Gayana

Gayana.BotanyOn-line versionISSN0717-6643

Gayana Bot.vol.68 no.2 Conception 2011

http://dx.doi.org/10.4067/S0717-66432011000200013

Gayana Bot.68 (2): 220-225, 2011

REGULAR ARTICLES

Panbiogeographic Analysis of some Cactaceae EcuadorPanbiogeographic analysis of some Cactaceae from Ecuador

Christian R. Loaiza S.1* & Juan J. Morrone21Institute of Ecology, Unit of Ecology and Plant Physiology, Technical University of Loja, Loja, Ecuador.*crloaiza@utpl.edu.ec2Museum of Zoology Alfonso L. Herrera, Department of Developmental Biology, Faculty of Science, National Autonomous University of Mexico (UNAM), Apartado Postal 70-399, 04510 Mexico, DF, Mexico.juanmorrone2001@yahoo.com.mx

ABSTRACTThe geographical distribution of 12 species of cacti included in five genera using a panbiogeographic approach in order to identify the biogeographic patterns of this group and contribute to the study of the biogeographical relationships of the flora of Ecuador was analyzed.The target species are mainly distributed in the dry forests of northern Peru and southern Ecuador.Through the analysis of stroke, three generalized tracks (dry coastal, north-central interandino interandino and South) and a node at the junction of the second and third lines level biogeographic province of western Ecuador were identified.

PALABRAS KEY:Cactaceae, panbiogeography, generalized tracks, dry forests, distribution.

ABSTRACTThe geographical distributions of 12 species of Cactaceae Assigned to five generates Were Analyzed using a panbiogeographic approach to identify identity Their Biogeographic patterns and Contribute to the study of the biogeographical relationships of the plant from Ecuador.The selected species are distributed mainly in the dry forests of northern Peru and southern Ecuador.Based on the track analysis, three generalized tracks (dry coastal, central-northern and southern interandean interandean) and one node Were Identified, the Latter in the overlap of the second and third tracks at the Biogeographic province Western Ecuador.

Keywords:Cactaceae, panbiogeography, generalized tracks, dry forests, distribution.

INTRODUCTIONThe Cactaceae is endemic to America.Its distribution includes the entire continent, with the exception of the highlands.From a dispersalist perspective, scholars have traditionally considered the dry tropical zone of South America as the probable center of origin of the family (Bravo 1978).A level of Ecuador, this family is represented by 16 genera and 43 native species (Loweet al., 2009), of which 13 species are endemic to the country, not including the Galapagos Islands (Madsen 1989, 2002, & Neill 2005 Ulloa Valenciaet al.2000).Among the areas with the highest endemism and diversity, is considered south of Ecuador (El Oro, Loja and Zamora Chinchipe).This region contains the greatest diversity of species, 13 genera (81.3%) and 28 species (58.3%) (Madsen 2002, Loweet al., 2009).Other provinces alsoare characterized by high diversity are Azuay, Canar and Chimborazo, with some equally endemic taxa asCleistocactus leonensisMadsen,Armatocereus godingianus(Britton & Rose) Backeb.E. Salisb ex.,Opuntia soederstromianaBritton & Rose andEspostoa frutescensMadsen (Britton & Rose 1920, Madsen 1989).Because of its endemic nature and the presence of certain primitive features like thorns into flowers and fruits, it is likely that genderArmatocereusandBrowningiarepresent relicts of xerophytic flora that would have been more widely distributed during the glacial period, because the conditions weather at that time were much more arid continent level (Madsen 2000).

Studies on cacti of Ecuador have been addressed mostly under a taxonomic and distributional approach without carefully analyzing biogeographic patterns and processes influencingthe current distribution of the species.Stand out from some of the most recent rankings of the flora of the dry forests of Ecuador contributions from Aguirreet al.(2006a, 2006b), who established a system of classification of vegetation based on the classification proposed by Sierra (1999 ).The proposed system Aguirre and colleagues is of great interest for the development of biogeographic studies in the dry forests of Ecuador and Peru.

Here we present an analysis panbiogeographic to contribute to the knowledge of the distribution patterns of flora cactus country.

MATERIALS AND METHODSData on the distribution of selected species (Fig. 1-8) were obtained from data collected in a previous study (Loaizaet al., 2009), which is systematized in a database collections cacti present in the major herbaria of Ecuador (QAP, QCA, QCNE and Loja), as well as some international databases herbaria, such as the University of Aarhus (AAU) and the Missouri Botanical Garden (MO), among others.The species selected were chosen based on two types of criteria: a) Known distribution in the dry forests of Ecuador, b) number of records.Species with low number of records and the distribution of which is not yet fully known was discarded.The classification proposed in this paper was properly analyzed and developed based on the results obtained in this investigation and based on the ratings of previous work (Valencia1999, Aguirreet al.2006a, 2006b, Morrone 2006, Loweet al.2009).

The panbiogeographic approach emphasizes the importance of spatial (geographic) dimension of biodiversity, to allow a better understanding of evolutionary patterns and processes, highlighting the relevance of geographical distribution as direct objects of analysis (Croizat 1958, 1964).The analysis basically consists in marking locations on maps the distribution of individual species or monophyletic groups and then record the locations of the species are joined by smaller distance lines, which are called individual strokes (Morrone & Crisci 1990, Espinosaet al .2002, Zunino & Zullini 2003, Morrone 2004).After obtaining the individual traces are superimposed and when multiple match is considered a generalized track, which allows maintaining the existence of an ancestral biota widely distributed and fragmented by a vicariant event.When two or more generalized tracks together in the same area, it is considered a node or composite area, which in turn implies a multiple biotic origin, possibly at timesdifferent geological.Nodes are areas of high biodiversity, which mainly due to its biological mixture are priorities for conservation (Espinosaet al.2002 2004 Morrone, 2009).

RESULTSThe geographical distribution of individual strokes and cactus species used in this analysis are shown inFigures 1-8.The strokeArmatocereus cartwrightianus(Britton & Rose) Backeb.ex AW Hill (Fig. 1a) is found in the provinces of El Oro, Guayas, Loja, Manabi and Santa Elena.This species is also present in the departments of Tumbes, Piura and Lambayeque in Peru (Ostolaza 2006).The strokeCleistocactus sepiumvar.morleyanus(Britton & Rose) Madsen (Fig. 1b) is located in the provinces of Chimborazo, Azuay and Canar.The stroke ofOpuntia cylindrica(Lam.) DC.(Fig. 2a) is found in the provinces of Chimborazo, Cotopaxi, Pichincha, and Tungurahua Canar.This species has also been recorded in northern Peru (Madsen 1989).The strokeArmatocereus laetus(Kunth) Backeb.ex AW Hill (Fig. 2b) is located in the province of Loja.This species is also present in the Department of Piura, Peru (Ostolaza 2006).The stroke ofOpuntia soederstromiana(Fig. 3a) is located in the provinces of Chimborazo, Cotopaxi, Imbabura Manabi Pichincha and Esmeraldas.The strokeCleistocactus icosagonus(Kunth) FAC Weber (Fig. 3b) is found in the provinces of Azuay and Loja.This species is also present in northern Peru (Madsen 1989).The stroke ofOpuntia quitensisFAC Weber (Fig. 4) is located in the provinces of Azuay, Chimborazo, El Oro, Loja, Pichincha, with a single record in the province of Cotopaxi.This species has a wide distribution and is found in northern Peru (Madsen 1989) and in parts of Colombia (Fernndez-Alonso & Estupin 2005).The strokePilosocereus tweedyanus(Britton & Rose) Byles & GD Rowley (Fig. 5a) is found in the provinces of Azuay, El Oro, Guayas, Loja, Manabi and Santa Elena.This species has also been recorded in northern Peru (Madsen 1989).The strokeCleistocactus sepium(Kunth) FAC Weber var.sepium(Fig. 5b) is present in the provinces of Chimborazo, Cotopaxi and Tungurahua.The strokeCleistocactus sepiumvar.ventimigliae(Riccob.) Madsen (Fig. 6) has a much broader than the previous two varieties distribution and is found in the provinces of Azuay, Canar, Carchi, Chimborazo, Pichincha, Imbabura and Esmeraldas.The stroke ofEchinopsis pachanoi(Britton & Rose) Friedrich & GD Rowley (Fig. 7) is present in the provinces of Azuay, Canar, Chimborazo, Imbabura, Loja, Pichincha and Tungurahua.This species is also present in northern Peru (Madsen 1989) and in Colombia, in

FIGURES 1-9.individual, generalized nodes and strokes.1-8 individual strokes: 1stArmatocereus cartwrightianus;1bCleistocactus sepiumvar.morleyanus;2a,Opuntia cylindrica,2b, A.laetus;3a, O.soederstromiana,3b, C.icosagonus;4 O.quitensis,5a,Pilosocereus tweedyanus,5b, C.sepiumvar.sepium;6, C.sepiumvar.ventimigliae;7Echinopsis pachanoi;8 O.pubescens;9 generalized tracks and node.FIGURES 1-9.Individual tracks, generalized and nodes.1-8 Individual tracks: 1aArmatocereus cartwrightianus;1bCleistocactus sepiumvar.morleyanus;2a,Opuntia cylindrica,2b, A.laetus;3a, O.soederstromiana,3b, C.icosagonus;4 O.quitensis,5a,Pilosocereus tweedyanus,5b, C.sepiumvar.sepium;6, C.sepiumvar.ventimigliae;7Echinopsispachanoi;8 O.pubescens;9 tracks and generalized node.

Department of Nario (Fernndez-Alonso & Estupin 2005).The strokeOpuntiapubescensJC Wendl.Pfeiff ex.(Fig. 8) is distributed in the provinces of Azuay, Canar, Chimborazo, Imbabura, Loja, Pichincha and Manabi.This species is also distributed in northern Peru(Madsen 1989).

The superposition of the eight individual traces obtained in this analysis allowed the identification of three generalized tracks (Fig. 9): dry coastal (I), north-central interandino (II) and South interandino (III).

Dry coastal line is located in the provinces of Loja, El Oro, Guayas, Santa Elena and Manab, and is supported by speciesArmatocereus cartwrightianus(Fig. 1a) andPilosocereus tweedyanus(Fig. 5a).Other species which are also present in this stroke areOpuntia quitensis(Fig. 4), O.soederstromiana(Fig. 3a),echinopsis pachanoi(Fig. 7) andO.pubescens(Fig. 8).This generalized track is in the biogeographical provinces of Ecuador, and Tumbes-Piura arid and the vegetation comprises of Dry Deciduous Forest (de), Bosque Seco Semideciduous (sd) and Dry Thorny Thicket (ms) (Madsenet al., 2001,Aguirreet al.2006a, 2006b).

The stroke interandino North Central is located in the provinces of Imbabura, Pichincha, Napo, Cotopaxi, Tungurahua, Bolvar, Chimborazo and part of the province of Canar.It is supported byCleistocactus sepiumvar.morleyanus(Fig. 1b),Opuntia cylindrica(Fig. 2a),O.soederstromiana(Fig. 3a),C.sepiumvar.sepium(Fig. 5b),O.quitensis(Fig. 4),C.sepiumvar.ventimigliae(Fig. 6),echinopsis pachanoi(Fig. 7) andO.pubescens(Fig. 8).This generalized track is located in the biogeographic province of western Ecuador and includes the dry forest vegetation of North Interandean (in) and Lower Montane Dry Forest (sm) (Aguirreet al.2006a).

The stroke interandino South is located in the provinces of Loja, Azuay province and part of Canar.It is supported byArmatocereus laetus(Fig. 2b),Cleistocactus icosagonus(Fig. 3b),Opuntia quitensis(Fig. 4),Echinopsis pachanoi(Fig. 7) andO.pubescens(Fig. 8).Other species are also included within this outline areCleistocactus sepiumvar.morleyanus(Fig. 1b) andPilosocereus tweedyanus(Fig. 5a).This generalized track is located in the biogeographic province of western Ecuador and includes the dry forest vegetation of South Interandean (is) and Dry Thorny Thicket (ms) (Aguirreet al.2006a, Loaizaet al., 2009).

Overlapping interandino southern and north-central interandino generalized tracks allowed us to identify the presence of a node (Fig. 9).This node is located between the provinces of Chimborazo and Canar in the biogeographic province of western Ecuador.

DISCUSSIONMost cacti species recorded in Ecuador are distributed in an area between the northwestern and central Peru and southwest of the country (Madsen 1989).Few species have a wide range of distribution, including several epiphytes (Madsen 1989, 2002, included Loaizaet al., 2009) and some other species such asEchinopsis pachanoiandOpuntia quitensis(Fernndez-Alonso & Estupin 2005).The genusOpuntiais characterized as one of the cactus genera with the largest distribution at the continental level (Britton & Rose 1919, Anderson 2001), so many of the species present in Ecuador could have had a much wider distribution in antiquity.However, this distribution would be clearly determined by the effectiveness of the dispersal mechanisms of certain species, as in the case ofOpuntia quitensisandO.pubescens,whose ease evolution of its stem segments has allowed them to have a very favorable distribution.Moreover, the presence of geographical barriers have been the primary factor limiting the expansion of the range of many species, this would explain the presence ofO.quitensisin Colombia and the total absence of other species.The case ofEchinopsis pachanoidiffers from other species, because it has been used since ancient times by man in Peru and Ecuador and is very likely to have been introduced in Colombia and in certain areas of the Ecuadorian coast for some culture ancestral (Madsenet al.2001).

The node located on overlapping South interandino interandino generalized tracks and north-central help to understand and explain the multiple origins of the lineages were present and the difference between cactus species present in both generalized tracks, many of which are endemic to each region.Some of the species itself originated in the identified node and that constitute the inter-line north-central would be:Armatocereus godingianus, leonensis Cleistocactus, C. sepium(three varieties),Espostoa frutescens, Opuntia aequatorialisBritton & Rose,O.bakeriMadsen,O.soederstromianaandWeberocereus rosei(Kimnach) Buxb.

Some authors, such as Valenciaet al.(1999) Sierra (1999), have also agreed to differentiate the inter dry valleys of the southern region (Loja) north central.To the north of the country, the valleys are higher and are located in altitudinal ranges between 1.8002.600 m.These valleys are isolated because both (eastern and western) slopes are covered with very wet montane forests.By contrast, in the south western slopes are relatively dry and the lower mountains, causing theAndean dry forests are located from 1,300 m up, which probably facilitates greater exchange between the dry forests of the coast and the Andean dry forests (Aguirreet al.2006a).The existence of these two generalized tracks within a biogeographic province in the future could allow recognition as districts.However, it should be noted that only the inter-line South have extended distribution to certain areas of northern Peru, being probably interrupted by the presence of geographic barriers that have limited the free distribution of species, promoted in some cases the formation subspecies within a vicarious context, as in the case ofmelocactus bellavistensisRauh & Backeb.subsp.bellavistensis(Lowe 2010).Other genres instead(ArmatocereusandBrowningia),whose origin is attributed to the dry forests of northern Peru, should be analyzed in conjunction with other endemic genera of that country, in order to establish the probable existence of some node not identified, which help to clearly understand the processes of geographical distribution of certain species.

Moreover, the node obtained in overlapping interandino South and interandino of the Ecuador, north-central generalized tracks would be of great importance to the establishment of a priority area for the conservation of cacti, since it combines representatives of two components different biotic, besides its importance could also be considered for the conservation of other groups of plants with similar patterns of geographical distribution.

AcknowledgementsThis research was supported by the Institute of Ecology and Herbarium (HUTPL) of the Technical University of Loja.We wish to thank the collaboration of the following persons and institutions: Carlos Ceron (QAP), Katya Romoleroux (QCA), Elsa Toapanta (QCNE) and Loja Zhofre Aguirre Herbarium for allowing access to and review of the collections in their care.Valuable suggestions and comments made by two anonymous reviewers are also appreciated.

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