        --         -
Pinaceae             --        +        native
Plantaginaceae       --        --       native
Plantaginaceae       --        --       native
Polygonaceae       native      --         -
Polygonaceae         --        --       native
Polygonaceae         --      native     native
Polygonaceae         --        --       native
Polygonaceae         --      native     native
Polygonaceae         --        --       native
Polygonaceae         --      native     native
Polygonaceae         --        --       native
Pontederiaceae       --        --         -
Pontederiaceae       --        --         +
Pontederiaceae       --        --         +
Pontederiaceae       --        --         +
Portulacaceae        +         --       native
Primulaceae          +       native     native
Punicaceae           --        +          -
Ranunculaceae        +         --         -
Rosaceae             --        --         +
Rubiaceae            --        --       native
Rubiaceae            --      native     native
Rutaceae             --      native     native
Salicaceae           +         --         -
Salviniaceae         --        --         -
Scrophulariaceae     --        --         -
Scrophulariaceae     --        --         +
Scrophulariaceae   native      --       native
Scrophulariaceae     --        --       native
Scrophulariaceae     --        --         +
Simaroubaceae        --        --         +
Solanaceae           --        --         -
Solanaceae           --        +          +
Solanaceae           +         --         -
Solanaceae           +         +          +
Solanaceae           --        +          -
Solanaceae           +         +          -
Solanaceae           --        --         -
Solanaceae           --        --         -
Solanaceae           --        --         -
Solanaceae           +       native     native
Solanaceae           --        --         -
Tamaricaceae       native      --         -
Tamaricaceae         --        --         -
Tamaricaceae       native    native     native
Tamaricaceae         --        --         -
Tamaricaceae         --        --         -
Ulmaceae             --        --         +
Ulmaceae             --        --         -
Ulmaceae             --        --         -
Umbelliferae         --        --       native
Umbelliferae         --        --         -
Umbelliferae         --        --         -
Umbelliferae         --        --         -
Umbelliferae         --        --         -
Umbelliferae         --      native     native
Urticaceae           --      native     native
Verbenaceae          +         --         -
Zygophyllaceae       --        --         -
Zygophyllaceae     native    native     native
Zygophyllaceae       +         +          +

Table 2

Tests of significance for dependent variable (invasiveness) and
predictors which are significantly related. Sigma-restricted
parameterization. Effective hypothesis decomposition.

              Sum of           Mean-
             squares    df     square    F-ratio    p-value

Intercept    425.0527    1    425.0527   1736.809   0.000000

Dispersal     2.8062     3     0.9354     3.822     0.010232
syndrome

Error        83.9431    343    0.2447

Intercept    620.3791    1    620.3791   2514.645   0.000000

Linking to    1.6356     1     1.6356     6.630     0.010446
disturbed
sites

Error        85.1137    345    0.2467

Table 3

Percentages of the number of alien species present in each area
studied and in all the areas as a whole, for each of the predictors
taken into account. The biological types are: C chamaephyte, G
geophyte, H hemicryptophyte, Hy hydrophyte, P phanerophyte, T
therophyte. The pollination modes are: An anemophilous, En
entomophilous, Hi hydrophilous. The dispersal syndromes are: An
wind (anemochorous), Hi water (hydrochorous), Zoo animals
(zoochorous), others (including barochorous, balistochorous, etc.).
The test [chi square] was used to determine if significant
differences exist between the results obtained in each of the
areas.

                                          Sonoran
                                Mojave    Desert    Atacama    Sahara

Biological       C                 3         3         2         0
type             G                4 4                  7         5
                 H                18        15        12         19
                 Hy                3         3         7         10
                 P                15        13         0         0
                 T                59        62        72         67
                 [chi square]   1.0528    0.0931    23.1296   34.0532
                 p-value        0.9582    0.9998    0.0003    < 0.0001

Clonality        no               79        82        85         74
                 yes              21        18        15         26
                 [chi square]   0.0625    0.2500    1.5625     2.2500
                 p-value        0.8026    0.6171    0.2113     0.1336

Linked to        no               51        54        48         48
moist soils      yes              49        46        52         52
                 [chi square]   0.0401    0.1603    0.6410     0.6410
                 p-value        0.8414    0.6889    0.4234     0.4234

Pollination      An               71        69        77         67
mode             En               28        30        23         33
                 Hi                1         1         0         0
                 [chi square]   1.1363    0.4091    5.8636     1.0152
                 p-value        0.5665    0.8150    0.0533     0.6019

Dispersal        An               50        38        34         29
syndrome         Hi                1         4         2         5
                 Zoo              13        19        21         12
                 others           36        38        43         55
                 [chi square]   12.1331   0.6766    2.1366     8.3881
                 p-value        0.0069    0.8787    0.5445     0.0386

Seed size        VS               25        20        18         31
                 S                39        44        51         50
                 M                14        16        10         10
                 L                14        14        13         5
                 VL               97                   8         5
                 [chi square]   2.5332    0.1436    3.1232    14.0294
                 p-value        0.6387    0.9975    0.5374     0.0072

Linked to        no               30        28        21         26
distubed sites   yes              70        72        79         76
                 [chi square]   0.0456    0.0455    3.1033     0.6625
                 p-value        0.8256    0.8257    0.0779     0.4157

Salinity         no               84        91        92         88
tolerance        yes              16         9         8         12
                 [chi square]   5.9829       --     0.1221     1.0989
                 p-value        0.0144       1      0.7268     0.2945

Longevity        annual           59        64        72         69
                 perenne          41        36        28         31
                 [chi square]   0.6864    0.0429    3.4749     1.5444
                 p-value        0.4074    0.8359    0.0623     0.2139

                                Tabernas   Monegros   Global

Biological       C                 9          3         3
type             G                 6          4         4
                 H                 13         11        45
                 Hy                0          7         3
                 P                 38         10        13
                 T                 34         65        62
                 [chi square]      --       7.2375      -
                 p-value        < 0.0001    0.2036      -

Clonality        no                66         79        80
                 yes               34         21        20
                 [chi square]   12.2500     0.0625      -
                 p-value         0.0005     0.8026      -

Linked to        no                62         44        52
moist soils      yes               38         66        48
                 [chi square]    4.0064     7.9807      -
                 p-value         0.0453     0.0047      -

Pollination      An                47         68        66
mode             En                53         32        33
                 Hi                0          0         1
                 [chi square]   18.5909     1.0909      -
                 p-value        < 0.0001    0.5796      -

Dispersal        An                19         28        35
syndrome         Hi                0          10        5
                 Zoo               25         21        19
                 others            56         42        41
                 [chi square]   19.6968     6.6349      -
                 p-value         0.0002     0.0845      -

Seed size        VS                19         17        20
                 S                 28         43        44
                 M                 19         26        15
                 L                 28         10        13
                 VL                6          4         7
                 [chi square]   24.3854    10.5174      -
                 p-value        < 0.0001    0.0326      -

Linked to        no                47         21        29
distubed sites   yes               53         79        71
                 [chi square]   15.7358     3.1083      -
                 p-value        < 0.0001    0.0779      -

Salinity         no                84         94        91
tolerance        yes               16         6         9
                 [chi square]    5.9829     1.0989      -
                 p-value         0.0144     0.2945      -

Longevity        annual            38         69        63
                 perenne           62         31        37
                 [chi square]      --       1.5444      -
                 p-value        <0.0001     0.2139      -
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Author:Sanz Elorza, Mario; Gonzalez Bernardo, Francisco; Serreta Olivan, Alfredo; Gavilan Iglesias, Laura P
Publication:Lazaroa
Article Type:Report
Date:Jan 1, 2010
Words:7764
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