THE POTENTIAL NICHE OF Guaiacum coulteri IN PROTECTED NATURAL AREAS OF SONORA TO OAXACA, MEXICO

Bruno Rodríguez-López, Jaime Sanchez, Gamaliel Castañeda G., Miguel Borja, Omag Cano Villegas, Raúl López García and Gisela Muro-Pérez

SUMMARY

The genus Guaiacum belongs to the family Zigophyllaceae, en- Sinaloa, data which matches that of the present study. The re- compassing six species of evergreen trees and shrubs. The geo- sult of ecological niche modeling for G. coulteri has a coverage graphical distribution model allows designing and creating prob- of 24,908.238 km2 and the surface of the ANPs is 13,545.76 km2; abilistic maps of sites where species are distributed. The Maxent when overlaying them, the niche area within these is 1,104.42 program projects patterns of geographic distributions by using km2, which corresponds to a coverage of 9.21%. The results sug- ecological niche models. In Mexico, several studies indicate that gest that modeling of the potential G. coulteri niche matches the the distribution of G. coulteri covers 11 Protected Natural Ar- current distribution of the species; although there are areas of eas (ANP, for their name in Spanish) covering 2,777km2; of this modeling that are not found within the ANP, the resulting model- area, 75% corresponds to the states of Oaxaca, Sonora and ing can be used as a basis for proposing specific fieldwork sites.

Introduction (Maryland-Report, 2000). The of Kricogonia lyside, which blue ‘matlali’ dye named after bud of flowers is blue-violet prefer to lay their eggs on the the Aztecs (Haude, 1998). At The countries with the great- and grow in clusters or isolated; leaves of G. coulteri, considered present, Sonoran inhabitants est biodiversity are tropical blooming occurring during the as a food source plant for but- use a mixture of green organic countries, and such is the case months of April and September terflies in Sonora (Honcoop, resin of G. coulteri, clay and of Mexico, where many species (Mielke, 1993). It can be con- 1997). water to produce the coloring are endemic (Grow and sidered as an endemic species Some medicinal uses are at- agent ‘Antezj kóil’ Schwartzman, 2001) or listed as to Mexico, since it is distributed tributed to this plant, as expec- (Haude,1998). rare (Contreras-Medina et al., naturally from Sonora to torant and blood purifying ele- The population size of G. 2010). The genus Guaiacum Oaxaca (Porter, 1972; Mielke, ment (Grow and Schwartzman, coulteri for the Mexican terri- belongs to the Zigophyllaceae 1993; CITES, 2000; Grow and 2001), as well as laxative, di- tory is unknown, although in family, which consists of six Schwartzman, 2001; Martínez uretic and in the treatment of the Maryland-Report (2000) it species of evergreen trees and and Galindo-Leal 2002, Dertien joint rheumatism (Maryland- is mentioned that this is a lo- shrubs distributed throughout and Duval 2009). The species is Report, 2000). G. coulteri and cally abundant species with Latin America (Maryland- exploited locally, and its natural G. sanctum share other medici- isolated specimens and grows Report, 2000; Grow and populations are under pressure nal properties such as anti-can- at low densities. Because Schwartzman, 2001). Some of due to land use changes that cer, diuretic and anti-inflamma- there are no recent studies, the species of the genus become a threat to their surviv- tory (CITE, 2000). In the case natural populations are likely Guaiacum are in a risk catego- al (Grow and Schwartzman, of G. coulteri, resin extraction to have decreased, as the dis- ry due to loss of their habitat 2001). It is considered a food is used in the manufacture of tributions of G. sanctum and (ARW, 1998; CITES, 2002; source for insects and larvae in special devices for the detection G. officinale have been re- Gordon et al., 2005). various plant associations, such of gastrointestinal bleeding duced due to land use changes Guaiacum coulteri is a small as in deciduous chaparral for- (Barsch et al., 2002). The flow- for agricultural purposes, ur- evergreen tree of slow growth, ests (Maryland-Report, 2000); er of G. coulteri is used in the banization, and for the tour- with a maximum height of 4m an example being the females production of an intense organic ism sector.

KEYWORDS / Distribution / Ecological Niche / Modeling / Protected Natural Areas / Received: 13/03/2020. Modified: 20/02/2021. Accepted: 08/02/2021.

Bruno Rodríguez-López. Master of Gamaliel Castañeda G. Biologist, and Environment of Arid Gisela Muro-Pérez. (Correspondig Science in Biodiversity and UJED, Mexico. Doctor of Scien- Zones, Universidad Autónoma author). Biologist, UJED, Mexico. Ecology, and doctoral student, ces in Ecology and Wildlife de Chapingo, Mexico. Doctoral Doctor of Sciences in Natural Universidad Juárez del Estado de Management, Universidad student, UJED, Mexico. Resources Management, UANL, Durango (UJED), Mexico. Autónoma de Nuevo León Raúl López García. Master of Mexico. Profeesor, UJED, Professor, UJED, Mexico. (UANL), Mexico. Science in Natural Resources Mexico. Address: Facultad de Jaime Sanchez. Biologist, UJED, Miguel Borja. Biologist and Doctor and Environment of Arid Ciencias Biológicas, UJED. Av. Mexico. Doctor of Sciences in in Biomedical Sciences, UJED, Zones, Universidad Autónoma Universidad s/n, Fracc. Natural Resources Management, Mexico. Professor, UJED, Mexico. de Chapingo, Mexico. Filadelfia, CP. 35010, Gómez UANL, Mexico. Professor, Omag Cano Villegas. Master of Doctoral student, UJED, Palacio, Durango, México. e- UJED, Mexico. Science in Natural Resources Mexico. mail: [email protected].

JANUARY 2021 • VOL. 46 Nº 1 0378-1844/14/07/468-08 $ 3.00/0 43 EL NICHO POTENCIAL DE Guaiacum coulteri EN ÁREAS NATURALES PROTEGIDAS DE SONORA A OAXACA, MÉXICO Bruno Rodríguez-López, Jaime Sanchez, Gamaliel Castañeda G., Miguel Borja, Omag Cano Villegas, Raúl López García y Gisela Muro-Pérez RESUMEN El género Guaiacum pertenece a la familia Zigophyllaceae y presente estudio. El resultado del modelaje del nicho ecológico abarca seis especies de árboles y arbustos siempre verdes. El de G.coulteri tiene una cobertura de 24.908,238km2 y la super- modelo de distribución geográfica permite diseñar y crear ma- ficie de las ANP es de 13.545,76km2, que al ser sobrepuestas pas probabilísticos de sitios donde se distribuyen las especies. revelan un área de nicho dentro de 1.104,42km2, la cual corres- El programa Maxent proyecta patrones de distribuciones geo- ponde a una cobertura de 9,21%. Los resultados sugieren que gráficos mediante el empleo de modelos de nicho ecológico. En el modelaje de nicho potencial de G. coulteri coincide con la México, diversos estudios registran 11 Áreas Nacionales Prote- distribución actual de la especie; aunque hay áreas del modela- gidas (ANP) donde se distribuye la especie en estudio, con una je que no se encuentran dentro de las ANP, el modelaje resul- cobertura de 2.777km2; un 75% del área corresponde a los es- tante puede usarse como base para proponer trabajo de campo tados de Oaxaca, Sonora y Sinaloa, datos que coinciden con el en sitios específicos.

O NICHO POTENCIAL DE Guaiacum coulteri EM ÁREAS NATURAIS PROTEGIDAS DE SONORA A OAXACA, NO MÉXICO Bruno Rodríguez-López, Jaime Sanchez, Gamaliel Castañeda G., Miguel Borja, Omag Cano Villegas, Raúl López García e Gisela Muro-Pérez RESUMO O gênero Guaiacum pertence à família Zigophyllaceae e te estudo. O resultado da modelagem do nicho ecológico de G. abrange seis espécies de árvores e arbustos sempre-verdes. O coulteri tem uma cobertura de 24.908,238km2 e a superfície das modelo de distribuição geográfica permite desenhar e criar ANP é de 13.545,76km2, que quando sobrepostas revelam uma mapas probabilísticos de locais onde estão distribuídas as es- área de nicho dentro de 1.104,42km2, que corresponde a uma pécies. O programa Maxent projeta padrões de distribuição cobertura de 9,21%. Os resultados sugerem que a modelagem geográfica mediante a modelagem de nicho ecológico. No Mé- de nicho potencial de G. coulteri coincide com a distribuição xico, diversos estudos registram 11 Áreas Nacionais Protegidas atual da espécie; embora existam áreas da modelagem que não (ANP) onde está distribuída a espécie em estudo, com uma co- se encontram dentro das ANP, a modelagem resultante pode ser bertura de 2.777km2; 75% da área corresponde aos estados de usada como base para propor trabalho de campo em locais es- Oaxaca, Sonora e Sinaloa, dados que coincidem com o presen- pecíficos.

The geographic distribution Natural Protected Areas these serve as a first contact Herbarium (US). Four hundred modelling of plant species is (ANP, for their name in with the species and provide and two records were obtained considered a vitally important Spanish) are sites that contrib- information such as taxonomic from the GBIF portal. tool, since it allows to design ute to the conservation of spe- characteristics and geographical Afterwards, a database with and create probabilistic maps cies, although they are often features that are in turn rooted taxonomic and geographical at- of sites where species are dis- established for political purpos- in websites such as the portal tributes was developed and a tributed (Martínez-Meyer, es, and the species do not have of the Global Fund for duplicate data elimination was 2005). In turn, it is comple- real protection (Paredes-García Biodiversity Information carried out, leaving a total of mented by the provision of et al., 2011). The present study (GBIF). The latter includes col- 139 locations. The requirement computer programs to obtain focused on modeling the poten- lections of the National of having a minimum of 50 lo- biogeographic information and tial ecological niche of Herbarium of Mexico, Instituto cations was met, so that the is also an advantage that can Guaiacum coulteri in ANPs in de Biología, UNAM (MEXU); modeling of the potential distri- be analyzed with a number of the Mexican states of Sonora, the Herbarium of the National bution is not overestimated or, existing records. For these Sinaloa, Nayarit, Jalisco, School of Biological Sciences, in its absence, underestimated models, the Maxent program Colima, Michoacán, Guerrero Instituto Politécnico Nacional (Scheldeman and Zonneveld, stands out, allowing to project and Oaxaca. (ENCB); the Herbarium of the 2011). Google Earth was used patterns of geographical distri- Instituto de Ecología in Xalapa for geo-referencing of the model butions through the use of eco- Methodology (XAL); and the Herbarium of following the methodology of logical niche models, built on the Instituto de Ecología in El Pérez-García and Liria (2013). the basis of specific records of Obtaining species data Bajío (IEB), as well as biologi- collection locations (Phillips et cal collections abroad as, for Modeling of the potential al., 2004, 2006; Hernández et The data for this study were example, the New York niche of the species al., 2006; Phillips and Dudík, collected from scientific collec- Botanical Garden (NY); the 2008; Illoldi-Rangel and tions (Soberon and Nakamura, Missouri Botanical Garden The program used to display Escalante, 2008). 2009; Soberon, 2012), since (MO) and the U.S. National the potential distribution was

44 JANUARY 2021 • VOL. 46 Nº 1 Maxent, since it is one of the the model was selected and in endemic species of flora and eight types of vegetation have best algorithms for modeling turn extrapolated to ArcGis fauna. It has three core areas been identified (Ceballos and the potential species niche 9.3.1 software (ESRI, 2008), that comprise the islands of Miranda, 1986; Castillo and (Ortega and Peterson, 2008). It and placed at the ANP loca- María Magdalena, María Gómez-Pompa, 1991; is considered an artificial intel- tions in order to calculate the Cleofás and San Juanito, as Ceballos, 1991; Ceballos et ligence method that applies the potential distribution on them well as the adjacent marine al., 1994). In the coastal plain principle of maximum entropy and estimate the percentage of portion. María Madre Island is of the Cuitzmala River, of to calculate the most suitable distribution. the largest island, the only one ~800ha, there are numerous geographical distribution of a with a human population and types of vegetation such as species. This method estimates Biodiversity in ANPs has been home to a Federal mangrove, riparian vegetation, the probability of occurrence of Criminal Colony since 1905. carrizal, aquatic vegetation, the species looking for the dis- Zicuirán-Infiernillo: The The Islas Marías Biosphere vegetation of coastal dunes tribution of maximum entropy Zicuirán-Infiernillo Biosphere Reserve was created on and grasslands. In the (as uniform as possible) subject Reserve was established in November 27, 2000, with an ‘lomeríos’ the low deciduous to the condition that the ex- November 2007. It is located in area of 641,285ha and a depth forest dominates, although pected value of each environ- the municipalities of Arteaga, of up to 200m. The Federal there are other types of vege- mental variable according to Churumuco, La Huacana and Criminal Colony is a peniten- tation that are more localized, this distribution coincides with Tumbiscatío, 103km southwest tiary center of the Federal such as the medium-sized the empirical mean. For model- of the city of Morelia, Government of Mexico, estab- sub-deciduous forest. ing, the Contreras-Medina et Michoacán, Mexico. Its exten- lished on May 12, 1905 by Huatulco: A deciduous forest al. (2010) methodology was sion is just over 265,117ha. Porfirio Díaz. At a given time, is located in the polygon of followed, performing a total of The predominating vegetation the worst criminals were held the Huatulco National Park, 100 repetitions to avoid model- in the area is a low deciduous there; subsequently, prisoners which presents a good state of ing biases, accounting 70% of and subcaducifolia forest, with that were not related to the conservation and a high biodi- the records for modeling and no degree of disturbance, cov- government or who had fought versity of flora and fauna. 30% for their evaluation. The ering about 75% of the territory against the state were allowed There are 430 species of flora, cross-validation option was in- of the ANP. Due to the size of to leave. On December 30, 15 species of amphibians, 291 cluded, which consists of repli- the area, this percentage makes 1939, former President Lázaro species of birds, 130 species cating the data for the elabora- it the most extensive area, na- Cárdenas authorized that those of mammals and 72 species of tion of the models and the tionwide, for the protection of prisoners, called there settlers, reptiles in the area. Of the to- same data used to evaluate the this type of vegetation. could live with their families tal, 146 species are under a next model. The parameters Vertebrates have an important (CONANP, 2007). protection status according to used to obtain these models presence in the region; of the Gulf islands of California: NOM-059-SEMARNAT-2010 were: regularization multiplier= 161 species of mammals de- Located in the states of Baja (CONANP, 2003). 1, maximum number of back- scribed for Michoacán, 86 spe- California Sur, Sonora and Cacaxtla Meseta: Located in ground points= 10000, conver- cies have been registered in the Sinaloa, with an area of the municipalities of San gence limit= 0.00001, and max- ANP, which implies that 418,910ha, it was established Ignacio and Mazatlan, in the imum iterations= 500. A set of Zicuirán-Infiernillo is a refuge as a reserve and refuge area state of Sinaloa, covers an area 19 bioclimatic layers were ob- for more than 53% of species. for migratory birds and wild- of 50,862ha. The ANP was de- tained from the WorldClim On the other hand, 253 species life in 1978. The origin of creed in 2000, as a protection portal, as proposed by Hijmans of birds have been registered, these islands is attributed to of flora and fauna area under et al. (2005). Additionally, the representing almost 47% of the processes linked to tectonic the name of ‘Cacaxtla digital elevation model of 539 species registered for the and volcanic activity. There Meseta’. It has a floristic list INEGI (1985) was used. state and about 25% of the spe- are 2,700 plant species in the of more than 200 species of cies registered for all of islands (Wiggins, 1980; Case vascular plants. Among the Bioclimatic layer correlation Mexico. Of them, 101 birds are and Cody, 1983). representative fauna species is analysis migratory. Regarding the herpe- Chamela-Cuixmala and Teopa the gila monster Heloderma tofauna, 69 species have been horrridum (Lavín et al., 2002), Given that bioclimatic layers beach: The establishment of registered, representing 37% of and six existing feline species can be correlated among them, the UNAM biology station in the state amphibian and reptile from Mexico have been regis- a bivariate correlation analysis Chamela took place in 1971. species (CONANP, 2007). tered for the ANP (Ceballos was performed with the SPSS The beaches of Cuixmala and and Oliva, 2005). version 18 software (SPSS, Islas Marías: Located 132km Teopa, located within the re- 2009). If two variables shared from San Blas, Nayarit, the ar- serve, were established as Chamela Bay islands: Located a correlation coefficient >0.8, chipelago called ‘Islas Marías’ sanctuaries for the protection off the coast of the that one was selected as most is made up of four islands, en- of sea turtles on October 29, Municipality of La Huerta, significant layer at the biologi- listed from larger to smaller as 1986. Since 1988, the Jalisco, they constitute the first cal level for each species or María Madre, María Ecological Foundation of sanctuary in Mexico estab- those simplest layers of Magdalena, María Cleofás and Cuixmala, A.C., and UNAM lished as an ANP on June 13, interpretation. San Juanito Islands. It has a conducted studies for the es- 2002. The only nesting colo- subtropical climate with rains tablishment of a biosphere re- nies on the entire Jalisco coast Distribution of the potential in the summer, and encompass- serve in the region, the decree for seabird species such as the niche within the ANPs es different ecosystems such as of which was published on brown pelican (Pelecanus oc- mangroves, low deciduous for- December 30, 1993. The re- cidentalis), the yellow-footed Upon completing modeling ests and medium sub-deciduous serve protects aquatic and ter- booby (Sula leucogaster) and of the potential distribution, forests, with numerous restrial environments, where the blue-footed booby (S.

JANUARY 2021 • VOL. 46 Nº 1 45 nebouxii) occur on these is- extension of 2.4km where this area corresponds to the lands. Other species that nest most of the sea turtles nest; to states of Oaxaca, Sonora and Modeling the ecological niche or take refuge in the islands the center of the Maruata coast Sinaloa, data that coincide with of G. coulteri are the common frigate line is a small beach of 90m the present study. Vanclay (Fregata magnificens), cormo- and at the west end is a beach (2001) and Jackson et al. The generated model fits rants (Phalancrocorax oliva- of ~150m. The average width (2009) have suggested that just the range of G. coulteri. ceus), night heron (Nycticorax of the beaches is 40m. On the because a species is found Moreover, it is worth mention- nycticorax), heron cinder main beach, where the bay is within an ANP, it does not ing that this model predicts (Ardea herodias), white heron located, the site ends abruptly guarantee its conservation; potential niches in the states (A. alba), golden-eared heron with vegetation composed therefore, it is proposed that of Tamaulipas and Baja (Egretta thula), ladle heron mainly of shrubs of the Acacia greater conservation efforts be California Sur; however, there (Cochlearius cochlearius), piji- genus, which marks the limit made. are no records of this species je duck (Dendrocygna autum- of the beach and the beginning in the collections. For nalis), white ibis (Eudocimus of a rocky edge. The beach is G. coulteri surface in ANPs Tamaulipas, the G. angusti- albus), pink spatula (Platalea bare of vegetation, although foluim record is available, and ajaja), dark ibis (Plegadis chi- only Ipomea pescaprae patch- The G. coulteri ecological for Baja that of the species G. hi), common buzzards es are found, which are orient- niche modeling resulted in a sanctum Also, when analyzing (Coragyps atratus), auras ed towards the sea. In the coverage of 24,908,238km2. the records, it was noticed that (Cathartes aura), as well as eastern limit, there is a strip of The surface of the ANPs there were locations in Baja several species of passerine vegetation of ~100m formed (Chamela-Cuixmala, El California and Chiapas, and birds and some migratory ones by shrubs of deciduous forest, Veladero, Chamela Bay Islands, such unconventional records (CONANP, 2008). mainly from the Prosopis, Marías Islands, Cacaxtla are due to a bad identification Escobilla beach: The ecosys- Acacia, Glirisidia, Plateau, Maruata Beach and of the species or to taxonomic tems that are protected in this Phitocellobium and Solanum Colola, Teopa Beach, Zicuiran- errors. Since there are two ANP are mangrove, medi- genera. Across this strip of Infiernillo and Huatulco) was other species and a variety: G. 2 um-sized deciduous forest, vegetation is the coastal road 13,545.76km . When overlaying angustifolium, G. unijugum thorny thicket, coastal dune (INEGI, 1985). the ANP polygons, the area of and G. coulteri var. palmeri, vegetation, tular, floating and the niche within them was according to McCauley et al. Veladero: It is an ecological 2 submerged vegetation, palm reserve located in the bay of 1,104.42km , which corre- (2010) it is likely that the lat- groves, halophyte plant clus- the port of Acapulco, sponds to a coverage of 9.21% ter has a greater similarity. ters, low deciduous forest, and Guerrero, southern Mexico. It (Figure 1). The ANP with the The bioclimatic layers with grassland. The main species of has a surface of 3,159ha and largest distribution areas were greater precision for modeling fauna to be protected is the Zicuirán-Infiernillo with were the seasonality of precip- within the site, there is a lo- 2 golf turtle (Lepidochelys oliva- 518.33km and the Marias itation (Bio 15, with a contri- cality that bears the same 2 cea), since this beach is the name as the park. The vegeta- Islands with 217.69km , while bution of 28,6) and the annual most important nationwide for tion includes, as in most of the highest percentage within average temperature (Bio 1, the number of females that an- the municipality, forests of the ANP is Playa Teopa, with contribution of 31,5), followed nually lay their eggs, although medium subcaducifolia type. 87.48%, followed by Huatulco by elevation (contribution of there are also birds such as the The oaks Quercus affinis and with 67.05% (Figure 1). 9.6%). Of the 139 unique blue hummingbird (Eupherusa Q. laurina predominate, as cyanophrys) that is endemic to well as a deciduous forest. the region and considered The abundant fauna consists threatened, as well as the of species of carnivorous green toucan (Aulacorhynchus birds, reptiles such as iguanas prasinus) and hawks consid- and snakes, boa (Constrictor ered under special protection. imperator), garrobo Marine mammals such as the (Ctenosaura pectinata), osprey false killer whale and several (Pandion haliaetus), scorpion species of dolphins have been (Heloderma horridum) and co- registered in the sea. jolite (Penelope purpurascens) Meanwhile, regarding the flo- (INE, 2007). ra, there is black mangrove (Avicennia germinans), red Results and Discussion mangrove (Rhizophora man- gle), white mangrove Protected natural areas (Laguncularia racemosa) and button mangrove (Conocarpus The distribution of G. coul- erectus) (CONAN, 2009). teri covers 11 Natural Maruata and Colola beach: Protected Areas (APN). These Maruata beach corresponds to results coincide with López et the bay of the same name; it al., 2011, who also recorded 11 is a marine and coastal wet- APNs where the species under land where there are three study is distributed in the beaches. The main beach lo- Mexican territory, with a cov- Figure 1. Distribution of localities of G. coulteri and Protected Natural cated to the east, with an erage of 2,777km2. The 75% of Areas near their distribution.

46 JANUARY 2021 • VOL. 46 Nº 1 distribution of the species. The Reserva de la Biosfera de model shows that the species is Chamela-Cuixmala. Fundación found only in 11 of the 57 Ecológica de Cuixmala-UNAM. México. 141 pp. ANPs close to its known distri- bution, so it is considered to be Ceballos G, Oliva G (2005) Mustela frenata. In Ceballos G, Oliva G insufficiently represented in the (Eds.) Los Mamíferos Silvestres current system of ANPs. de México. CONABIO-UNAM- FCE. México. pp. 380-381. REFERENCES CITES (2000) Transfer of Guaiacum sanctum. From Appendix I. XI ARW (1998) Conservation and sus- Meeting of the Conference of tainable management of trees, the Parties. Convention on Costa Rica. Guaiacum sanctum. International Trade of In IUCN Red List of Threatened Endangered Species Nairobi, Species. Version 2009. Americas Kenya. 62 pp. Regional Workshop. International Union for Conservation of CITES (2002) Inclusion of Nature. Geneva, Switzerland. Guaiacum spp. in Appendix II, http://www.iucnredlist.org (Cons. in accordance with Article II. 26/08/2009). Annotation: Designates all parts and derivative, including wood, Barsch F, Honnef S, Melisch R bark and extract. Convention on (2002) Handel mit Guaiacum International Trade of coulteri in Deutschland. Endangered Species. -TRAFFICEurope Germany im Figure 2. Distribution of the potential niche of G. coulteri in natural Collar NJ (1996) The reasons for red Auftrag des Bundesamtes für protected areas of Mexico. data books. Oryx 30: 121-130. Naturschutz. 5 pp. CONANP (2003) Programa de Case T, Cody M (1983) Synthesis: Manejo de Parque Nacional Pattern and processes in island Huatulco, México. Comisión biogeography. In Case TJ, Cody locations, only 17 records oc- CADNR043 in Nayarit. The Nacional de Áreas Naturales ML (Eds.) Island Biogeography cur within the protected natu- resulting model can be used as Protegidas. Mexico. 209 pp. in the Sea of Cortez. University ral areas, which confers a foundation for proposing field of California Press. Berkeley, CONANP (2007) Memoria de la 12.23% of the total data for work in specific sites, such as CA, USA. 307-341 pp. Consulta Pública de la Reserva the modeling of the ecological some priority areas for conser- de la Biosfera Zicuirán- Castillo G, Gómez-Pompa A (1991) La Infiernillo, en el Estado de niche (Figure 2 and Table I). vation, where the presence of vegetación de Cuixmala. In Gómez- the species has been suggested Michoacán. Comisión Nacional Pompa A (Ed.) Estudio Botánico de Áreas Naturales Protegidas. Conclusions by the model. Yet, there is no Ecológico de las Reservas de Mexico. 39 pp. evidence of the potential niche Cuixmala y Jabalí. Report to UICN CONANP (2007) Programa de The results obtained suggest of the species in the model. For and Botanical Gardens Conservation International. Conservación y Manejo. that the G. coulteri potential this propose, specific sites with Reserva de la Biosfera Islas niche modeling coincides with suitable conditions should be Ceballos G (1991) Estudio Base para Marías. 1st ed. Comisión the current distribution de- proposed strategically for their la Creación de la Reserva de Nacional de Áreas Naturales Cuixmala. Fundación Ecológica Protegidas. México. 220 pp. scribed for the species, although development regarding conser- de Cuixmala. México. 141 pp. there were also areas of the vation purposes. When overlay- CONANP (2008) Programa de Ceballos G, Miranda A (1986) Los Conservación y Manejo del model that are not located with- ing the modeling of G. coulteri Mamíferos de Chamela, Jalisco. in the Natural Protected Areas, on the map of the Natural Santuario Islas de la Bahía de Manual de Campo. Instituto de Chamela. Comisión Nacional de such as Sierra de Álamos-Rio, Protected Areas system, it was Biología, UNAM. México. 436 Áreas Naturales Protegidas. Cuchujaqui in Sonora, Sierra of observed that 91% is not includ- pp. Mexico. 145 pp. Manantlán in Colima, National ed within the limits of these Ceballos G, García A, Rodríguez P CONANP (2009) Estudio Previo Marshes in Nayarit and areas close to the known (1994) Plan de Manejo de la Justificativo para Establecer el Área Natural Protegida en Categoría de Santuario Playa de Escobilla, Santa María Tonameca, Oaxaca. Comisión TABLE I Nacional de Áreas Naturales SURFACE (KM2) OF G. coulteri IN PROTECTED NATURAL AREAS Protegidas. Mexico. 78 pp. Name Area Km2 Niche area in the ANP Percentage Contreras-Medina R, Luna-Vega I, Ríos-Muñoz CA (2010) Zicuirán-Infiernillo 2637.04 518.33 19.66 Distribución de Taxus globosa Marías Islands 6353.12 217.69 3.43 (Taxaceae) en México: modelos Golfo de California Islands 3745.53 201.15 5.37 ecológicos de nicho, efectos del Chamela-Cuixmala 130.74 80.16 61.31 cambio del uso de suelo y con- Huatulco 119.44 80.08 67.05 servación. Rev. Chil. Hist. Nat. Meseta de Cacaxtla 503.18 6.38 1.27 83: 421-433. Bahía de Chamela Islands 19.69 0.21 1.09 Dertien J, Duval M (2009) Teopa Beach 0.12 0.10 87.48 Biogeography and divergence in Escobilla Beach 0.3 0.3 100 Guaiacum sanctum Maruata y Colola Beach 0.33 0.01 3.67 (Zygophyllaceae) revealed in chloroplast DNA: Implications El Veladero 36.27 0.01 0.03 for conservation in the Florida Total 13545.76 1104.42 Keys. Biotropica 41: 120-127.

JANUARY 2021 • VOL. 46 Nº 1 47 ESRI (2008) ArcGIS 9.3.1. INEGI (1985) Síntesis Geográfica Mexico. Rev. Mex. Biodivers. Porter DM (1972) The genera of Environmental Scientific del Estado de Michoacán. 81: 745-758. Zygophyllaceae in the southeast- Research Institute. Redlands, Instituto Nacional de Estadística ern United States. J. Arnold Mielke J (1993) Native Plants for CA, USA. y Geografía. Secretaría de Arboret. 53: 531-552. Southwestern Landscapes. Programación y Presupuesto. Gordon JE, González MA, University of Texas Press. Scheldeman X, Zonneveld MV México. 316 pp. Hernández JV, Lavariega RO, Austin, TX, USA. 310 pp. (2011) Manual de Capacitación Reyes-García A (2005) Jackson SF, Walker K, Gaston KJ en Análisis Espacial de Guaiacum coulteri: An over- (2009) Relationship between Ortega Huerta MA, Peterson AT Diversidad y Distribución de logged dry forest tree of Oaxaca, distributions of threatened (2008) Modeling ecological Plantas. Biodiversity Mexico. Oryx 39: 82-85. plants and protected areas in niches and predicting geograph- International. Roma, Italia. 186 ic distributions: a test of six Grow S, E Schwartzman (2001). The Britain. Biol. Conserv. 142: pp. 1515-1522. presence-only methods. Rev. status of Guaiacum species in Mex. Biodivers. 79: 205-216. SEMARNAT (2010) Norma Oficial trade. Medicinal Plant López-Toledo L, González-Salazar Mexicana NOM-059- Conservation, 7: 19-21, IUCN C, Burslem DFRP, Martínez- Paredes-García DM, Ramírez- SEMARNAT-2010. Protección Medicinal Plant Specialist Group. Ramos M (2011) Conservation Bautista A, Martínez-Morales Ambiental - Especies Nativas de MA (2011) Distribución y repre- Haude ME (1998) Identification of Assessment of Guaiacum sanc- México y de Flora y Fauna sentatividad de las especies del colorants on maps from the ear- tum and Guaiacum coulteri: Silvestres Categorías de Riesgo género Crotalus en las áreas ly colonial period of New Spain Historic Distribution and Future y Especificaciones para su naturales protegidas de México. (Mexico). J. Am. Inst. Conserv. Trends in México. Biotropica Inclusión, Exclusión o Cambio Rev. Mex. Biodivers 82: 37: 240-270. 43: 246-255. - Lista De Especies En Riesgo. 689-700. Secretaría de Medio Ambiente y Martínez E, Galindo-Leal C (2002) Hernandez PA, Graham CH, Master Recursos Naturales. Diario La vegetación de Calakmul: Pérez-García B, Liria J (2013) LL, Albert DL (2006) The ef- Oficial de la Federación. Descripción, composición y dis- Modelos de nicho ecológico fun- fect of sample size and species Mexico. 78 pp. characteristics on performance tribución. Bol. Soc. Bot. Mex. damental para especies del géne- of different species distribution 71: 7-32. ro Thraulodes (Ephemeroptera: Soberón J (2012) Nichos y Áreas de Leptophlebiidae: Atalophlebiinae). Distribución. CONABIO. modeling methods. Ecography Martínez-Meyer E (2005) Climate Rev. Mex. Biodivers 84: Mexico. https://sites.google.com/ 29: 773-785. change and biodiversity: some 600-611. site/nichesandareasofdistribution/ Hijmans RJ, Cameron SE, Parra JL, considerations in core casting sobre-losautores/jorge-soberon. Jones PG, Jarvis A (2005) Very shifts in species potential distri- Phillips SJ, Dudik M, Schapire RE high resolution interpolated cli- butions. Biodivers. Inf. 2: (2004) A maximum entropy Soberón J, Nakamura M (2009) mate surfaces for global land 42-55. approach to species distribution Niches and distributional areas: modeling. In Proc. 21st Int. concepts, methods, and assump- areas. Int. J. Climatol. 25: Maryland-Report (2000) Guaiacum Conf. on Machine Learning. tions. Proc. Nat. Acad. Sci. USA 1965-1978. sanctum: Population status and ACM Press. New York, USA. 106: 19644-19650. Honcoop J (1997) Notes on the mi- trade in Mexico with CITES pp. 655-662. gratory sulphur, Kricogonia ly- recommendations. The 2000 SPSS (2009) SPSS for Windows side. https://www.sasionline.org/ Problem-Solving Team, Phillips SJ, Anderson RP, Schapire (Statistical Package for Social sulf/kric.html University of Maryland. College RE (2006) Maximum entropy Sciences). Version 18. SPSS Inc. Illoldi-Rangel P, Escalante T (2008) Park, MD, USA. 51 pp. modeling of species geographic Chicago, IL, USA. distributions. Ecol. Model. 190: De los modelos de nicho McCauley RA, Cortés-Palomec AC, Vanclay JK (2001) The effectiveness 231-259. ecológico a las áreas de distri- Oyama K (2010) Distribution, of parks. Science 293: bución geográfica. Biogeografía genetic structure, and conserva- Phillips SJ, Dudík M (2008) 1007-1008. 3: 7-12. tion status of the rare microen- Modeling of species distribu- Wiggins I (1980) Flora of Baja INE (2007) Parque Nacional El demic species, Guaiacum uni- tions with Maxent: new exten- California. Stanford University Veladero. Instituto Nacional de jugum (Zygophyllaceae) in the sions and a comprehensive eval- Press. Stanford, CA, USA. 1025 Ecología. México. Cape Region of Baja California, uation. Ecography 31: 161-175. pp.

48 JANUARY 2021 • VOL. 46 Nº 1