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Gigantic Cactus Perhaps the most extraordinary and grotesque forms in the vegetable world are to be met with in various species of Cactus, of which are seven tribes containing no less than sixty different kinds. Cactaceæ are exclusively confined to the tropics of America extend a little way north and south. When met with elsewhere they have been introduced—are not patives of the soil. The Pitahaya or Gigantic Cactus is the very Grand Master of the order. It is found in the rocky valleys and slopes of New Mexico, Arizona, and California, and is called by different names according to the language or dialect of each country. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear ; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. The Chihuahuan Desert hosts several protected areas, both at federal and state level, and most Thelocactus species can be found in some of them, although the percentage of localities occurring in protected areas is generally low (Hernández & Gómez-Hinostrosa, 2011a). The situation is worse for microendemic taxa that occur in very small areas, e.g. some T. bicolor and T. conothelos subspecies and T. hastifer, which do not occur in any protected area and for which the creation of small reserve areas was already proposed in view of its efficacy and as a complement to largest protected areas (Fos et al., 2017; Hernández & Gómez-Hinostrosa, 2011a). SDMs are the main tool to predict species distributions based on environmental suitability, and are very effective to render spatial models from sparse observations available from biological surveys and natural history collections (Franklin, 2010). They have the potential to support conservation actions and contribute to the decision-making process. SDMs may be used to identify and protect critical habitats that are necessary for the persistence of threatened species; to select areas for the establishment of reserves; to identify suitable sites for reintroduction or translocation as an aid to lessen the threat of climate changes or the impact of change of land use (Guisan et al., 2013). Most Thelocactus species can be considered vulnerable to global warming as a result of many factors like a low seed dispersal efficiency, a limited plant recruitment caused by seedling sensitivity to high temperatures (Aragón Gastélum et al., 2016), a direct effect on their physiology (Nobel, 1996), or a change in the biotic interactions, as climate change could affect also the presence of pollinators and the animals required for seed dispersal or have an effect on the vegetation community and the nurse plants belonging to it (Ibisch & Mutke, 2015). The necessity of increasing protected areas in the CDR has been already underlined and the results presented in this study could be relevant for improving preservation actions and guiding reintroduction programs for a better conservation of Thelocactus species, taking into account the ecological requirements of focal species. Niche overlap values between Thelocactus species are mostly low, reflecting the difference in the environmental suits each species is adapted to. Except for the pair hastiferleucacanthus, the niche equivalency test was rejected for all other species, showing that environmental spaces of Thelocactus species are significantly different from each other (Warren et al., 2010). The results of the niche similarity tests were quite varied, for some species pairs the niche similarities were higher than expected by chance, for others the null hypothesis was rejected, while in other species the significant results of the similarity test in only one direction, and not significant in the counter-direction, probably depend on the differences in the environmental background for the species pairs (Table 4) (Nakazato et al., 2010). The niches of the pair T. hastifer and T. leucacanthus were more similar than expected by chance, which was expected as their niches are equivalent, while when compared to the other species the similarity was not significative. Considering that these two species have also an verlapping geographical distribution, the obtained results support the hypothesis of niche conservatism. Niche similarity higher than expected by chance was also found in most of the other pair-wise comparisons, suggesting that habitat conservatism is common among Thelocactus species. Thelocactus bicolor has the widest niche breadth, therefore being capable to exploit a larger set of environmental conditions, which is in agreement with its large geographic distribution that overlaps with the range of most species, with the exclusion of the 2 southernmost, which is probably the explanation for the similarity test being significantly more similar when compared to the other species (Table 4). The reverse was not always true. For T. buekii, T. conothelos, and T. hexaedrophorus the results were not significantly similar, suggesting that these species are not suited to the habitat conditions in which T. bicolor can grow. For what concerns T. rinconensis, the similarity test was rejected when paired to T. hexaedrophorus, but the reverse comparison showed that the similarity test was accepted. These results suggest that T. rinconensis has rather different environmental requirements of T. hexaedrophorus, which exploits a more heterogeneous habitat and therefore its niche overlaps that of T. rinconensis. Five species, T. buekii, T. conothelos, T. hexaedrophorus, T. multicephalus, and T. tulensis, showed a similarity greater than expected by chance. All of them are geographically distributed in part or only in the Galeana, Mier y Noriega, and Huizache subregions of the CDR (Hernández & Bárcenas, 1996), areas rich in species number and endemicity of cacti, whose diversification is related to increased aridity in response to the uplift of the Sierra Madre Oriental and the development of the TransMexican Volcanic Belt (Vázquez-Sánchez et al., 2013) in the late Miocene (Arakaki et al., 2011; HernándezHernández et al., 2014). Pleistocene glacial maximum (Wisconsin, 11,000 years ago) brought a cooler and wetter climate affecting the areas occupied by desert communities. Climate fluctuations driven by advances and retreats of the Laurentide Continental Glacier promoted contractions, retractions and displacements of the geographic range of the species involved (Cartron et al., 2005). It has been shown that niche conservatism can be traced back to Late Pliocene Maximum, for example when the distribution range of Schiffornis turdina WiedNeuwied fragmented in many areas that correspond to possible Pleistocene refugia (Martínez-Meyer & Peterson, 2006; Peterson & Nyári, 2008). Significant ecological niche conservatism is also found for most Thelocactus species pairs, although the observed geographic ranges of Thelocactus species rarely overlap, and then mostly partially, and that species distribution is mainly allopatric. This result indicates that many Thelocactus species have conserved their ecological niche traits over time (Wiens et al., 2010), corroborating the hypothesis that the richest areas in cactus taxa in the CDR acted as glacial refugia, leading to isolation and species diversification as well as shaping actual cactus distribution (Gómez-Hinostrosa & Hernández, 2000; Hernández & Bárcenas, 1995).Gigantic Cactus

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Arizona & California Perhaps the most extraordinary and grotesque forms in the vegetable world are to be met with in various species of Cactus, of which are seven tribes containing no less than sixty different kinds. Cactaceæ are exclusively confined to the tropics of America extend a little way north and south. When met with elsewhere they have been introduced—are not patives of the soil. The Pitahaya or Gigantic Cactus is the very Grand Master of the order. It is found in the rocky valleys and slopes of New Mexico, Arizona, and California, and is called by different names according to the language or dialect of each country. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear ; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. The Chihuahuan Desert hosts several protected areas, both at federal and state level, and most Thelocactus species can be found in some of them, although the percentage of localities occurring in protected areas is generally low (Hernández & Gómez-Hinostrosa, 2011a). The situation is worse for microendemic taxa that occur in very small areas, e.g. some T. bicolor and T. conothelos subspecies and T. hastifer, which do not occur in any protected area and for which the creation of small reserve areas was already proposed in view of its efficacy and as a complement to largest protected areas (Fos et al., 2017; Hernández & Gómez-Hinostrosa, 2011a). SDMs are the main tool to predict species distributions based on environmental suitability, and are very effective to render spatial models from sparse observations available from biological surveys and natural history collections (Franklin, 2010). They have the potential to support conservation actions and contribute to the decision-making process. SDMs may be used to identify and protect critical habitats that are necessary for the persistence of threatened species; to select areas for the establishment of reserves; to identify suitable sites for reintroduction or translocation as an aid to lessen the threat of climate changes or the impact of change of land use (Guisan et al., 2013). Most Thelocactus species can be considered vulnerable to global warming as a result of many factors like a low seed dispersal efficiency, a limited plant recruitment caused by seedling sensitivity to high temperatures (Aragón Gastélum et al., 2016), a direct effect on their physiology (Nobel, 1996), or a change in the biotic interactions, as climate change could affect also the presence of pollinators and the animals required for seed dispersal or have an effect on the vegetation community and the nurse plants belonging to it (Ibisch & Mutke, 2015). The necessity of increasing protected areas in the CDR has been already underlined and the results presented in this study could be relevant for improving preservation actions and guiding reintroduction programs for a better conservation of Thelocactus species, taking into account the ecological requirements of focal species. Niche overlap values between Thelocactus species are mostly low, reflecting the difference in the environmental suits each species is adapted to. Except for the pair hastiferleucacanthus, the niche equivalency test was rejected for all other species, showing that environmental spaces of Thelocactus species are significantly different from each other (Warren et al., 2010). The results of the niche similarity tests were quite varied, for some species pairs the niche similarities were higher than expected by chance, for others the null hypothesis was rejected, while in other species the significant results of the similarity test in only one direction, and not significant in the counter-direction, probably depend on the differences in the environmental background for the species pairs (Table 4) (Nakazato et al., 2010). The niches of the pair T. hastifer and T. leucacanthus were more similar than expected by chance, which was expected as their niches are equivalent, while when compared to the other species the similarity was not significative. Considering that these two species have also an verlapping geographical distribution, the obtained results support the hypothesis of niche conservatism. Niche similarity higher than expected by chance was also found in most of the other pair-wise comparisons, suggesting that habitat conservatism is common among Thelocactus species. Thelocactus bicolor has the widest niche breadth, therefore being capable to exploit a larger set of environmental conditions, which is in agreement with its large geographic distribution that overlaps with the range of most species, with the exclusion of the 2 southernmost, which is probably the explanation for the similarity test being significantly more similar when compared to the other species (Table 4). The reverse was not always true. For T. buekii, T. conothelos, and T. hexaedrophorus the results were not significantly similar, suggesting that these species are not suited to the habitat conditions in which T. bicolor can grow. For what concerns T. rinconensis, the similarity test was rejected when paired to T. hexaedrophorus, but the reverse comparison showed that the similarity test was accepted. These results suggest that T. rinconensis has rather different environmental requirements of T. hexaedrophorus, which exploits a more heterogeneous habitat and therefore its niche overlaps that of T. rinconensis. Five species, T. buekii, T. conothelos, T. hexaedrophorus, T. multicephalus, and T. tulensis, showed a similarity greater than expected by chance. All of them are geographically distributed in part or only in the Galeana, Mier y Noriega, and Huizache subregions of the CDR (Hernández & Bárcenas, 1996), areas rich in species number and endemicity of cacti, whose diversification is related to increased aridity in response to the uplift of the Sierra Madre Oriental and the development of the TransMexican Volcanic Belt (Vázquez-Sánchez et al., 2013) in the late Miocene (Arakaki et al., 2011; HernándezHernández et al., 2014). Pleistocene glacial maximum (Wisconsin, 11,000 years ago) brought a cooler and wetter climate affecting the areas occupied by desert communities. Climate fluctuations driven by advances and retreats of the Laurentide Continental Glacier promoted contractions, retractions and displacements of the geographic range of the species involved (Cartron et al., 2005). It has been shown that niche conservatism can be traced back to Late Pliocene Maximum, for example when the distribution range of Schiffornis turdina WiedNeuwied fragmented in many areas that correspond to possible Pleistocene refugia (Martínez-Meyer & Peterson, 2006; Peterson & Nyári, 2008). Significant ecological niche conservatism is also found for most Thelocactus species pairs, although the observed geographic ranges of Thelocactus species rarely overlap, and then mostly partially, and that species distribution is mainly allopatric. This result indicates that many Thelocactus species have conserved their ecological niche traits over time (Wiens et al., 2010), corroborating the hypothesis that the richest areas in cactus taxa in the CDR acted as glacial refugia, leading to isolation and species diversification as well as shaping actual cactus distribution (Gómez-Hinostrosa & Hernández, 2000; Hernández & Bárcenas, 1995).Arizona & California

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Perhaps the most extraordinary and grotesque forms in the vegetable world are to be met with in various species of Cactus, of which are seven tribes containing no less than sixty different kinds. Cactaceæ are exclusively confined to the tropics of America extend a little way north and south. When met with elsewhere they have been introduced—are not patives of the soil. The Pitahaya or Gigantic Cactus is the very Grand Master of the order. It is found in the rocky valleys and slopes of New Mexico, Arizona, and California, and is called by different names according to the language or dialect of each country. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear ; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. The Chihuahuan Desert hosts several protected areas, both at federal and state level, and most Thelocactus species can be found in some of them, although the percentage of localities occurring in protected areas is generally low (Hernández & Gómez-Hinostrosa, 2011a). The situation is worse for microendemic taxa that occur in very small areas, e.g. some T. bicolor and T. conothelos subspecies and T. hastifer, which do not occur in any protected area and for which the creation of small reserve areas was already proposed in view of its efficacy and as a complement to largest protected areas (Fos et al., 2017; Hernández & Gómez-Hinostrosa, 2011a). SDMs are the main tool to predict species distributions based on environmental suitability, and are very effective to render spatial models from sparse observations available from biological surveys and natural history collections (Franklin, 2010). They have the potential to support conservation actions and contribute to the decision-making process. SDMs may be used to identify and protect critical habitats that are necessary for the persistence of threatened species; to select areas for the establishment of reserves; to identify suitable sites for reintroduction or translocation as an aid to lessen the threat of climate changes or the impact of change of land use (Guisan et al., 2013). Most Thelocactus species can be considered vulnerable to global warming as a result of many factors like a low seed dispersal efficiency, a limited plant recruitment caused by seedling sensitivity to high temperatures (Aragón Gastélum et al., 2016), a direct effect on their physiology (Nobel, 1996), or a change in the biotic interactions, as climate change could affect also the presence of pollinators and the animals required for seed dispersal or have an effect on the vegetation community and the nurse plants belonging to it (Ibisch & Mutke, 2015). The necessity of increasing protected areas in the CDR has been already underlined and the results presented in this study could be relevant for improving preservation actions and guiding reintroduction programs for a better conservation of Thelocactus species, taking into account the ecological requirements of focal species. Niche overlap values between Thelocactus species are mostly low, reflecting the difference in the environmental suits each species is adapted to. Except for the pair hastiferleucacanthus, the niche equivalency test was rejected for all other species, showing that environmental spaces of Thelocactus species are significantly different from each other (Warren et al., 2010). The results of the niche similarity tests were quite varied, for some species pairs the niche similarities were higher than expected by chance, for others the null hypothesis was rejected, while in other species the significant results of the similarity test in only one direction, and not significant in the counter-direction, probably depend on the differences in the environmental background for the species pairs (Table 4) (Nakazato et al., 2010). The niches of the pair T. hastifer and T. leucacanthus were more similar than expected by chance, which was expected as their niches are equivalent, while when compared to the other species the similarity was not significative. Considering that these two species have also an verlapping geographical distribution, the obtained results support the hypothesis of niche conservatism. Niche similarity higher than expected by chance was also found in most of the other pair-wise comparisons, suggesting that habitat conservatism is common among Thelocactus species. Thelocactus bicolor has the widest niche breadth, therefore being capable to exploit a larger set of environmental conditions, which is in agreement with its large geographic distribution that overlaps with the range of most species, with the exclusion of the 2 southernmost, which is probably the explanation for the similarity test being significantly more similar when compared to the other species (Table 4). The reverse was not always true. For T. buekii, T. conothelos, and T. hexaedrophorus the results were not significantly similar, suggesting that these species are not suited to the habitat conditions in which T. bicolor can grow. For what concerns T. rinconensis, the similarity test was rejected when paired to T. hexaedrophorus, but the reverse comparison showed that the similarity test was accepted. These results suggest that T. rinconensis has rather different environmental requirements of T. hexaedrophorus, which exploits a more heterogeneous habitat and therefore its niche overlaps that of T. rinconensis. Five species, T. buekii, T. conothelos, T. hexaedrophorus, T. multicephalus, and T. tulensis, showed a similarity greater than expected by chance. All of them are geographically distributed in part or only in the Galeana, Mier y Noriega, and Huizache subregions of the CDR (Hernández & Bárcenas, 1996), areas rich in species number and endemicity of cacti, whose diversification is related to increased aridity in response to the uplift of the Sierra Madre Oriental and the development of the TransMexican Volcanic Belt (Vázquez-Sánchez et al., 2013) in the late Miocene (Arakaki et al., 2011; HernándezHernández et al., 2014). Pleistocene glacial maximum (Wisconsin, 11,000 years ago) brought a cooler and wetter climate affecting the areas occupied by desert communities. Climate fluctuations driven by advances and retreats of the Laurentide Continental Glacier promoted contractions, retractions and displacements of the geographic range of the species involved (Cartron et al., 2005). It has been shown that niche conservatism can be traced back to Late Pliocene Maximum, for example when the distribution range of Schiffornis turdina WiedNeuwied fragmented in many areas that correspond to possible Pleistocene refugia (Martínez-Meyer & Peterson, 2006; Peterson & Nyári, 2008). Significant ecological niche conservatism is also found for most Thelocactus species pairs, although the observed geographic ranges of Thelocactus species rarely overlap, and then mostly partially, and that species distribution is mainly allopatric. This result indicates that many Thelocactus species have conserved their ecological niche traits over time (Wiens et al., 2010), corroborating the hypothesis that the richest areas in cactus taxa in the CDR acted as glacial refugia, leading to isolation and species diversification as well as shaping actual cactus distribution (Gómez-Hinostrosa & Hernández, 2000; Hernández & Bárcenas, 1995).Perhaps

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Diversification Perhaps the most extraordinary and grotesque forms in the vegetable world are to be met with in various species of Cactus, of which are seven tribes containing no less than sixty different kinds. Cactaceæ are exclusively confined to the tropics of America extend a little way north and south. When met with elsewhere they have been introduced—are not patives of the soil. The Pitahaya or Gigantic Cactus is the very Grand Master of the order. It is found in the rocky valleys and slopes of New Mexico, Arizona, and California, and is called by different names according to the language or dialect of each country. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear ; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. The Chihuahuan Desert hosts several protected areas, both at federal and state level, and most Thelocactus species can be found in some of them, although the percentage of localities occurring in protected areas is generally low (Hernández & Gómez-Hinostrosa, 2011a). The situation is worse for microendemic taxa that occur in very small areas, e.g. some T. bicolor and T. conothelos subspecies and T. hastifer, which do not occur in any protected area and for which the creation of small reserve areas was already proposed in view of its efficacy and as a complement to largest protected areas (Fos et al., 2017; Hernández & Gómez-Hinostrosa, 2011a). SDMs are the main tool to predict species distributions based on environmental suitability, and are very effective to render spatial models from sparse observations available from biological surveys and natural history collections (Franklin, 2010). They have the potential to support conservation actions and contribute to the decision-making process. SDMs may be used to identify and protect critical habitats that are necessary for the persistence of threatened species; to select areas for the establishment of reserves; to identify suitable sites for reintroduction or translocation as an aid to lessen the threat of climate changes or the impact of change of land use (Guisan et al., 2013). Most Thelocactus species can be considered vulnerable to global warming as a result of many factors like a low seed dispersal efficiency, a limited plant recruitment caused by seedling sensitivity to high temperatures (Aragón Gastélum et al., 2016), a direct effect on their physiology (Nobel, 1996), or a change in the biotic interactions, as climate change could affect also the presence of pollinators and the animals required for seed dispersal or have an effect on the vegetation community and the nurse plants belonging to it (Ibisch & Mutke, 2015). The necessity of increasing protected areas in the CDR has been already underlined and the results presented in this study could be relevant for improving preservation actions and guiding reintroduction programs for a better conservation of Thelocactus species, taking into account the ecological requirements of focal species. Niche overlap values between Thelocactus species are mostly low, reflecting the difference in the environmental suits each species is adapted to. Except for the pair hastiferleucacanthus, the niche equivalency test was rejected for all other species, showing that environmental spaces of Thelocactus species are significantly different from each other (Warren et al., 2010). The results of the niche similarity tests were quite varied, for some species pairs the niche similarities were higher than expected by chance, for others the null hypothesis was rejected, while in other species the significant results of the similarity test in only one direction, and not significant in the counter-direction, probably depend on the differences in the environmental background for the species pairs (Table 4) (Nakazato et al., 2010). The niches of the pair T. hastifer and T. leucacanthus were more similar than expected by chance, which was expected as their niches are equivalent, while when compared to the other species the similarity was not significative. Considering that these two species have also an verlapping geographical distribution, the obtained results support the hypothesis of niche conservatism. Niche similarity higher than expected by chance was also found in most of the other pair-wise comparisons, suggesting that habitat conservatism is common among Thelocactus species. Thelocactus bicolor has the widest niche breadth, therefore being capable to exploit a larger set of environmental conditions, which is in agreement with its large geographic distribution that overlaps with the range of most species, with the exclusion of the 2 southernmost, which is probably the explanation for the similarity test being significantly more similar when compared to the other species (Table 4). The reverse was not always true. For T. buekii, T. conothelos, and T. hexaedrophorus the results were not significantly similar, suggesting that these species are not suited to the habitat conditions in which T. bicolor can grow. For what concerns T. rinconensis, the similarity test was rejected when paired to T. hexaedrophorus, but the reverse comparison showed that the similarity test was accepted. These results suggest that T. rinconensis has rather different environmental requirements of T. hexaedrophorus, which exploits a more heterogeneous habitat and therefore its niche overlaps that of T. rinconensis. Five species, T. buekii, T. conothelos, T. hexaedrophorus, T. multicephalus, and T. tulensis, showed a similarity greater than expected by chance. All of them are geographically distributed in part or only in the Galeana, Mier y Noriega, and Huizache subregions of the CDR (Hernández & Bárcenas, 1996), areas rich in species number and endemicity of cacti, whose diversification is related to increased aridity in response to the uplift of the Sierra Madre Oriental and the development of the TransMexican Volcanic Belt (Vázquez-Sánchez et al., 2013) in the late Miocene (Arakaki et al., 2011; HernándezHernández et al., 2014). Pleistocene glacial maximum (Wisconsin, 11,000 years ago) brought a cooler and wetter climate affecting the areas occupied by desert communities. Climate fluctuations driven by advances and retreats of the Laurentide Continental Glacier promoted contractions, retractions and displacements of the geographic range of the species involved (Cartron et al., 2005). It has been shown that niche conservatism can be traced back to Late Pliocene Maximum, for example when the distribution range of Schiffornis turdina WiedNeuwied fragmented in many areas that correspond to possible Pleistocene refugia (Martínez-Meyer & Peterson, 2006; Peterson & Nyári, 2008). Significant ecological niche conservatism is also found for most Thelocactus species pairs, although the observed geographic ranges of Thelocactus species rarely overlap, and then mostly partially, and that species distribution is mainly allopatric. This result indicates that many Thelocactus species have conserved their ecological niche traits over time (Wiens et al., 2010), corroborating the hypothesis that the richest areas in cactus taxa in the CDR acted as glacial refugia, leading to isolation and species diversification as well as shaping actual cactus distribution (Gómez-Hinostrosa & Hernández, 2000; Hernández & Bárcenas, 1995).Diversification

Micro Regular

Sample

Perhaps the most extraordinary and grotesque forms in the vegetable world are to be met with in various species of Cactus, of which are seven tribes containing no less than sixty different kinds. Cactaceæ are exclusively confined to the tropics of America extend a little way north and south. When met with elsewhere they have been introduced—are not patives of the soil. The Pitahaya or Gigantic Cactus is the very Grand Master of the order. It is found in the rocky valleys and slopes of New Mexico, Arizona, and California, and is called by different names according to the language or dialect of each country. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear ; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. For the first few years of its existence it is globular, then it shoots up for ten or twelve feet and blossoms, then the trunk or stem shoots out again, and frequently rises to the height of sixty feet. It has few branches, but these are generally covered with flowers, which are clustered together. The seed-vessel or fruit, which falls to the ground in clusters in July and August, is in the shape of a reddish green pear; inside is a rich crimson pulp, which tastes like al fresh fig, is nutritious and much valued by natives and others. The Night Blooming Cereus has a singular different from all others of its family. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. During the summer months it begins to open flowers between seven and eight o’clock in the evening. At eleven they are fully expanded, and while thus they emit the most fragrance, which in their native home perfumes the air to a considerable distance. Each flower when open is nine inches in diameter, the inside is a splendid yellow colour, resembling the rays of a star, the stamens are a pure white, which adds to the illusion. The outside of the flower is brown. The flowers close between three and four o’clock in the morning, and rapidly decay. This cactus is represented in the centre of the small engraving. Another very rare and curious species is the Cactus Senilis, or Old Man’s Head. This is globular, and entirely covered with long, hair like filaments of a greyish colour, which hang down like grey hair. The Chihuahuan Desert hosts several protected areas, both at federal and state level, and most Thelocactus species can be found in some of them, although the percentage of localities occurring in protected areas is generally low (Hernández & Gómez-Hinostrosa, 2011a). The situation is worse for microendemic taxa that occur in very small areas, e.g. some T. bicolor and T. conothelos subspecies and T. hastifer, which do not occur in any protected area and for which the creation of small reserve areas was already proposed in view of its efficacy and as a complement to largest protected areas (Fos et al., 2017; Hernández & Gómez-Hinostrosa, 2011a). SDMs are the main tool to predict species distributions based on environmental suitability, and are very effective to render spatial models from sparse observations available from biological surveys and natural history collections (Franklin, 2010). They have the potential to support conservation actions and contribute to the decision-making process. SDMs may be used to identify and protect critical habitats that are necessary for the persistence of threatened species; to select areas for the establishment of reserves; to identify suitable sites for reintroduction or translocation as an aid to lessen the threat of climate changes or the impact of change of land use (Guisan et al., 2013). Most Thelocactus species can be considered vulnerable to global warming as a result of many factors like a low seed dispersal efficiency, a limited plant recruitment caused by seedling sensitivity to high temperatures (Aragón Gastélum et al., 2016), a direct effect on their physiology (Nobel, 1996), or a change in the biotic interactions, as climate change could affect also the presence of pollinators and the animals required for seed dispersal or have an effect on the vegetation community and the nurse plants belonging to it (Ibisch & Mutke, 2015). The necessity of increasing protected areas in the CDR has been already underlined and the results presented in this study could be relevant for improving preservation actions and guiding reintroduction programs for a better conservation of Thelocactus species, taking into account the ecological requirements of focal species. Niche overlap values between Thelocactus species are mostly low, reflecting the difference in the environmental suits each species is adapted to. Except for the pair hastiferleucacanthus, the niche equivalency test was rejected for all other species, showing that environmental spaces of Thelocactus species are significantly different from each other (Warren et al., 2010). The results of the niche similarity tests were quite varied, for some species pairs the niche similarities were higher than expected by chance, for others the null hypothesis was rejected, while in other species the significant results of the similarity test in only one direction, and not significant in the counter-direction, probably depend on the differences in the environmental background for the species pairs (Table 4) (Nakazato et al., 2010). The niches of the pair T. hastifer and T. leucacanthus were more similar than expected by chance, which was expected as their niches are equivalent, while when compared to the other species the similarity was not significative. Considering that these two species have also an verlapping geographical distribution, the obtained results support the hypothesis of niche conservatism. Niche similarity higher than expected by chance was also found in most of the other pair-wise comparisons, suggesting that habitat conservatism is common among Thelocactus species. Thelocactus bicolor has the widest niche breadth, therefore being capable to exploit a larger set of environmental conditions, which is in agreement with its large geographic distribution that overlaps with the range of most species, with the exclusion of the 2 southernmost, which is probably the explanation for the similarity test being significantly more similar when compared to the other species (Table 4). The reverse was not always true. For T. buekii, T. conothelos, and T. hexaedrophorus the results were not significantly similar, suggesting that these species are not suited to the habitat conditions in which T. bicolor can grow. For what concerns T. rinconensis, the similarity test was rejected when paired to T. hexaedrophorus, but the reverse comparison showed that the similarity test was accepted. These results suggest that T. rinconensis has rather different environmental requirements of T. hexaedrophorus, which exploits a more heterogeneous habitat and therefore its niche overlaps that of T. rinconensis. Five species, T. buekii, T. conothelos, T. hexaedrophorus, T. multicephalus, and T. tulensis, showed a similarity greater than expected by chance. All of them are geographically distributed in part or only in the Galeana, Mier y Noriega, and Huizache subregions of the CDR (Hernández & Bárcenas, 1996), areas rich in species number and endemicity of cacti, whose diversification is related to increased aridity in response to the uplift of the Sierra Madre Oriental and the development of the TransMexican Volcanic Belt (Vázquez-Sánchez et al., 2013) in the late Miocene (Arakaki et al., 2011; HernándezHernández et al., 2014). Pleistocene glacial maximum (Wisconsin, 11,000 years ago) brought a cooler and wetter climate affecting the areas occupied by desert communities. Climate fluctuations driven by advances and retreats of the Laurentide Continental Glacier promoted contractions, retractions and displacements of the geographic range of the species involved (Cartron et al., 2005). It has been shown that niche conservatism can be traced back to Late Pliocene Maximum, for example when the distribution range of Schiffornis turdina WiedNeuwied fragmented in many areas that correspond to possible Pleistocene refugia (Martínez-Meyer & Peterson, 2006; Peterson & Nyári, 2008). Significant ecological niche conservatism is also found for most Thelocactus species pairs, although the observed geographic ranges of Thelocactus species rarely overlap, and then mostly partially, and that species distribution is mainly allopatric. This result indicates that many Thelocactus species have conserved their ecological niche traits over time (Wiens et al., 2010), corroborating the hypothesis that the richest areas in cactus taxa in the CDR acted as glacial refugia, leading to isolation and species diversification as well as shaping actual cactus distribution (Gómez-Hinostrosa & Hernández, 2000; Hernández & Bárcenas, 1995).Perhaps

TypefaceThelo

DesignerTassiana Nuñez Costa

Release 2020

Fonts In Use

Specimen

9 Styles

Roman

Italic

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Thelo in Use