Geographic distribution and conservation of Cactaceae ... - Springer Link

Biodiversity and Conservation (2005) 14:2483–2506 DOI 10.1007/s10531-004-0217-y

Ó Springer 2005

Geographic distribution and conservation of Cactaceae from Tamaulipas Mexico JOSE´ GUADALUPE MARTIŃEZ-AVALOS1,* and ENRIQUE JURADO2 1

Instituto de Ecologıá y Alimentos, Universidad Autońoma de Tamaulipas, 13 Blvd. Lo´pez Mateos 928, Cd. Victoria, Tamaulipas, Me´xico; 2Facultad de Ciencias Forestales, UANL. Km. 145 Carretera Cd. Victoria – Linares, Linares, N.L Me´xico, C.P 67700; *Author for coresspondence (e-mail: jmartin@cactus.uat.mx, sucoland@hotmail.com; phone/fax: +52-834-316-27-21) Received 28 July 2003; accepted in revised form 13 April 2004

Key words: Cactaceae, Cacti, Endemism, Flora, Me´xico, Squares, Tamaulipas Abstract. An inventory of the species of Cactaceae from Tamaulipas, Mexico was carried out. The study area was divided into 48 squares of 30 min of latitude and 30 min of longitude. Collections were made at 60 sampling sites distributed over an area extending in to the bordering states of San Luis Potosı´ , Nuevo Leoń, Veracruz and Texas (USA). Species diversity per squares and their geographic distribution range were obtained. Results showed a diversity of 120 species and 53 subspecies, of total 49 are considered as endangered by the CITES, IUCN, and SEMARNAT. The genera most represented were Mammillaria, Coryphantha, Echinocereus and Opuntia. Index of Geographical Expansion (IGE) was calculated for each species. Highest diversity per squares as well as the IGE for each species was found at square 66 (Jaumave) IGE = 52%, 72 (Tanque Blanco) IGE = 22% and 59 (Cd. Victoria) IGE = 7%. It was for these sites that had the highest number of endemic taxa and thus are recommended as priority sites for conservation. This information may help establish natural protected areas by the government of the State of Tamaulipas.

Introduction Northeastern Mexico has a very interesting floristic composition, with a great diversity of climatic, topographic and environmental conditions. More than 26 vegetation types at different altitudinal ranges have been recognized by Miranda and Hernańdez (1985) and Rzedowski (1978). However, rosetophyllous scrubs, piedmont scrubs, deciduous low forests, mesquite forests, pine-oak forests and halophilic vegetation are among the most important ones on the basis of their distribution in this part of Mexico (INEGI 1993). There is a high diversity of species of Cactaceae, Compositae and Gramineae that are also abundant in arid and semiarid regions from the north and center of the country (Rzedowski 1991a, b). Hunt (1999) recognizes 48 genera and 570 species (plus some 350 provisionally accepted species), of Cactaceae in Mexico, of which 529 of these are endemic and 63 are endangered and are listed in Appendix 1 of CITES (CITES 1990). This family is widely distributed in the Sonoran and Chihuahuan Deserts in the central and northern zones of Mexico (McMahon 1993) however, there are some other smaller regions such as the Tehuacań and

2484 Cuicatlań valleys in the south of Mexico where there is a high number of species of columnar cacti growing under different climatic conditions (A´rias et al. 1997, Da´vila et al. 2002). However, Hernańdez and Ba´rcenas (1995, 1996), Go´mezHinostrosa and Hernańdez (2000), and Martı´ nez-Avalos (1998) pointed out that the southern part of the Chihuahuan Desert, consisting of the southern part of the states of Coahuila and Nuevo Leoń, northern San Luis Potosı´ and southeastern Tamaulipas, together with the northeastern region of Mexico, are the most important regarding the diversity and endemic taxa of Cactaceae in Mexico. Some examples of the latter are the valley of Cuatro Cienegas in Coahuila with 48 spp. (Pinkava 1984), the southern part of the state of Nuevo Leoń and more specifically the municipalities of Mier y Noriega in Nuevo Leoń with 54 spp. (Go´mez-Hinostrosa and Hernańdez 2000), Doctor Arroyo, Nuevo Leon with 43 spp. (Hernańdez 1981), the locality known as El Huizache, San Luis Potosı´ with 71 spp. (Hernańdez et al. 2001), and the Biosphere Reserve ‘‘El Cielo’’ in Tamaulipas with 44 spp. (Martı´ nez-Avalos and Jimeńez 1993; Martı´ nez-Avalos et al. 1996). Cacti belong to a young family in evolutionary times (Barthlott and Hunt 1993) and also to one of the most currently threatened. Because of this, many studies dealing with taxonomy (Anderson 2003; Bravo-Hollis 1978; Bravo-Hollis and Sańchez-Mejorada 1991a, b; Guzmań et al. 2003), demography (ValienteBanuet 1991a, b; Martı´ nez-Avalos et al. 1993, 1994), reproductive biology (Godı´ nez and Valiente-Banuet 1998; Valiente-Banuet et al. 1997a, b), biogeography and inventories have been conducted throughout the American Continent (Go´mez-Hinostrosa and Hernańdez 2000; Hernańdez and Ba´rcenas 1996, 1991). This study is part of a broader study to evaluate cacti species of the State of Tamaulipas, Mexico. In here we aim at describing species richness, geographic distribution span, and priority areas for conservation with large numbers of rare species. The number of specimens of the species included in the herbarium was substantially increased and a computer database with all relevant data on taxonomy and geography of the species was developed.

Study area The state of Tamaulipas is located in the northeast of Mexico (22 120 , 27 400 north latitude and 97 080 ,100 080 west longitude). It neighbors the USA on the north, the Gulf of Me´xico on the east, the state of Veracruz on the south, and the states of Nuevo Leoń and San Luis Potosı´ on the west. The state of Tamaulipas has an area of 7980 km2 and its physiography is composed of elements of the North Gulf Coastal province, the Great North American Plains and the Sierra Madre Oriental. Climatic conditions of the latter regions are either hot and humid or semidesertic (Garcı´ a 1983). The vegetation of Tamaulipas has been grouped into 13 types by Miranda and Hernańdez (1985). The most large and important types with high number of endemic plant species are: rosetophyllus scrubs, piedmont thornscrubs, medium subperennial forest, thornscrubs, pine-oak forests and halophytic vegetation. These vegetation

2485 types are recognized by the Mexican System for Biological Inventories (Sistema Nacional de Inventarios Biolo´gicos: SNIB, (Hernańdez et al. 1993). Methods Fieldwork All accepted species and subspecies of cacti for Tamaulipas were included in this study. A large number of subspecies are endemic to the study site and are listed on Appendix 1 form CITES. To determine the size of the bhome range of species and subspecies the State of Tamaulipas was divided in 48 grid squares of 30 min of latitude and 30 min of longitude (Figure 1). With this we could identify species that were more narrowly distributed and the areas with the highest number of rare species according to ‘‘Norma Oficial Mexicana (NOM059-ECOL-2001)’’ (SEMARNAT 2002), the IUCN Red List of Threatened Species (www.redlist.org) and those species included on Appendix 1 of the Convention International Trade in Endangered Species of Wild Fauna and Flora (CITES). The selection of the sites was determined by presence of well preserved, natural vegetation. Highly disturbed sites were avoided following Go´mez-Hinostrosa and Hernańdez (2003). At each square an intensive collection of all species of cacti was made. A total of 24 field trips were made and 60 sites were sampled and located using GPS technology. At each one of the localities observed species were counted using either a band transect methodology or a line transect as reported for counting birds and mammals (Davis and Winstead 1987; Baillie 1991; Bukland et al. 1993), as well as for plant species diversity in several vegetation types. Two or three specimens per species and locality were collected and included at the herbarium of the Universidad Autońoma de Tamaulipas and herbarium of the Instituto de Biologı´ a, Universidad Nacional Autońoma de Me´xico. Database A database with all the field data was made using Fox pro v.2.5. Fifteen different fields were used. Following the format of the Sistema Nacional de Inventarios Biolo´gicos (SNIB), the data were organized in tables with different options, and information is shown in a synoptic way so that new records and updating of information will be easy to accommodate Hernańdez et al. 1993). Index of Geographical Expansion To estimate the breath of the geography index occupied by each species and subspecies an Index of Geographical Expansion (IGE) was used. This has been used in other studies to quantify the geographic extent of a given species in a

2486

Figure 1. Location of square samples in the region of the state of Tamaulipas.

defined area (Hernańdez and Ba´rcenas 1995, 1996; Arita et al. 1997; Go´mezHinostrosa and Hernańdez 2000). Thus the state of Tamaulipas was divided in squares of 30 min of latitude by 30 min of longitude, following Hernańdez and

2487 Ba´rcenas (1995) to determine patterns of distribution of threatened cacti of the Chihuahuan Desert. In addition to field visits, data bases from over 8500 cacti samples (Hernańdez et al. 1993) were used to determine their presence in each square, IGE was calculated according to the formula IGE = Ss/Sm, where Ss is the number of grid squares from which the species has been recorded and Sm the number of grid squares occupied by the species with the most extensive geographical range (Go´mez-Hinostrosa and Hernańdez 2000). Cylindropuntia leptocaulis was the most frequent species, together with Opuntia engelmannii subsp. engelmannii, as they were distributed in 47 of 48 grid squares respectively in the study area. Cylindropuntia leptocaulis was found in the maximum number of grid squares and was used in order to obtain the IGE of the other species. The total number of grid squares where each species was recorded was then divided by 47. Following this procedure, some geographically restricted species such as Turbinicarpus vierecki subsp. viereckii and T. saueri subsp. ysabelae with a IGE = 0.021 were registered. Cylindropuntia leptocaulis has a wide distribution area since it was found in 47 grid squares and it has an IGE = 0.979. It is important to emphasize than the IGE’s were calculated considering the whole distribution range of the species.

Analysis of complementarity The analysis of complementarity has been used in other studies in order to determine priorities to optimize the conservation of areas with a high biodiversity in specific areas (Ackery and Vane-Wright 1984; Humphries et al. 1991; Williams et al. 1991; Pressey et al. 1993; Arita et al. 1997). In Mexico it has been used by Hernańdez et al. (2001); Hernańdez and Ba´rcenas (1995; 1996) and Go´mez-Hinostrosa and Hernańdez (2000) to determine areas with the highest diversity of cacti in the South part of the Chihuahuan Desert. The principle is based upon the identification of a first priority area, which is defined by having the highest number of species. The additional areas are ordered as second, third priority, etc. according to the contribution of additional species not found in the areas of higher priority (Go´mez-Hinostrosa and Hernańdez 2000). This method was used at 48 grid squares in the state of Tamaulipas. Complementarity values were calculated according to the formula CV = RC 100/CO, where CV is the percentage of Complementarity, RC the residual complement, and CO the complement. The complement is defined as the total number of species present in all grid squares from Tamaulipas (CV = 134). The residual complement is the number of species not found at the areas of higher priority. A second complementarity analysis was made following Hernańdez et al. (2001) and Go´mez-Hinostrosa and Hernańdez (2000) using the endangered species method.

2488 Results and discussion Database A database with 1457 records of herbarium specimens was developed; this is compatible with other database systems used at national and international herbaria. Using the works by Anderson (2001) and Guzmań et al. (2003) with other taxa found here, a total of 35 genera, 120 species and 53 subspecies of cacti were identified. This is the result of 24 field trips made at 48 quadrants at an equivalent area of 2700 km2 each in all the state of Tamaulipas (see Appendix 1). The larger than previously reported number of taxa in this study was a result of new species (Turbinicarpus nieblae), and new nomenclature by Anderson (2001) and Guzmań et al. (2003) as well as new records for Tamaulipas (Echinocereus knippelianus, Ferocactus histrix and F. glaucescens).

Diversity The square with the highest numbers of species and subspecies (square 66), at Jaumave, had 70 cacti taxa (species and subspecies), from which 18 are listed as endangered; this was followed by square 59 (Cd. Victoria) with 67 species and 18 endangered, square 72 (Tanque Blanco) with 64 species and 21 endangered, square 71 (La´zaro Ca´rdenas) with 58 species and 17 endangered, square 65 (Bustamante) with 57 species and 18 endangered, square 64 (Mier y Noriega) with 51 species and 15 endangered, and square 58 (Miquihuana) with 50 species and 18 endangered (Figure 2). These results showed a high diversity of cacti and large numbers of endangered species in the southeastern region. The large number of endemic cacti at square 66 (Jaumave), 72 (Tanque Blanco) and 59 (Cd. Victoria), could be associated to aridity. As these squares are at the Edge of the Chihuahuan Desert and away from moist winds to the Sierra Madre Oriental. Figure 4 shows the three sites with the lowest rainfall ranging from 200 to 500 mm, with arid land vegetation. It is interesting to note that in this region that has the lowest rainfall ranges and the rosetophyllous scrub is the main vegetation type where cacti are found with 113 species and subspecies, 39 of which are classified as endangered. This is followed by piedmont scrub with 41 species and 7 endangered; medium subperennial forests 36 species and 6 endangered; thornscrubs with 26 species and 4 endangered; pine-oak forests with 21 species and 2 endangered; and halophytic vegetation with 15 species and 0 endangered (Figure 3). The 120 species and 53 subspecies of cacti found for the state of Tamaulipas belong to 35 genera. The genus Mammillaria had the highest number of species with 23 (21 subspecies), followed by the genus Coryphantha, Echinocereus and Opuntia with 13 species (1, 4 and 9 subsp)., Turbinicarpus with 7 (4 subsp)., Ferocactus and Thelocactus with 5 species (1 and 5 subsp). All other genera had less than 5 species each (Table 3).

2489

Figure 2. Distribution of Cacti per squares, and squares with the highest diversity in the region of Tamaulipas.

Considering that 1457 specimens of cacti were collected from the state of Tamaulipas and added to the herbarium collection of the Universidad Autońoma de Tamaulipas, this might be one of the best cacti collections from Mexico, particularly from the northeastern region.

2490 Endangered 160

Not Endangered

Number of species

140 120 100 80 60 40 20 0 Rs

Pt

Msf

T

PQf

Hv

Vegetation Type

Figure 3. Distribution of the species of Cactaceae and vegetation type in the region of Tamaulipas. Rs: Rosetophyllous scrubs; Pt: Piedmont thornscrubs; Msf: Medium subperennial forests; T: Thornscrubs; POF: Pine-Oak forests; Hv: Halophytic vegetation. Table 1. Cacti species diversity in Northeaster Me´xico regioń. State

Genus

Species

Endemic

Endangered

Coahuila Nuevo Leońa Tamaulipas San Luis Potosı´ b

17 24 35 33

126 136 120 151

20 14 12 9

49 61 49 60

a Gonza´lez-Botello, M. and L. Rocha; b M. Martı´ nez pers. obs. (Martı´ nez-Avalos 1998; Anderson 2001; Guzmań et al. 2003).

The diversity of cacti from the state of Tamaulipas was compared with that of other states from the northeastern region that are considered important for their high diversity of cacti. Considering work from Anderson (2001), Guzman et al. (2003) and Martı´ nez-Avalos (1998), as well as information form local herbaria, Table 1 was constructed for several states. Thus the state of San Luis Potosı´ was the one with the highest cacti species richness (151 species and 61 threatened) from northern Mexico, followed by Nuevo Leoń (136 species and 61 threatened), Coahuila (126 species and 49 threatened) and Tamaulipas (120 species and 46 threatened).

Size of geographical range Figure 2 shows squares 66 (Jaumave), 59 (Cd.Victoria) and 72 (Tanque Blanco) as the sites with more than 61 species of cacti in the state of Tamaulipas. Other important squares are the 71 (La´zaro Ca´rdenas), 65 (Bustamante), 64 (Mier y Noriega) and 58 (Miquihuana), since they have from 41 to 60 cacti species. All other square have from 1 to 40 species each. This high number

2491 might be associated with the lowest rainfall (200–500 mm) and the influence of the Chihuahuan Desert that reaches valley de Jaumave (Figure 4). Index Geographical Expansion The index geographical expansion(IGE) is an important tool used to determine the conservation status of the species in their area of distribution. Value gradients for the IGE of the species and subspecies is shown in Figure 5 (Appendix 2). The median of all the values for the GWI of the species (M = 0.106) was used in order to classify the species into two categories: species with a wide distribution and species with a restricted distribution. Of the 134 taxa (120 species and 53 subspecies found in the state of Tamaulipas, 54% (73 spp.) had a restricted distribution, while 46% (46 spp.) had a wider distribution. Among some of the restricted-distribution species are: Turbinicarpus jauernigii, T. nieblae, T. saueri subsp. saueri, it T. saueri subsp. ysabelae and T. viereckii subsp. viereckiihe, seven species of the genus Mammillaria, Cumarina odorata, Echinocereus knippelianus, etc, and they showed a IGE = 0.021. Species with the highest distribution range were: Cylindropuntia leptocaulis, Opuntia engelmannii subsp. engelmannii, (IGE = 0.979) and were found in 47 out of 48 squares, followed by Ferocactus hamatacanthus subsp. hamatacanthus, with an IGE = 0.936, Mammillaria heyderii subsp. heyderii with an IGE = 0.915 and Echinocactus texensis, with an IGE = 0.915. The latter species are consideredby Hernańdez et al. (2001) and Go´mez-Hinostrosa and Hernańdez (2000) as outside the Chihuahuan Desert. Endemism From the 73 species and subspecies of cacti with a restricted IGE, only 63% (i.e., 42 spp.) are essentiality endemic to the Chihuahuan Desert Region. Included among these, are species of Ariocarpus, Astrophytum, Coryphantha, Leuchtenbergia, Neolloydia, Thelocactus and Turbinicarpus, all of which are genera endemic to this region. Also, 22 species and subspecies of cacti were found in 1 square only. All species of Turbinicarpus were restricted to one square, as well as 7 species from the genus Mammillaria, Mammillaria carmenae, M. laui subsp. dasyacantha, M. laui subsp. laui and M. laui subsp. subducta, M. plumose, M. prolifera subsp. zublerae and M. shiediana subsp. giselae. Other species found only in one square were: Cumarina odorata, Echinocereus knippelianus, Epithelantha micromeres subsp. micromeris, Ferocactus glaucescens, F. histrix, Grusonia dumetorum, Pelecyphora strobiliformis, Stenocactus multicostatus subsp. multicostatus, Thelocactus conothelos subsp. garciae (Figure 5) (Appendix 2) . From all species and subspecies found only on a small number squares (73 taxa), 15 are endemic to Tamaulipas, (Guzmań et al. 2003). In Mexico there are 913 species and 669 subspecies (Guzmań et al. 2003), in 63 genera. From these

2492

Figure 4. Mean annual rainfall distribution at the region of Tamaulipas.

724 taxa (518 species and 206 subspecies) are considered endemic for Me´xico, Thus in here we found that Tamaulipas has 2% of endemic species and subspecies of Mexico (Appendix 1). In contrast, the species that had a IGE wide (i.e. 48%) were found at more than five squares and they occur throughout most of the Chihuahuan Desert.

Figure 5. Geographical Expansion Index of Cactaceae species in the region of the state of Tamaulipas.

2493

2494 Table 2. Complementarity values for the cacti species used to determine priority sites for conservation proposals at the region of the state of Tamaulipas. Priority

Name

Square

No. of unique species

Complementarity residual value

Complementarity value (%)

First Second Third Fourth Fourth Fourth Fourth Fourth Fifth Fifth Sixth Sixth Sixth Sixth

Jaumave Tanque Blanco Cd. Victoria Hidalgo Mante San Carlos San Rafael Miquihuana La´zaro Ca´rdenas Lomas del Real Bustamante Mendez Vaquerı´ as Zapata

66 72 59 52 74 46 16 58 71 76 65 39 38 17

70 64 67 23 25 26 24 50 58 15 57 23 24 24

70 28 9 4 4 4 4 3 2 2 1 1 1 1

52 22 7 3 3 3 3 3 1 1 0.746 0.746 0.746 0.746

Endangered species First Tanque Blanco Second Cd. Victoria Third Jaumave Fourth La´zaro Ca´rdenas Fourth Miquihuana Fourth San Carlos Fifth Bustamante Sixth El Picacho

72 59 66 71 58 46 65 68

21 18 18 17 18 5 18 2

21 12 4 3 3 3 2 1

43 24 8 6 6 6 4 2

Examples of the latter are Cylindropuntia leptocaulis, Opuntia engelmannii subsp. engelmannii and Ferocactus amatacanthus subsp. hamatacanthus, which were found in 47 and 44 of 48 squares. According to the literature, these species can be found in most of the states from northern and central Me´xico – a wide geographic distribution (Figure 5) (Appendix 2).

Complementarity analysis According to the complementarity analysis (CV) made for all species, square 66 (Jaumave) is considered as first priority with 70 species and a CV = 52% for all the species in the area. Square 72 (Tanque Blanco) had the second priority range with a CV = 22%; square 59 (Cd. Victoria) had a third priority range with a CV = 7%; squares 52 (Hidalgo), 74 (Mante), 46 (San Carlos), 16 (San Rafael) and 58 (Miquihuana) had a fourth priority range with a CV = 3%; squares 71 (La´zaro Ca´rdenas) and 76 (Lomas del Real) fifth priority with a CV = 1% each, a sixth priority range was for squares 65 (Bustamante), 39 (Mendez), 38 (Vaquerı´ as) and 17 (Zapata) with a CV = 0.746.

2495 Table 3. Taxonomic composition of species and subspecies of cacti present from Tamaulips region.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35

Genus

No. of species

No. of subspecies

Mammillaria Coryphantha Echinocereus Opuntia Turbinucarpus Ferocactus Thelocactus Cylindropuntia Ariocarpus Echinocactus Selenicereus Astrophytum Grusonia Stenocactus Stenocereus Acanthocereus Cumarina Echinomastus Epithelantha Escobaria Glandulicactus Leuchtembergia Lophophora Marginatocereus Mammilloydia Myrtillocactus Neobuxbaumia Neolloydia Obregonia Pelecyphora Pereskia Pereskiopsis Pilosocereus Rhipsalis Sclerocactus

23 13 13 13 7 5 5 4 3 3 3 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 120

21 1 9 4 4 1 5 1 2 1 0 0 0 1 0 0 0 0 1 0 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 53

Regarding complementary analysis for endangered species, square 72 (Tanque Blanco) showed the highest number of endangered species with 21 species (CV = 43%) and was thus considered as first priority for conservation. This means that conservation of square 72 would help protect 43% of the endangered species of Tamaulipas. Square 59 (Cd. Victoria) was considered as a second priority with a CV = 27%; square 66 (Jaumave) was third priority with a CV = 11%; squares 71 (L. Ca´rdenas), 58 (Miquihauan), and 46 (San Carlos) were fourth priority with a CV = 6%; square 65 (Bustamante) was considered as fifth priority with a CV = 4%, square 68 (El Picacho) was

2496 considered the sixth priority with a CV = 2%. All other squares had a seventh priority designation (Table 2). Squares 72 (Tanque Blanco), 59 (Victoria) and 66 (Jaumave), are firstpriority sites for conservation of cacti species. The latter is due to the high species diversity found, to the higher number of endangered species and to their restricted geographic distribution at each squares. Thus with results presented here and those of Hernańdez et al. (2001) and Go´mez-Hinostrosa and Hernańdez (2000), the region of Tula, Bustamante, Miquihuana, Valle de Jaumave (squares 72, 71, 58 and 66) should be considered as natural areas for the conservation of cacti by the State Authorities of Tamaulipas, and perhaps be promoted Natural Protected areas. Similar diversity values have been used to declare Natural protected areas for cacti at Huizache by the San Luis Potosı´ government and Mier y Noriega (in process) by the government of Nuevo Leoń. With results presented here, Mammillaria prolifera subsp. zublerae, M. schiedeana susbp. giselae, Thelocactus conothelos subsp. garciae, restricted to one locality in Tamaulipas should be listed as threatened. Rosetophyllous scrub and piedmont thornscrubs in the arid and semiarid zone located at the south of the state are the most important vegetation types in terms of biodiversity protection, as they have both, the largest number of cacti taxa and endemisms as well as other plant and birds and mammals (Escalante et al. 1995; Fa and Morales 1995; Rzedowski 1991a, b).

Acknowledgements Authors would like to thank Fortunato Garza-Ocanãs, Hector M. Hernandez, Terry Martin, Jorge S. Marroquı´ n and Gerardo Sańchez-Ramos for their comments and suggestions to this paper. Thanks are also given to anonymous reviewers and to all other people that made this study and its publication possible. We would like to thank the support given by the Consejo Tamaulipeco de Ciencia y Tecnologı´ a, Consejo Nacional de Ciencia y Tecnologı´ a and Comision Nacional para el Conocimiento y Uso de la Biodiversidad through projects P120 and H149, FOMIX: TAMPS-2002-CO1-3536 and PROMEPUAT.

Acanthocereus Ariocarpus Ariocarpus Ariocarpus

Astrophytum Astrophytum Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Cumarinia Cylindropuntia Cylindropuntia Cylindropuntia Cylindropuntia Echinocactus Echinocactus Echinocactus Echinocereus

1 2 3 4

5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28

SPECIES tetragonus (Linnaeus) Hummelinck agavoides (Castan˜eda) E. F. Anderson* kotschoubeyanus (Lemaire) K. Shumann* retusus Sheidweiler subsp. retusus* retusus Sheidweiler subsp. trigonus* asterias (Karwinski ex Zucc) Lemaire* myriostigma Lemaire* delicata L. Bremer* georgii Boedeker* glanduligera (Otto ex A. Dietrich) Lemaire* guerkeana (Boedeker) Br. & Rose macromeris (Engelmann) Lemaire subsp. macromeris nickelsiae (K. Brandegee) Br. & Rose* octacantha (D C) Br. & Rose pulleineana (Backeberg) Glass* radians (D C) Br. & Rose salinensis (Poselger) A. D. Zimmerman ex Dicht & A. Lu¨ty sulcata (Engelmann) Br. & Rose vaupelianaBoedeker** wohlschlageri Holzeis* odorata (Boedeker) Buxbaum* imbricatta (C.B. Wolf) Rebman & Pinkava subsp. imbricata kleiniae (D C) F. M. Knuth leptocaulis (D C) F. M Knuth tunicata (Lehmann) Link Otto ex Pfeiffer horizonthalonius Lemaire susbp. horizonthalonius platyacanthus Link & Otto* texensis Hoffer berlandieri (Engelmann) Haage

0–774 1050–1128 1050–1189 850–1524 304–670 189–254 600–731 650–745 1036–1950 750–1900 670–1128 220–300 400–576 1050–1780 914–1100 750–1900 400–600 350–700 1432–1900 1050–1100 670–1430 0–1649 1000–1200 0–1400 1000–1750 1000–1189 975–1950 230–1700 1615–2250

ALTIDUDE (m) MA 0405 MA 0468 MA 0437 MA 0054 MA 0383 MA 0540 MA 0063 MA 0720 MA 0619 HMH 2045 MA 0459 MA 0851 MA1023 MA 0366 MA 0469 MA 0142 MA 1112 MA 1040 MA 0275 MA 0998 CGH 1288 MA 0126 MA 0525 MA 0748 MA 0260 MA 0438 MA 0474 MA 0304 MA 0038

VOUCHER UAT UAT UAT UAT UAT UAT UAT UAT UAT MEXU UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT MEXU UAT UAT UAT UAT UAT UAT UAT UAT

HERBARIUM

Appendix 1. List of cacti found in the State of Tamaulipas and adjacent regions. Taxonomic concepts based on Anderson (2001) and Guzman et al. (2003).

2497

29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57

Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinocereus Echinomastus Epithelantha Escobaria Ferocactus Ferocactus Ferocactus Ferocactus Ferocactus Glandulicactus Grusonia Grusonia Leuchtenbergia Lophophora Mammillaria Mammillaria Mammillaria Mammillaria

SPECIES

Appendix 1. Continued.

cinerascens (Bravo) N. P. Taylor subsp. tulensis enneacanthus Engelmann subsp. enneacanthus knippelianus Liebner* + papillosus Linke ex Ruëmpler parkeri (N. P. Taylor) N. P. Taylor subsp. gonzalezii pectinatus (Sheidweiler) Engelmann subsp. pectinatus pentalophus (D C) Lemiare subsp. pentalophus poselgeri Lemiare* reichenbachii (Terscheck ex Walp.) Haage subsp. reichenbachii stramineus (Engelmann) Seitz subsp. stramineus + triglochidiatus Engelmann subsp. triglochidiatus + viereckii Werdermann subsp. viereckii* gautii (L. D. Benson) Mosco & Zanovello micromeres (Engelmann) Weber ex Br. & Rose subsp. micromeres* + emskoetteriana (Quehl) Borg echidne (D C) Br. & Rose glaucescens (D C) Br. & Rose + hamatacanthus (Muehlenpfordt) Br. & Rose subsp. hamatacanthus histrix (D C) Linsay* + pilosus (Galeotti ex Salm-Dyck) Werdermann* uncinatus (Galeotti ex Pfeiff. & Otto) Backemberg subsp. uncinatus* dumetorum (A. Berger) E. F. Anderson** shottii (Engelmann) H. Rob. principis Hooker* williamsii (Lemaire ex Salm-Dyck) J. Coulter* albicoma Boedeker* anniana Glass & Foster* ** baumii Boedeker* carmenae Castan˜eda & Nu`n˜ez* **

900–1900 200–400 2000–2682 250–792 1450–1900 400–1493 200–1500 732–1300 350–400 1900–2100 1500–2100 300–1450 1500–2255 1900–2000 332–420 630–2133 550–670 200–2100 2400–2800 1780–2300 1050–2100 1600–19001 274–450 1493–1900 350–1127 1859–2400 108–200 600–740 914–1250

ALTIDUDE (m) MA 0037 MA 0533 MA 0056 MA 0522 MA 0792 MA 0341 MA 0826 MA 0625 MA 1154 MA 0338 MA 0989 MA 0122 MA 0059 MA 1234 MA 0767 MA 0263 MA 1275 MA 0693 MA 1124 MA 0368 MA 0615 MA 1203 MA 0427 MA 0350 MA 0105 MA 0001 LAU 1332 MA 0328 MA 0601

VOUCHER UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT MEXU UAT UAT

HERBARIUM

2498

Mammillaria

Mammillaria

Mammillaria

Mammillaria

Mammillaria Mammillaria

Mammillaria Mammillaria Mammillaria

Mammillaria Mammillaria

Mammillaria Mammillaria Mammillaria

Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Mammilloydia Marginatocereus Myrtillocactus

58

59

60

61

62 63

64 65 66

67 68

69 70 71

72 73 74 75 76 77 78 79

compressa (Reppenhagen) Hunt subsp. centralifera compressaD C. subsp. compressa formosa Galeotti ex Scheidweiler subsp. formosa formosa(Muelenpfort) Hunt subsp. microthele glassii (Reppenhagen) Hunt subsp. ascensionis glassii Foster subsp. glassii heyderi (Engelmann) Hunt subsp. gummifera heyderi (Engelmann) Hunt subsp. hemiesphaerica heyderi Mulenpfordt subsp. heyderii heyderi (Engelmann) Hunt subsp. meiacantha klissingiana Boedeker* laui (Hunt) Hunt subsp. dasyacantha* ** laui Hunt subsp. laui* ** laui (Hunt) Hunt subsp. subducta* ** magnimamma Harworth melaleuca Karwinski ex Salm-Dyck*** melanocentra Poselger subsp. melanocentra melanocentra(Reppenhagen & Lau) Hunt subsp. rubrograndis* prebella Hildmann ex Schumann picta Meinsausen subsp. picta picta(Boedeker) Hunt subsp. viereckii pilispina Purpus* plumose (F.A.C.) Weber* prolifera (Miller) Haworth subsp. arachnoideae prolifera (Reppenhagen) Hunt subsp. zublerae roseoalba Boedeker* scheideana subsp. giselae (J.G. Martı´ nez-Avalos & Glass) J. Lu¨ty sphaerica A. Dietrich surculosa Boedeker* winterae Boedeker subsp. winterae candida (Scheidweiler) Buxbaum subsp. candida* marginatus (D C) Backeberg geometrizans (Martius ex Pfeiffer) Console

1000–1280 1050–1250 1050–1767 1000–1250 1450–1500 1500–1900 400–790 400–750 250–750 50–750 630–1320 800–1200 800–1200 800–1200 1000–1280 600–1900 1500–1900 1200–1450 780–1036 600–1402 600–1584 1600–2050 430–500 400–1250 980–1200 600–731 1400–2100 274–400 1000–1250 800–1500 600–1250 700–1900 1402–2400

MA 0490 MA 0503 MA 0360 MA 0694 MA 0776 MA 0720 OB 1263 HMH 2040 MA 0746 MA 0672 MA 0177 MA 0859 MA 0153 MA 0860 MA 0120 MA 0783 MA 0779 MA 0152 MA 0614 MA 0356 MA 0217 MA 0775 MA 0976 MA 0167 MA 0330 MA 0507 MA 0135 MA 0402 MA 0662 MA 0706 MA 0663 MA 0061 MA 0203

UAT UAT UAT UAT UAT UAT ANSM MEXU UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT

2499

80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98 99 100 101 102 103 104 105 106 107 108 109

Neobuxbaumia Neolloydia Obregonia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Opuntia Pelecyphora Pereskiopsis Pereskia Pilosocereus Rhipsalis Sclerocactus Selenicereus Selenicereus Selenicereus Stenocactus Stenocactsu Stenocereus Stenocereus Thelocactus

SPECIES

Appendix 1. Continued.

euphorbioides (Haworth) Buxbaum ex Bravo conoidea (DC) Br. & Rose denegrii Fric* ** decumbens Sal-Dyck dejecta Salm-Dyck dillenii (Ker-Gawl.) Haw. engelmannii Salm-Dyck ex Engelmann subsp. lindheimeri humifusa (Raf.) Raf. subsp. humifusa** leucotricha D C. macrorhiza Engelmann subsp. macrorhiza microdasys (Lehmann) Pfeiffer subsp. microdasys phaeacantha Engelmann pubescens Wedland ex Pfeiffer rastrera Weber stenopetala Engelmann strigil Engelmann strobiliformis (Werdermann) Fric & Schelle ex Kreuzinger* aquosa(Weber) Britton lychnidiflora DC. leucocephalus (Poselger) Byles & Rowley baccifera (Muller) Stearn scheeri (Salm-Dyck) N.P. Taylor boeckmannii (Otto ex Salm-Dyck) Br.& Rose coniflorus (Wenigart) Br. & Rose spinulosus (D C.) Br. & Rose crispatus (D C.) Berger ex Hill multicostatus (Hildmann ex Shumann) Berger ex Hill subsp. multicostatus griseus (Harwoerth) Buxbaum pruinosus (Otto & Pfeiffer) Buxbaum bicolor (Backeberg) Taylor subsp. shwarzii*

350–600 400–1524 650–731 300–600 330–500 1600–2900 91–850 600–800 900–1320 850–1400 600–1402 800–1450 330–500 1500–1700 1400–2100 1200–1900 1584–1900 250–670 15–100 350–450 100–450 61–350 91–400 15–500 330–1600 1050–1189 700–1100 244–700 300–600 300–600

ALTIDUDE (m) MA 0660 MA 0054 MA 0062 MA 1134 MA 0166 MA 1010 MA 0646 MA 1222 MA 0182 MA 1254 MA 0202 MA 0928 MA 0157 MA 0307 MA 0600 MA 0879 MA 0320 PC 0207 BRAVO S/N MA 0661 MA 0658 MA 0528 MA 0649 MA 0719 MA 0161 MA 0439 MA 0774 MA 0534 MA 0877 MA 0544

VOUCHER UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT UAT MEXU MEXU UAT UAT UAT UAT MEXU UAT UAT UAT UAT UAT UAT

HERBARIUM

2500

Thelocactus Thelocactus Thelocactus Turbinicarpsu Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus

111 112 113 114 115 116 117 118

119 120

Thelocactus

110

conothelos (Regel & Klein) Knuth subsp. conothelos conothelos Glass & M. Mendoza subsp. garciae ** hexaedrophorus (Lemiare) Br. & Rose subsp. hexaedrophorus setispinus (Engelmann) Anderson tulensis (Poselger) Br. & Rose subsp. tulensis gielsdorfianus (Werdermann) John & Riha* jauernigii Gerhart Frank* nieblae Garcı´ a-Morales, Martı´ nez-Avalos ex Bergmann* ** pseudopectinatus (Backeberg) Glass & Foster* saueri (Boedeker) John & Rika subsp. saueri* saueri (Boedeker) John & Rika subsp. yisabelae* ** schmiedickeanus (Boedeker) Buxbaum & subsp. schmiedickeanus* ** viereckii (Werdermann) John & Riha subsp. viereckii* **

600–1400 1850–1900 1000–1300 10–400 800–1300 600–760 600–800 360–400 1200–1402 600–792 1100–1200 1200–1400 650–1892

MA 0357 MA 0917 MA 0318 MA 0673 MA 0353 PC 5002 MA 1153 MA 0890 MA 0351 MA 0700 MA 0657 MA 0155 MA 0150

UAT UAT UAT UAT UAT HNT UAT UAT UAT UAT UAT UAT UAT

2501

2502 Appendix 2. Index Geographical Expansion values for the cacti found in each square in the region of the state of Tamaulipas.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49

Genus

Species

Presence in square

IGE

Cylindropuntia Opuntia Ferocactus Mammillaria Echinocactus Stenocereus Echinocereus Thelocactus Sclerocactus Acanthocereus Echinocereus Mammillaria Mammillaria Selenicereus Echinocereus Pilosocereus Echinocereus Mammillaria Opuntia Cylindropuntia Escobaria Lophophora Ferocactus Selenicereus Selenicereus Echinocereus Mammillaria Echinocereus Grusonia Astrophytum Echinocereus Astrophytum Coryphantha Coryphantha Coryphantha Coryphantha Coryphantha Cylindropuntia Cylindropuntia Echinocactus Echinocereus Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Mammilloydia Marginatocerus Neolloydia

leptocaulis engelmannii subsp. lindheimeri hamatacanthus subsp. hamatacanthus heyderi subsp. heyderi texensis griseus pentalophus subsp. pentalophus setispinus scheeri tetragonus papillosus sphaerica prolifera subsp. arachnoideae spinulosus enneacantus subsp. enneacanthus leucocephalus berlandieri heyderi subsp. gummifera dejecta kleiniae emskoetteriana williamsii echidne boeckmannii coniflorus poselgeri heyderi subsp. hemiesphaerica pectinatus subsp. pectinatus shottii. asterias viereckii subsp. viereckii myriostigma georgii glanduligera guerkeana macromeris subsp. macromeris vaupeliana imbricata subsp. imbricata tunicata platyacanthus cinerascens subsp. tulensis compressa subsp. compressa formosa subsp. formosa magnimamma perbella picta subsp. picta candida subsp. candida marginatus conoidea

47 47 44 43 43 41 40 37 36 35 33 26 24 24 22 18 17 15 15 14 14 14 13 13 13 12 12 11 11 10 8 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7 7

0.979 0.979 0.936 0.915 0.915 0.872 0.851 0.787 0.766 0.745 0.702 0.553 0.511 0.511 0.468 0.383 0.362 0.319 0.319 0.298 0.298 0.298 0.277 0.277 0.277 0.255 0.255 0.234 0.234 0.213 0.170 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149 0.149

2503 Appendix 2. Continued.

50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86 87 88 89 90 91 92 93 94 95 96 97 98

Genus

Species

Presence in square

IGE

Opuntia Opuntia Opuntia Opuntia Thelocactus Ariocarpus Mammillaria Myrtillocactus Opuntia Opuntia Aricocarpus Echinocereus Echinocereus Echinomastus Ferocactus Glandulicactus Leuchtenbergia Mammillaria Thelocactus Thelocactus Turbinicarpus Coryphantha Coryphantha Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Opuntia Opuntia Opuntia Stenocereus Thelocactus Turbinicarpus Coryphantha Coryphantha Echinocereus Echinocereus Mammillaria Pereskiopsis Pereskia Ariocarpus Ariocarpus Coryphantha Coryphantha Coryphantha Echinocactus Mammillaria Mammillaria

microdasys subsp. microdasys phaeacantha rastrera stenopetala conothelos subsp. conothelos retusus subsp. retusus wintereae subsp. winteriae geometrizans humifusa subsp. humifusa strigil retusus subsp. trigonus parkeri subsp. gonzalezii stramineus subsp. stramineus gautii pilosus uncinatus subsp. uncinatus principis albicoma hexaedrophorus subsp. hexaedrophorus tulensis subsp. tulensis schmiedickeanus subsp. schmiedickeanus octacantha pulleineana compressa subsp. centralifera formosa subsp. microthele melanocentra subsp. melanocentra melanocentra subsp. rubrograndis surculosa dillenii leucotricha macrorhiza subsp. macrorhiza pruinosus bicolor subsp. shwarzii pseudopectinatus delicata nickelsiae reichenbachii subsp. reichenbachii triglochidiatus subsp. triglochidiatus heyderi subsp. meiacantha aquosa lychnidiflora agavoides kotschoubeyanus radians salinensis sulcata horizonthalonius anniana baumii

7 7 7 7 7 6 6 6 6 6 5 5 5 5 5 5 5 5 5 5 5 4 4 4 4 4 4 4 4 4 4 4 4 4 3 3 3 3 3 3 3 2 2 2 2 2 2 2 2

0.149 0.149 0.149 0.149 0.149 0.128 0.128 0.128 0.128 0.128 0.106 0.106 0.106 0.106 0.106 0.106 0.106 0.106 0.106 0.106 0.106 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.085 0.064 0.064 0.064 0.064 0.064 0.064 0.064 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043

2504 Appendix 2. Continued.

99 100 101 102 103 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134

Genus

Species

Presence in square

IGE

Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Neobuxbaumia Obregonia Opuntia Opuntia Rhipsalis Stenocactus Turbinicarpus Coryphantha Cumarinia Echinocereus Epithelantha Ferocactus Ferocactus Grusonia Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Mammillaria Pelecyphora Stenocactus Thelocactus Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus Turbinicarpus

glassii subsp. ascencionis glassii subsp. glassii klissingiana melaleuca picta subsp. viereckii pilispina roseoalba euphorbioides denegrii decumbens pubescens baccifera crispatus gielsdorfianus wohlschlageri odorata knippelianus micromeris subsp. micromeris glaucescens histrix dumetorum carmenae laui subsp. dasyacantha laui subsp. laui laui subsp. subducta plumosa prolifera subsp. zublerae schiedeana subsp. giselae strobiliformis multicostatus subsp. multicostatus conotehlos subsp. garciae jauernigii nieblae saueri subsp. saueri saueri subsp. yisabelae viereckii subsp. viereckii

2 2 2 2 2 2 2 2 2 2 2 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.043 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021 0.021

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