Article

Molecular phylogenetics of Echinopsis (Cactaceae): Polyphyly at all levels and convergent evolution of pollination modes and growth forms

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Abstract

Premise of the study: In its current circumscription, Echinopsis with 100-150 species is one of the largest and morphologically most diverse genera of Cactaceae. This diversity and an absence of correlated characters have resulted in numerous attempts to subdivide Echinopsis into more homogeneous subgroups. To infer natural species groups in this alliance, we here provide a plastid phylogeny and use it to infer changes in growth form, pollination mode, and ploidy level. Methods: We sequenced 3800 nucleotides of chloroplast DNA from 162 plants representing 144 species and subspecies. The sample includes the type species of all genera close to, or included in, Echinopsis as well as a dense sample of other genera of the Trichocereeae and further outgroups. New and published chromosome counts were compiled and traced on the phylogeny, as were pollination modes and growth habits. Key results: A maximum likelihood phylogeny confirms that Echinopsis s.l. is not monophyletic nor are any of the previously recognized genera that have more than one species. Pollination mode and, to a lesser extent, growth habit are evolutionarily labile, and diploidy is the rule in Echinopsis s.l., with the few polyploids clustered in just a few clades. Conclusions: The use of evolutionary labile floral traits and growth habit has led to nonnatural classifications. Taxonomic realignments are required, but further study of less evolutionary labile traits suitable for circumscribing genera are needed. Surprisingly, polyploidy seems infrequent in the Echinopsis alliance and hybridization may thus be of minor relevance in the evolution of this clade.

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... As already reported (Anceschi & Magli 2018, 36: 74), since Wallace's study (1995, 13: 1-12), over the last decades, changes at the genericus level and the higher taxa in the family Cactaceae, have almost always been followed by new evidence emerging from molecular analysis. Examples are Nyffeler (1999); Nyffeler & Eggli (2010) ;Schlumpberger & Renner (2012); Schlumpberger (2012); Charles (2012); Anceschi & Magli (2013a, 2013b; Hunt (2013); Lodé (2015). In the creation of their taxonomic systems, all these authors invoke the principle of monophyly to support the formation of their groups (even with opposite results), as opposed to the principles of paraphyly and polyphyly. ...
... data & Nyffeler et al. umpubl. data in Nyffeler & Eggli 2010;Schlumpberger & Renner 2012;Anceschi & Magli 2013a, 2013b). Molecular evidence also shows just as clearly that the most part of the genera of the Trichocereeae/Trichocereinae (Schlumpberger & Renner 2012;Anceschi & Magli 2013a, 2013b, are part of a well-supported monophyletic macrogenus Echinopsis s.l. ...
... data in Nyffeler & Eggli 2010;Schlumpberger & Renner 2012;Anceschi & Magli 2013a, 2013b). Molecular evidence also shows just as clearly that the most part of the genera of the Trichocereeae/Trichocereinae (Schlumpberger & Renner 2012;Anceschi & Magli 2013a, 2013b, are part of a well-supported monophyletic macrogenus Echinopsis s.l. (i.e. ...
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After returning from our sixth South American trip (16 Nov. 2015/2 Aug. 2016), in a cactusinhabitat.org News dated September 2016, we hoped to update the website with the materials of the last two trips (2013-2014, 2015-2016), for the beginning of 2018. In reality, the processing of the huge amount of data collected in the habitats, in addition to the publication of some wide-ranging articles, namely the synopsis of Parodia Spegazzini s.l., which appeared in Bradleya (2018, 36: 70-161), and the two articles on the genera Parodia s.l. and Echinopsis s.l., created for the special issue of CactusWorld dedicated to South American Cacti (2020, Vol. 38 Special Issue: 1-52), made us postpone the deadlines. … The present publication brings to 346 the accepted taxa at the specific level and to 46 those at the generic level, compared to 252 and 40 respectively considered in the previous 2013 edition. Through the molecular outcomes we are aware (Nyffeler & Eggli 2010; Barcenas et al. 2011; Schlumpberger & Renner 2012), that most of the genera still recognized within the family Cactaceae de Jussieu simply do not exist in phylogenetic terms, as the morphological differences still used to distinguish them do not correspond to real differences at the genetic level (Nyffeler & Eggli 2010). In this sense, to the cases of Echinopsis s.l., Parodia s.l., Eriosyce s.l., already treated in the previous booklets (Anceschi & Magli 2010, 2013), based on the evidences of Franco et al. (2017) we now also add Cereus s.l. to the macrogenera considered, with the already suspected inclusion of Cipocereus F. Ritter and Praecereus Buxbaum, the second already assimilated by us in Cereus in 2013. The 608 new surveys conducted during the last two trips are documented by 6300 photos, which together with the previous 6500, bring the total images illustrating the taxa in cactusinhabitat.org to 12800. In the main text of the booklet, Taxonomy (part III), we trace some guidelines which, retracing the milestones of Western philosophical and scientific thought, lead to the current propensity of human mind, in its definitions about something approximately true in nature, to a predisposition to proceed always and only through inductive methods aimed at dissecting the real, and not to the understanding of a totality of the same reality through a deductive and unifying method. … , we remember that this approach comes from afar. Precisely by the Fathers of Western culture, philosophical, scientific, poetic, ethical, political, etc: the ancient Greeks. Especially with Plato's and Aristotle's approach to knowledge, who, although aware of what the deductive method was, were both "spitters" in their understanding of the world. That is, the sensible being (in Plato and Aristotle) and the intelligible Being (in Plato), were constituted (albeit in a different way in the two doctrines), by a "many". While Parmenides and Plotinus, with their most integral and univocal visions of Being, were the progenitors of a more unifying although anti-phenomenal approach to reality. Through the transition from the Aristotelian qualitative science, to the quantitative one, sustained in the early 1600s by Descartes, Mersenne, and of course Galileo and exemplified by the Baconian principle of the “dissectio naturae”, that “it is better to dissect than to abstract nature” [melius autem est natura secare, quam abstrahere] (Bacon 1620, book 1, section 51), we tried to retrace the achievements gained by this interpretation of reality. Namely the Newton's physics first, and then the achievements of the men of quantum mechanics, remembering that the two great contemporary technological revolutions, that of the transistor (1948) and the laser (1950 c.), they are both progeny of the second. Continuing, we also point out the current "impasse" due to the exclusive use of this approach, highlighting also its subsequent defeats (the string theory, “a theory of everything", the wandering for an inclusion of human mind as part of the measuring apparatus in quantum measurement etc.), and the "divertissements" (such as the search for "exoplanets", for example), sustaining the intelligence fundamental unifying value in the approach to knowledge. Returning to taxonomic science, and to the paradigms that regulate its current use, we point out the limit constituted by the predilection given to the sense of sight in the understanding of the sensible reality that surrounds us, emphasizing that this predilection often leads us to surface interpretations. The long journeys conducted through the most arid and semi-arid ecosystems on the planet, have made us aware that species are not interested in maintaining an identity through reproductive barriers, but they simply want to continue to exist or to be, transforming to each other in space and time through reproduction and crossing. On the basis of genetic arguments, we underline how a taxonomic science with a more universal vision, can help us to overcome together with many useless names that reassure us so much, also as many useless barriers, in the direction of a more empathic and ethical understanding of the world. Coming back to what we would like, it should be the approach to knowledge, ontological in general and scientific in particular, we believe that true science is based on the intuition of the principles and not on inductive methods, probabilities supported by "solid" mathematical quantities, opinion and relative consensus, i.e. the paradigms dear to contemporary epistemology. To substantiate our hypothesis, we bring the testimonies of three great men: Aristotele (Aristotle, Posterior Analytics, II, 19, 100 b), Albert Einstein (Einstein 1936) and Willi Hennig (Hennig 1966, 128-129), convinced supporters of the fundamental value of intuition as the "principle of principle" (Aristotle, ibidem), of the scientific procedure. In the conclusions we formulate as a proposal for a preparation for a new method of approach to scientific knowledge, a return to a way of proceeding that favours theoretical-speculative thinking as the basis for understanding reality. An invitation to grasp the visible through reasoning, and the invisible through intuition. For this purpose, a re-reading of the Classics of Western philosophy, also by scientists, physicists included, would probably be a good starting point.
... Anderson (2001Anderson ( , 2005, Hoffmann & Walter (2004) and Hunt & al. (2006Hunt & al. ( , 2013 adopted the broad generic concept, but somewhat modified the classification at specific and infrageneric (using "subgenus" or "group" instead of sections and subsections) levels. Furthermore, morphological cladistics analyses gave support to the broad circumscription of Eriosyce (Wallace in Kattermann, 1994;Nyffeler & Eggli, 1997), but since vegetative and reproductive morphological characters in Cactaceae have been documented as convergent, these results must be contrasted with molecular data (Schlumpberger & Renner, 2012). ...
... On the other hand, the two sympatrically growing species E. heinrichiana and E. simulans from Kattermann's subsection Horridocactus were placed within the Neoporteria clade in our analyses, although their flowers are funnel-form and bee-pollinated. This result might be considered as another example for the finding that closely related species may have different pollination syndromes, as floral characters and pollination syndromes in cacti are evolutionary labile (Nyffeler & Eggli, 2010;Schlumpberger & Renner, 2012). The three northernmost distributed taxa in the clade, E. villosa (Monv.) ...
... Former hypotheses of the infrageneric classification of Eriosyce were based on morphological characters alone, and were shown to be non-natural by our results. Convergent evolution of species living in similar habitats and potential hybridization might be the reason why morphology-based classifications are not always compatible with molecular phylogenetic hypotheses (Ritz & al., 2007;Guerrero & al., 2011b;Schlumpberger & Renner, 2012). ...
... Echinopsis s.l. includes more than 100 species, and exhibits a great diversity in architecture and in pollination systems (bee, hummingbird, or sphingid pollinated) (Schlumpberger and Renner 2012). Molecular phylogenies have shown Echinopsis s.l. to be nonmonophyletic (Schlumpberger and Renner 2012;Ritz et al. 2007). ...
... includes more than 100 species, and exhibits a great diversity in architecture and in pollination systems (bee, hummingbird, or sphingid pollinated) (Schlumpberger and Renner 2012). Molecular phylogenies have shown Echinopsis s.l. to be nonmonophyletic (Schlumpberger and Renner 2012;Ritz et al. 2007). In this genus, distant species may exhibit similar morphological traits, while close relatives may present very different morphological traits (Schlumpberger and Renner 2012;Ritz et al. 2007). ...
... Molecular phylogenies have shown Echinopsis s.l. to be nonmonophyletic (Schlumpberger and Renner 2012;Ritz et al. 2007). In this genus, distant species may exhibit similar morphological traits, while close relatives may present very different morphological traits (Schlumpberger and Renner 2012;Ritz et al. 2007). Interestingly, morphological homoplasy was triggered by convergence to similar selective forces rather than hybridization, which seems to be rare in the group (Schlumpberger and Renner 2012). ...
Article
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Members of the cactus family are keystone species of arid and semiarid biomes in the Americas, as they provide shelter and resources to support other members of ecosystems. Extraordinary examples are the several species of flies of the genus Drosophila that lay eggs and feed in their rotting stems, which provide a model system for studying evolutionary processes. Although there is significant progress in understanding the evolution of Drosophila species, there are gaps in our knowledge about the cactus lineages hosting them. Here we review the current knowledge about the evolution of Cactaceae, focusing on phylogenetic relationships and trends revealed by the study of DNA sequence data. During the last several decades, the availability of molecular phylogenies has considerably increased our understanding of the relationships, biogeography, and evolution of traits in the family. Remarkably, although succulent cacti have very low growth rates and long generation times, they underwent some of the fastest diversifications observed in the plant kingdom, possibly fostered by strong ecological interactions. We have a better understanding of the reproductive biology, population structure and speciation mechanisms in different clades. The recent publication of complete genomes for some species has revealed the importance of phenomena such as incomplete lineage sorting. Hybridization and polyploidization are common in the family, and have been studied using a variety of phylogenetic methods. We discuss potential future avenues for research in Cactaceae, emphasizing the need of a concerted effort among scientists in the Americas, together with the analyses of data from novel sequencing techniques.
... The more a splitter's classification is used, the more likely hybrids between species from closely related genera would be classified as intergeneric, while the very same combination of parent species becomes an intrageneric hybrid within the concept of a lumper's classification. The genus Echinopsis s.l. is a good example - Hunt & al. (2006) and Anderson & Eggli (2011) use a wide circumscription (hence Echinopsis s.l.), while the recent molecular study of Schlumpberger & Renner (2012) finds Echinopsis s.l. as both polyphyletic and vastly paraphyletic relative to the majority of the remaining genera of subtribe Trichocereinae, and at least half a dozen clades of their phylogeny would have to be recognized at generic level to arrive at monophyletic units (see Schlumpberger 2012 for the necessary new combinations). ...
... published data) between the suspected parents (Table 1), and the flowers also combine characters of both putative parent species (Fig. 9). In the currently preferred classification of Anderson & Eggli (2011) and Hunt & al. (2006), the plants conform to an intrageneric Echinopsis hybrid, while when applying the classification suggested by Schlumpberger & Renner (2012), it would be a Leucostele × Soehrensia hybrid, viz. a hybrid between two rather distantly related clades within Echinopsis s.l. in the phylogeny suggested by these authors (Schlumpberger & Renner 2012). This could explain its sterility. ...
... published data) between the suspected parents (Table 1), and the flowers also combine characters of both putative parent species (Fig. 9). In the currently preferred classification of Anderson & Eggli (2011) and Hunt & al. (2006), the plants conform to an intrageneric Echinopsis hybrid, while when applying the classification suggested by Schlumpberger & Renner (2012), it would be a Leucostele × Soehrensia hybrid, viz. a hybrid between two rather distantly related clades within Echinopsis s.l. in the phylogeny suggested by these authors (Schlumpberger & Renner 2012). This could explain its sterility. ...
Article
A solitary plant of the putative intercladal cross Echinopsis (Leucostele) atacamensis subsp. atacamensis × Echinopsis (Soehrensia) formosa was observed in the contact zone of the mostly allopatric populations of the parent species in the Salar de Atacama region in the Chilean Andes. The plant is intermediate between the parents in its characters. It was found to be sterile, despite normal-looking anthers (with abundant pollen) and stigma. In addition, an apparently typical individual of E. atacamensis with yellow flowers has been found, but it remains unknown whether this is merely a flower colour variant, or whether hybridization could be involved. Resumen: Se observó un individuo solitario de un supuesto cruce intercladal entre Echinopsis (Leucostele) atacamensis subsp. atacamensis × Echinopsis (Soehrensia) formosa en la zona de solape entre las poblaciones principalmente alopátricas de las especies progenitoras, cerca del Salar de Atacama en la zona norte de Chile. De acuerdo a sus características, la planta es morfológicamente intermedia entre sus progenitores y es estéril, pese al polen y al estigma de aspecto normal. Además se encontró un individuo aparentemente normal de E. atacamensis, pero con flores de color amarillo. Sin embargo no se pudo establecer en que medida se trata simplemente de una variación floral caprichosa no más o si está implicada alguna hibridización.
... constitutes a well-defined lineage within the angiosperm order Caryophyllales Berchtold & J. Presl (see e.g., Brockington et al. 2015, Cuénoud et al. 2002, Schäferhoff et al. 2009). The general understanding of evolutionary relationships in Cactaceae has improved in recent years as a result of molecular phylogenetic studies (Hernández-Hernández et al. 2011, Korotkova et al. 2011, Korotkova et al. 2010, Nyffeler 2002, Nyffeler & Eggli 2010, Schlumpberger & Renner 2012, Vázquez-Sánchez et al. 2013. Nevertheless, parts of the Cactaceae phylogenetic tree remain to be resolved, in particular concerning the relationships of major clades and relationships at the species level. ...
... Cactaceae show complex patterns of convergent evolution in life forms, pollination syndromes and other traits (e.g. Gibson & Nobel 1986, Hernández-Hernández et al. 2011, Schlumpberger & Renner 2012. The obvious morphological characters and their states associated with these traits have frequently been used for diagnosing genera but they are often homoplastic and genera based on those characters are therefore often shown as not monophyletic. ...
... The most recent taxonomic backbone of Caryophyllales that summarizes the current understanding of genus concepts in this order still had to accept several poly-or paraphyletic genera in Cactaceae (Hernández-Ledesma et al. 2015). Striking examples are Echinopsis Zuccarini (Anceschi & Magli 2013, Schlumpberger & Renner 2012 or Ferocactus Britton & Rose (Vázquez-Sánchez et al. 2013). ...
Article
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The tribe Hylocereeae are represented by mainly Central American-Mexican epiphytic, hemi-epiphytic and climbing cacti. They are popular due to their spectacular nocturnal flowers and have some importance as crops grown for their edible fruits. We present the first comprehensive phylogenetic study of the Hylocereeae sampling 60 out of the 63 currently accepted species and 17 out of 19 infraspecific taxa. Based on four plastid regions (trnK/matK, the rpl16 intron, rps3-rpl16, and trnL-F) we find a highly supported core Hylocereeae clade that also includes Acanthocereus and Peniocereus p.p., while Strophocactus is depicted as polyphyletic and is resolved outside of the Hylocereeae tribe. The clades found within Hylocereeae agree, in general terms, with the currently accepted genera but none of the genera are entirely monophyletic in their current circumscription. A new concept for the Hylocereeae is presented to include the genera Acanthocereus (incl. Peniocereus p.p.), Aporocactus, Disocactus, Epiphyllum, Selenicereus (incl. Hylocereus and Weberocereus p.p.), Pseudorhipsalis, Kimnachia gen. nov., and Weberocereus. New nomenclatural combinations are provided to make these genera monophyletic. The genus Deamia is reinstated for Strophocactus testudo and S. chontalensis, while Strophocactus is newly circumscribed to include S. wittii, Pseudoacanthocereus brasiliensis, and P. sicariguensis. Both genera are excluded from Hylocereeae. A taxonomic synopsis of Hylocereeae is provided.
... In 2001, Anderson published what has become the most traditionally used treatment based on the consensus classification of the Cactaceae Working Party of the International Organization for Succulent Plant Study (IOS) (Hunt andTaylor 1986, 1990), which presented the tribe Cereeae comprising 11 genera (Table 1). More recent molecular phylogenetic treatments of Cactaceae have produced evidence pointing to a non-monophyletic Cereinae as currently circumscribed; however, these studies included an extremely reduced sampling of the group (Applequist and Wallace 2002;Nyffeler 2002;Crozier 2005;Ritz et al. 2007;B arcenas et al. 2011;Hern andez-Hern andez et al. 2011;Schlumpberger and Renner 2012;Lendel 2013), leading all to question if a more comprehensive analysis would recover the same results. ...
... Previous phylogenetic studies using different samplings of taxa have indicated a non-monophyletic Cereinae (Applequist Nyffeler 2002;Crozier 2005;Ritz et al. 2007;B arcenas et al. 2011;Hern andez-Hern andez et al. 2011;Schlumpberger and Renner 2012;Lendel 2013;Soffiatti 2003). Most of those studies included Stetsonia coryne (Salm-Dyck) Britton & Rose and Espostoopsis dybowski (Rol.-Goss.) ...
Article
Cereinae comprises 14 genera distributed in Neotropical dry forest formations such as in the Caatingas of northeastern Brazil or in rocky outcrops in the north of southeastern Brazil. Historically, the taxonomy of the group has been very controversial, especially regarding generic circumscriptions, and phylogenetic relationships within the group are still poorly understood. To investigate the delimitation of thesubtribe and infra-subtribal relationships, we performed a phylogenetic analysis including 50 taxa representing 13 genera using one nuclear ( PhyC ) and four cpDNA ( petL-psbE , trnL-trnT , trnS-trnG , and rpl16 ) regions. Our results show a monophyletic Cereinae with high support in Bayesian, maximum parsimony, and maximum likelihood analyses based on combined matrices. Although our results expand the knowledge of generic relationships, we emphasize the need for further molecular phylogenetic studies combined with ecological evidence to clarify relationshipsat the more inclusive nodes of the subtribe.
... Several family-wide phylogenetic studies have been published (Nyffeler 2002;Nyffeler & Eggli 2010;Bárcenas & al. 2011;Guerrero & al. 2019a), representatives of most Cactaceae genera have by now been included in detailed molecular phylogenetic analyses, including some of the most speciose clades of Cactaceae such as Echinopsis Zucc. (Schlumpberger & Renner 2012), Mammillaria Haw. (Breslin & al. 2021) and Opuntia Mill. ...
... And for still other genera, the phylogenetic results have so far remained inconclusive for various reasons, mostly attributable to insufficient taxon sampling or low node support. Examples include some genera in Cacteae (Vázquez-Sánchez & al. 2013) or the Echinopsis alliance (Schlumpberger & Renner 2012). Final conclusions regarding the monophyly and generic limits of Coryphantha (Engelm.) ...
Article
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This data paper presents a largely phylogeny-based online taxonomic backbone for the Cactaceae compiled from literature and online sources using the tools of the EDIT Platform for Cybertaxonomy. The data will form a contribution of the Caryophyllales Network for the World Flora Online and serve as the base for further integration of research results from the systematic research community. The final aim is to treat all effectively published scientific names in the family. The checklist includes 150 accepted genera, 1851 accepted species, 91 hybrids, 746 infraspecific taxa (458 heterotypic, 288 with autonyms), 17,932 synonyms of accepted taxa, 16 definitely excluded names, 389 names of uncertain application, 672 unresolved names and 454 names belonging to (probably artificial) named hybrids, totalling 22,275 names. The process of compiling this database is described and further editorial rules for the compilation of the taxonomic backbone for the Caryophyllales Network are proposed. A checklist depicting the current state of the taxonomic backbone is provided as . All results are also available online on the website of the Caryophyllales Network and will be constantly updated and expanded in the future. Citation: Korotkova N., Aquino D., Arias S., Eggli U., Franck A., Gómez-Hinostrosa C., Guerrero P. C., Hernández H. M., Kohlbecker A., Köhler M., Luther K., Majure L. C., Müller A., Metzing D., Nyffeler R., Sánchez D., Schlumpberger B. & Berendsohn W. G. 2021: Cactaceae at Caryophyllales.org – a dynamic online species-level taxonomic backbone for the family. – Willdenowia 51: 251–270. Version of record first published online on 31 August 2021 ahead of inclusion in August 2021 issue. Data published through: http://caryophyllales.org/cactaceae/Checklist
... 110, 154), in which diploids were only detected in P. tuberosus f. tuberosus. Although the two P. tuberosus forms show different chromosome numbers, they cannot be separated as different species because they are morphologically very similar, and there are many species of cacti with two or more cytotypes (e.g., Pinkava, 2002;Schlumpberger & Renner, 2012;Las Peñas & al., 2014). Pterocactus megliolii had a DNA content close to P. tuberosus f. tuberosus; thus, we can infer that, although we have not counted its chromosomes, it would be diploid. ...
... The ancestral diploid state was exclusively present in P. tuberosus f. tuberosus. Although there are some gaps in our knowledge, diversification in the genus was accompanied by numerous chromosomal variations, as also observed in the Opuntioideae (Negrón-Ortiz, 2007;Las Peñas & al., 2017 and other Cactaceae (e.g., Cota & Philbrick, 1994;Schlumpberger & Renner, 2012). On the other hand, the genome size mapping, which also had missing data for several species, showed no clear trend. ...
Article
Pterocactus has 10 species endemic to restricted areas of the Argentine Monte and Patagonia regions, also growing in neighbouring Chile. We analysed phylogenetic relationships between its species, based on plastid DNA sequences (psbA‐trnH, rpl16, trnH‐matK), exomorphological features (spine type and size, stem segment form, flower position, epidermis colour, areole pubescence, stem surface) and cytogenetical traits (genome size, somatic chromosome number). Traits were mapped using character mapping and ancestral state reconstruction to understand the dynamics of evolutionary changes. Pterocactus was monophyletic. Two lineages were recovered: clade A (with P. fischeri diverging early and P. reticulatus, P. valentinii, P. megliolii, P. gonjianii, and both P. tuberosus forms) and clade B (with P. neuquensis diverging first and P. araucanus and P. hickenii). Both samples examined of P. australis were paraphyletic. It was estimated that Pterocactus split from its sister group at 9.38 mya. The crown group age was estimated to be 7.69 myr. Species showed x = 11 with different ploidy levels (2n = 22, 44, 99, ca. 110, and 154); only P. tuberosus f. tuberosus was diploid. The numbers were typical of each taxon studied, and polyploidy was important in its diversification. Two 18S‐5.8S‐26S sites were detected regardless of the ploidy level, except P. fischeri with four sites and 2n = ca. 110. The number of 5S sites was 2 in P. gonjianii and the diploid P. tuberosus f. tuberosus, 4 in the tetraploid P. tuberosus f. lelongii, 10 in the probably decaploid P. fischeri and 8 in P. araucanus of unknown chromosome number. The genome size ranged from 2C = 4.94 pg in P. megliolii of unknown chromosome number to 2C = 16.68 pg in the tetradecaploid P. hickenii. The ancestor is reconstructed as a dwarf shrub with cylindrical central spines of 16.35–31.2 mm long, non‐woolly areoles, cylindrical to globose/obpyriform stem segments, non‐tuberculate stem surface, lateral flower position, green/brown to violet epidermis, probably diploid, and with a genome size between 1.10 and 1.23 pg.
... DNA analyses (Wallace 1997, Schlumpberger & Renner 2012, Albesiano & Terrazas 2012, Franck 2013 suggest that Mediocactus hahnianus belongs to the large and polyphyletic Echinopsis genus sensu lato, but the authors suggest using either Echinopsis, Trichocereus or Soehrensia as the genus name. We chose Trichocereus in the broad sense, i.e., based on morphological characters and including large columnar plants as well as small and low growing plants. ...
... The two well-identified clones are cultivated in the Systematics, Evolution and Cytogenetics Laboratory of Cactaceae (IMBIV-CONICET-UNC). Molecular phylogeny studies will be done to ascertain the position of this new clones within the phylogenies published by Schlumpberger & Renner (2012) and Franck et al. (2013), using the same markers and others frequently used in these studies. Furthermore, cytogenetical and morphological characters will be mapped. ...
... (Tribu Trichocereeae -Subtribu Trichocereinae; Barboza et al., 2016). Este género agrupa a una variedad de cactus columnares, con pilosidad en las areolas florales y en general presenta flores blancas y nocturnas; incluye aproximadamente 45 especies, pero su delimitación ha estado históricamente en discusión (Kiesling, 1978;Anderson, 2001;Hunt et al., 2006;Kiesling et al., 2008;Schlumpberger & Renner, 2012). Análisis filogenéticos con datos morfológicos y de ADN posicionaron a Trichocereus como monofilético, si también se incluyen dos especies del género Harrisia (Albesiano & Terrazas, 2012). ...
... Kiesling (1978) considera a estas entidades como especies diferentes, criterio adoptado en el presente trabajo, y ha propuesto que T. pseudocandicans podría ser una especie de reciente formación, derivada de T. candicans, presumiblemente un híbrido entre T. candicans y T. vatteri. Hunt et al. (2006) proponen que ambos taxones son sinónimos; en tanto que, Schlumpberger & Renner (2012) y Albesiano & Terrazas (2012) en sus filogenias incluyen solamente ejemplares de T. candicans. Recientemente, Albesiano (2015) en su tesis doctoral concluye que las diferencias exomorfológicas entre ellas son suficientes para considerar a T. pseudocandicans como una subespecie de T. candicans. ...
Article
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Introducción y objetivos: Las Sierras de Famatina son un área importante de endemismo, allí conviven dos representantes de Trichocereus, de los que se discute su identidad taxonómica. Trichocereus pseudocandicans es endémica, con flores coloridas y diurnas; mientras que, T. candicans presenta distribución más amplia, con flores blancas y nocturnas. Teniendo en cuenta estas diferencias, llevamos a cabo un estudio morfo-anatómico del perigonio, gineceo y fruto, realizando comparaciones cualitativas y cuantitativas. El objetivo es generar información novedosa para el género, aportar caracteres taxonómicos confiables y realizar inferencias sobre su biología reproductiva. M&M: Se realizaron preparados histológicos temporarios y permanentes de flores y frutos, utilizando técnicas clásicas. Se analizaron y compararon estadísticamente 43 variables morfo-anatómicas. Resultados: Las entidades presentan características morfo-anatómicas similares, pero se diferencian por el color del perigonio y el tamaño de las flores. Se destacan los siguientes caracteres: el estilo presenta canal estilar y tejido transmisor, el receptáculo y la pared del ovario son notoriamente diferentes pero íntimamente fusionados, incluso luego de la fecundación, óvulos circinótropos, entre otros. Solo el 27,9% de las variables analizadas resultaron con diferencias estadísticamente significativas, siendo las flores de T. candicans levemente mayores que las de T. pseudocandicans. Conclusiones: La similitud encontrada entre los taxones reafirma su alto grado de parentesco, siendo posible que T. pseudocandicans sea una subespecie o híbrido de T. candicans. Por otro lado, los rasgos florales diferentes podrían deberse a una alta variabilidad fenotípica, influenciada por las características particulares que presentan las Sierras de Famatina.
... Among angiosperms, the species of the Cactaceae family have one of the most impressive evolutionary labile reproductive systems (Mandujano et al., 2010;Schlumpberger & Renner, 2012;Hernández-Hernández et al., 2014;Guerrero et al., 2019a). Cacti have evolved conspicuous flowers that attract a wide range of animal pollinators, including vertebrates (e.g., bats, hummingbirds, passerine birds, and lizards) and insects, such as moths, flies, and bees (Mandujano et al., 2010;Guerrero et al., 2012). ...
... This result and the phylogenetic topology of the clade (Guerrero et al., 2019b), suggest the reversal of a plesiomorphic state in the Neoporteria from bird pollination back to bee pollination. Previous studies in the Cactaceae have revealed that changes in pollination systems can increase diversification rates by promoting speciation (Schlumpberger & Renner, 2012;Hernández-Hernández et al., 2014). Finally, our study highlights the necessity of further studies that could test the existence and potential effects of inbreeding depression, particularly in species with some degree of autogamous reproduction. ...
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Background Sympatric congeneric plants might share pollinators, or each species might avoid competition by evolving specialized traits that generate partitions in pollinator assemblages. In both cases, pollen limitation (a decrease in the quality and quantity of compatible reproductive pollen) can occur, driving the plant mating system to autogamy as a mechanism of reproductive assurance. We assessed the relationships between pollinator assemblages and mating systems in a group of sympatric congeneric plants. We attempted to answer the following questions: (i) How similar are pollinator assemblages among sympatric cactus species? (ii) Which mating systems do sympatric cactus species use? Methods We studied sympatric Eriosyce taxa that inhabit a threatened coastal strip in a mediterranean-type ecosystem in central Chile. We performed field observations on four taxa and characterized pollinators during the years 2016 and 2017. We estimated differences in the pollinator assemblages using the Bray–Curtis index. To elucidate the mating systems, we conducted hand-pollination experiments using three treatments: manual cross-pollination, automatic self-pollination, and control (unmanipulated individuals). We tested differences in seed production for statistical significance using Kruskal–Wallis analysis. Results Eriosyce subgibbosa showed a distinctive pollinator assemblage among the sympatric species that we studied (similarity ranged from 0% to 8%); it was visited by small bees and was the only species that was visited by the giant hummingbird Patagona gigas . Pollinator assemblages were similar between E. chilensis (year 2016 = 4 species; 2017 = 8) and E. chilensis var. albidiflora (2016 = 7; 2017 = 4); however, those of E. curvispina var. mutabilis (2016 = 7; 2017 = 6) were less similar to those of the aforementioned species. E. curvispina var. mutabilis showed the highest interannual variation in its pollinator assemblage (18% similarity). Reproduction in E. subgibbosa largely depends on pollinators, although it showed some degree of autogamy. Autonomous pollination was unfeasible in E. chilensis , which depended on flower visitors for its reproductive success. Both E. chilensis var. albidiflora and E. curvispina var. mutabilis showed some degree of autogamy. Discussion We observed differences in pollinator assemblages between E. subgibbosa and the remaining Eriosyce taxa, which depend on hymenopterans for pollen transfer. Pollinator assemblages showed considerable interannual variation, especially those of E. subgibbosa (ornithophilous syndrome) and E. curvispina var. mutabilis (melitophilous syndrome). Autogamous reproduction in these taxa may act as a reproductive assurance mechanism when pollinator availability is unpredictable. Our study contributes to improving our understanding of the reproductive systems of ecological interactions between threatened species in a Chilean mediterranean-type ecosystem.
... b Owhi, 1992 two North American, one Japanese, and some other Chinese huge trumpet-shaped lily species and coming into contact with the floral sexual organs (Skinner, 1988;Yokota and Yahara, 2012;references in Table 3), implying that these lilies may have floral mechanisms similar to that revealed here. Echinopsis provides another example (Schlumpberger and Renner, 2012), with recurrent transitions from short trumpet-shaped ancestors putatively pollinated by bees to huge trumpet-shaped flowers putatively pollinated by hawkmoths. As observed for Lilium, huge trumpet-shaped Echinopsis can be visited and probably pollinated by hawkmoths with a wide range of tongue lengths (references in Table 3). ...
... For a flower with a short trumpet-shaped morphology, once a pollinator shift from diurnal insects to hawkmoths has occurred, the subsequent evolutionary elongation of perianths will probably follow with ease if trade-offs in using hawkmoth pollinators are insignificant, as shown herein. This scenario may explain why trumpet-shaped flowers have evolved to be huge in multiple clades of Lilium and Echinopsis and possibly other taxa (Schlumpberger and Renner, 2012; Table 3). ...
... 2017). Previous studies have documented several aspects of the species' biology, including their phylogenetic relationships (Schlumpberger and Renner 2012), reproduction (Ortega-Baes et al. 2011), population structure (Quipildor et al. 2017), chemical composition (Padró and Soto 2013;Padró et al. 2022) and threats to their persistence (e.g., Peco et al. 2011;Ortega-Baes and Lowry 2017;Gorostiague et al. 2018). Further, the evolutionary model of cactophilic Drosophila of South America has revealed the key role of the cactus chemistry in niche partitioning , phenotypic and genetic adaptations (Padró et al. 2014;De Panis et al. 2016), host-plant specificity (Soto et al. 2017;Bouzas et al. 2021), sexual isolation (Padró et al. 2019), microbial composition (Mongiardino Koch et al. 2015) and ultimately in the phylogenetic radiation of the group (Oliveira et al. 2012;Colines et al. 2018). ...
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Desert ecosystems are currently threatened by human activities resulting in the rapid decline of xerophytic plants and specialized fauna. In South America, the demise of cactus species already resulted in the population decline of > 30% of the iconic giant columnar cactus Trichocereus terscheckii. The increasing vulnerability of these keystone species could trigger a cascade of secondary extinctions in highly dependent organisms. Thus, necrotic cacti constitute an important habitat for desert arthropods, yet little is known about the hidden diversity of this neglected niche. We used DNA barcode techniques to survey the diversity of arthropods in a threatened cactus forest dominated by T. terscheckii in northwestern Argentina. We obtained a total of 542 mitochondrial barcode sequences, resulting in 323 Molecular Taxonomic Units (MOTUs) associated to the xerophytic forest and 21 MOTUs exclusive to the giant cactus necrosis. Our results indicated that the area is a biodiversity hotspot within the harsh Andean desert and suggests that nearly 30 species could occur in the decaying cactus, representing the highest richness of cactophilic arthropods recorded in any cactus on the continent to date (6 orders and 16 families). The community structure of cactophilic arthropods showed a phylogenetic clustering pattern, suggesting the coexistence of closely related species. Overall, our study indicates that the giant cactus necrosis sustains a particular phylogenetic diversity of desert arthropods, while demonstrating the efficacy of DNA barcodes for biodiversity assessments in complex and poorly understood ecological systems.
... Looking at the presence of the different pollinators across the main taxonomic groups, we could presume that the more ancestral pollinators are bees, since they were found in almost every tribe. This type of pollination could have originated others through pollinator shifts, as it has been previously suggested for Cactaceae (Pimienta-Barrios & del Castillo, 2002;Schlumpberger & Renner, 2012) and other plant families (Alcantara & Lohmann, 2010). ...
Article
Biotic interactions are said to be more specialised in the tropics, and this was also proposed for the pollination systems of columnar cacti from North America. However, it has not yet been tested for a wider set of cactus species. Here, we use the available information about pollination in the Cactaceae to explore the geographic patterns of this mutualistic interaction, and test if there is a latitudinal gradient in its degree of specialisation. We performed a bibliographic search of all publications on the pollination of cacti species and summarized the information to build a database. We used generalised linear models to evaluate if the degree of specialisation in cacti pollination systems is affected by latitude, using two different measures: the number of pollinator guilds (functional specialisation) and the number of pollinator species (ecological specialisation). Our database contained information about the pollination of 148 species. The most frequent pollinator guilds were bees, birds, moths and bats. There was no apparent effect of latitude on the number of guilds that pollinate a cactus species. However, latitude had a small but significant effect on the number of pollinator species that services a given cactus species. Bees are found as pollinators of most cactus species, along a wide latitudinal gradient. Bat and bird pollination is more common in the tropics than in the extra‐tropics. The available information suggests that cacti pollination systems are slightly more ecologically specialised in the tropics, but it does not support any trend with regard to functional specialisation.
... For instance, Espostoopsis dybowskii was recovered within the Cereinae clade, although this taxon is traditionally considered a member of the subtribe Trichocereinae. In the phylogenies of Ritz et al. [3], Schlumpberger and Renner [71], Lendel et al. [72], and Bombonato et al. [17], it is also possible to observe the members of these subtribes as nonmonophyletic taxa. Finally, our results found the same contentious relationships involving the genera Cereus, Cipocereus, and Praecereus inferred with plastid [50] and RAD-Seq data [17]. ...
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The molecular phylogenies of Cactaceae have enabled us to better understand their systematics, biogeography, and diversification ages. However, most of the phylogenetic relationships within Cactaceae major groups remain unclear, largely due to the lack of an appropriate set of molecular markers to resolve its contentious relationships. Here, we explored the genome and transcriptome assemblies available for Cactaceae and identified putative orthologous regions shared among lineages of the subfamily Cactoideae. Then we developed a probe set, named Cactaceae591, targeting both coding and noncoding nuclear regions for representatives from the subfamilies Pereskioideae, Opuntioideae, and Cactoideae. We also sampled inter- and intraspecific variation to evaluate the potential of this panel to be used in phylogeographic studies. We retrieved on average of 547 orthologous regions per sample. Targeting noncoding nuclear regions showed to be crucial to resolving inter- and intraspecific relationships. Cactaceae591 covers 13 orthologous genes shared with the Angiosperms353 kit and two plastid regions largely used in Cactaceae studies, enabling the phylogenies generated by our panel to be integrated with angiosperm and Cactaceae phylogenies, using these sequences. We highlighted the importance of using coalescent-based species tree approaches on the Cactaceae591 dataset to infer accurate phylogenetic trees in the presence of extensive incomplete lineage sorting in this family.
... multigeniculata and C. whipplei: Baker 2016), Echinocereus (E. acanthosetus and E. pulchellus: Sánchez et al. 2020), andEscobaria (E. guadalupensis andE. ...
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Pilosocereus is one of the Cactaceae family’s most relevant genera in terms of the number of species and its wide geographical range in the Americas. Within Pilosocereus , five informal taxonomic groups have been recognized, one of which is P. leucocephalus group s.s. , whose phylogenetic relationships remain unresolved. Therefore, our objectives are to recognize the circumscriptions of the species in P. leucocephalus group s.s. and to corroborate the monophyly and phylogenetic relationships of this group through a set of morphological and molecular characters. This study is based on representative sampling along the broad distribution of this group in Mexico and Central America using multivariate and phylogenetic analyses. The morphological characters identified to contribute to species recognition and group formation are branch diameter, areole length, the areole length-width ratio, the distance between areoles, the length of the longest radial spine, and branch and spines colors. The chloroplast markers rpl16 , trnL-trnF , and petL-psbE and the nuclear marker AT1G18270 support the monophyly of the P. leucocephalus group s.s. , and two probable synapomorphies are suggested, including one transversion in rpl16 and another in petL-psbE . Together, our results demonstrate that sampled species of P. leucocephalus group s.s. encompass six species distributed in Mexico and Central America: P. alensis and P. purpusii in the western region, P. chrysacanthus and P. collinsii in the central region, and P. gaumeri and P. leucocephalus in the eastern region. A taxonomic key to recognized species is provided.
... H. Friedrich & G.D. Rowley and Echinopsis schickendantzii F.A.C. Weber) Schlumpberger. However, a phylogenetic study suggested that Echinopsis comprises several divergent lineages and that T. spachianus falls within the genus Trichocereus (Schlumpberger & Renner 2012). Trichocereus spachianus is a multistemmed shrub with upright stems between 1 and 2.5 m high. ...
Article
Cactaceae are among the most problematic invasive alien plants in South Africa, posing serious negative consequences to agriculture and natural ecosystems. Fortunately, South Africa has a long and successful history of controlling cactus weeds using biological control (biocontrol). This paper reviews all the biocontrol programmes against invasive alien Cactaceae in South Africa, focusing on the decade since the last review published in 2011, up to, and including 2020. Biocontrol programmes against 16 target weeds are summarised, all of which rely on either the galling mealybug, Hypogeococcus sp. (Pseudococcidae), or various species or intraspecific lineages of cochineal insects (Dactylopius spp., Dactylopiidae) as agents. New agents are being considered for the three target weed species, Opuntia elata Salm-Dyck, Opuntia megapotamica Arechav. and Trichocereus spachianus (Lem.) Riccob., while permission to release a new agent against Cylindropuntia pallida (Rose) F.M. Knuth has recently been granted. The biocontrol agent, Dactylopius opuntiae (Cockrell) stricta, which has been utilised for the successful control of Opuntia stricta Haw., has shown some promise as an agent against one of the worst cactus weeds in the country, the North Cape/Free State variety of Opuntia engelmannii Salm-Dyck. Post-release monitoring and recent observations of the status of control for the 11 other cactus weeds, all of which have well-established agents, are provided. Taxonomic uncertainties and misidentifications of both target weeds and agents has been a constraint to biocontrol efforts, but this has been partially overcome through the use of genetic techniques. Biocontrol is particularly successful in controlling cactus weeds compared to most other taxonomic groups, and it is likely that past successes can be repeated with new target weeds. Mass-rearing and redistribution of agents are essential to gain the maximum possible benefit from cactus biocontrol agents, and recent increases in mass-rearing outputs have been beneficial.
... The ability to successfully exploit columnar cacti would comprise T. terscheckii and the more closely related species such as T. atacamensis. But regardless of the adaptation that allows the exploitation of these plants, it does not even cover all the species of the genus Trichocereus as observed for the phylogenetically more distant T. tarijensis and T. schikendantzi (Schlumpberger & Renner, 2012).The quantitative and qualitative differences in chemical defenses among these species (Trout, 2014) might be preventing the implementation of a more general detoxification mechanism (i.e. the existence of a Jack of all trades for columnar cacti). The pattern observed in the CVs points to the same conclusion. ...
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1. We assessed the host‐related niche breadth for D. koepferae and D. buzzatii, a pair of sibling cactophilic species with contrasting backgrounds of host use. We tested for the ‘Jack of all trades‐ Master of none’ scenario predicting a more evident exhibition in D. buzzatii rather than in the supposedly specialist D. koepferae. 2. Additionally, using laboratory strains of both species selected for tolerance to extremely high concentrations of a columnar cacti's secondary metabolites, we tested whether adaptation to a high‐demanding host involved the loss of performance capabilities in other hosts. 3. D. buzzatii was more affected by the artificial host shifts than D. koepferae which presented an overall better performance when rearing in novel columnar hosts. 4. Artificially selected strains of D. buzzatii performed poorer in both novel and native natural hosts compared with control strain indicating that adaptation carried associated costs regarding the potential to exploit other cacti. Conversely, artificial evolution of the D. koepferae's strains did not translate into decreased performance in other hosts. 5. D. buzzatii complied better with the predictions of the Jack of all trades‐Master of none hypothesis. 6. Host specialization is a dynamic feature in the repleta group and a major driver of diversification in its evolutionary history. As the group presents an Opuntia breeder as the ancestral condition, D. buzzatii would represent not only a plesiomorphic state of host use but also the ancestral ecological strategy of specialization.
... The development of new phylogenetic techniques in recent decades has revolutionized the study of plant evolution (Willis and McElwain 2002, Donoghue 2008, Endress 2011). In the field of plant reproductive biology, phylogenetics has begun to provide useful insights regarding breeding system evolution (Weiblen et al. 1999, Truyers et al. 2005, Goldberg and Igic 2012, Kafer 2013) and floral trait evolution (Perez et al. 2006, DeWitt Smith 2010, Marten-Rodriguez et al. 2010, Schlumpberger and Renner 2012. The obvious benefit of this approach is the accurate identification of trait shifts (e.g., changes in corolla tube morphology) that may have arisen multiple times in a taxonomic group, compared to traditional morphological taxonomy that often groups taxa based on shared characters without taking convergence into consideration. ...
... Out of 117 hawkmoth species recorded from Argentina, only the short-tongued Hyles euphorbiarum (proboscis length shorter than 30 mm), is consistently recorded beyond 40 • south latitude (Moré et al., 2014). Interestingly, other plant lineages with significant amounts of sphingophily, such as Nicotiana (Solanaceae) and the family Cactaceae, show the same geographic and altitudinal patterns described here (Goodspeed, 1954;Schlumpberger and Renner, 2012). It is worth mentioning that in addition to metabolic limits, other factors may be involved in the distribution of hawkmoth fauna. ...
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Flower phenotype may diverge within plant lineages when moving across pollination climates (geographic differences in pollinator abundance or preference). Here we explored the potential importance of pollinators as drivers of floral color diversification in the nightshade genus Jaborosa taking into account color perception capabilities of the actual pollinators (nocturnal hawkmoths vs saprophilous flies) under a geographic perspective. We analyzed the association between transitions across environments and perceptual color axes using comparative methods. Our results revealed two major evolutionary themes in Jaborosa: 1) a ‘warm subtropical sphingophilous clade’ composed of three hawkmoth-pollinated species found in humid lowland habitats, with large white flowers that clustered together in the visual space of a model hawkmoth (Manduca sexta) and a ‘cool-temperate brood-deceptive clade’ composed of largely fly-pollinated species with small dark flowers found at high altitudes (Andes) or latitudes (Patagonian Steppe), that clustered together in the visual space of a model blowfly (Lucilia sp.). Our findings suggest that the ability of plants to colonize newly formed environments during Andean orogeny and the ecological changes that followed were concomitant with transitions in flower color as perceived by different pollinator functional groups. Our findings suggest that habitat and pollination mode are inextricably linked in the history of this South American plant lineage.
... The factors that influence the net species diversification ratethat is to say, the net result of speciation and extinction for each taxon-are multiple and complex (Scott and Arnold, 1995;Magallón and Sanderson, 2001;Magallón and Castillo, 2009;Arakaki et al., 2011;Schlumpberger, 2012;Schlumpberger and Renner, 2012;Van der Niet and Johnson, 2012;Hernández-Hernández et al., 2014). These include extrinsic factors, such as physical space, climate, other organisms, or available habitats, or intrinsic factors, such as morphological or physiological traits, characters that affect the body's adequacy through its growth, survival, and/or reproduction (Glor, 2010;Losos, 2010;Bouchenak-Khelladi et al, 2015). ...
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Agave sensu lato is one of the most diverse and complex genera of Asparagaceae, with more than 250 species. The morphological, ecological, and evolutionary diversity of the group has complicated its taxonomical study. We conducted phylogenetic analyses of DNA sequence data to reconstruct the phylogenetic relationships of the Agave genus. We included 107 species of the Asparagaceae family from which 83 correspond to the Agave sensu lato clade (Agave sensu stricto + Polianthes + Manfreda and Prochnyanthes, which together represent 30% of the genus) and as outgroups the genera Dasylirion, Hesperoyucca, Chlorogalum, Camassia, Hesperaloe, Yucca, Beschorneria, and Furcraea, in order to estimate the age and propose the history of their diversification. Previous studies postulated the relevance of the Miocene in the speciation rates of the agaves, as well as the relevance of the type of inflorescence in its diversification. However, these assertions have not been well supported. The analysis of chloroplast regions resulted in low resolution, which could be the consequence of the few variable sites. On the other hand, the internal transcribed spacer (ITS) implemented in our analysis ensued in higher resolution and better support values. Our phylogenetic analyses recovered five groups; one is the Striatae group, which is the sister group to Agave sensu stricto clade. Within this clade, we found three main groups with high support; these groups are not related with previous morphological proposals. We also analyzed the dates of origin and diversification rates. A Bayesian analysis of macroevolutionary mixtures indicated two significant shifts; the first was identified at 6.18 Ma, where the speciation rate increased to 4.10 species/Mya, this shift occurred during the late Miocene period, characterized by the emergence of arid biomes in North America. The second was identified at a stem age of 2.68 Ma Frontiers in Plant Science | www.frontiersin.org 1 November 2020 | Volume 11 | Article 536135 Jiménez-Barron et al. Phylogeny and Diversification in Agave where the speciation rate increased to 6.04 species/Mya. Concerning the ancestral reconstruction state of the inflorescence type in the Agave sensu stricto clade, the spike inflorescence character was predominant in the early-diverging groups, whereas the late-diverging groups present panicle inflorescences as the predominant character and higher speciation rates.
... At the time of publication of Borzicactus hoxeyi it was decided to use the genus Borzicactus with a circumscription including Loxanthocereus based on similarities of the hummingbird adapted flower morphology. A molecular study (Schlumpberger & Renner, 2012) (Lodé, 2014). Further molecular evidence (unpublished) has confirmed this close relationship between Loxanthocereus and Haageocereus with Borzicactus a more distant relative. ...
... The first study to evaluate pollination ecology in Cleistocactus was developed in the Dry Chaco ecoregion and showed that C. baumannii was pollinated exclusively by hummingbirds, while C. smaradigoflorus was possibly pollinated by hummingbirds and bees (Gorostiague & Ortega-Baes 2016). It is interesting that the two species are closely related and that both are ornithophilous (Schlumpberger & Renner 2012). According to Gorostiague & Ortega-Baes (2016), many ornithophilous cacti may have generalized pollination systems (e.g. ...
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Cleistocactus baumannii is the only ornithophilous cactus species in the Brazilian Chaco. In addition, this species of Cactaceae invests heavily in flowering in the ecoregion. Such characteristics motivated us to evaluate the temporal availability of flowers in the context of its floral visitors. The reproductive system of C. baumannii, the number of individuals in flowering, flower abundance and the frequency and richness of floral visitors were evaluated and quantified. Nectar robbery was a frequent phenomenon in the studied population; therefore, we compared the pollen load deposited on the stigma of damaged (robbery) and undamaged flowers. In the Brazilian Chaco, C. baumannii is self-incompatible and has a continuous flowering pattern, providing floral resources throughout the year for nine species of floral visitors. One hummingbird species acted as a potential pollinator, and we consider Xylocopa splendidula to be a nectar robber. We found stigma of flowers damaged by nectar robbers to have lower pollen loads than those of undamaged flowers. This study highlights the importance of studying reproductive traits in different populations to understand changes in the reproductive success of plant species at different scales and possible causes, such as availability of floral visitors, incidence of robbers and flowering patterns.
... Echinopsis possui 100-150 espécies com grande diversidade na arquitetura e são polinizadas por abelhas, beija-flores ou esfingídeos [1] . Polinização por todos estes grupos (mista) pode ocorrer em espécies com flores noturnas, cuja longevidade floral se estende até parte da manhã [2,3] . ...
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Introdução Echinopsis possui 100-150 espécies com grande diversidade na arquitetura e são polinizadas por abelhas, beija-flores ou esfingídeos [1]. Polinização por todos estes grupos (mista) pode ocorrer em espécies com flores noturnas, cuja longevidade floral se estende até parte da manhã [2,3]. Autoincompatibilidade predomina no gênero [4]. Apresentamos o estudo da polinização de Echinopsis rhodotricha no Chaco brasileiro visando contribuir com futuros programas de conservação. Metodologia O estudo foi realizado em remanescentes de vegetação chaquenha brasileira (Savana Estépica Arbórea), em Porto Murtinho, MS. A morfologia floral foi estudada a partir de flores frescas. A viabilidade polínica foi testada com carmin acético a receptividade estigmática pela presença de exsudados. Para a biologia floral foram marcados 20 botões em pré-antese, acompanhados da abertura à murcha. O sistema de reprodução foi estudado a partir de experimentos de autopolinização espontânea e manual, geitonogamia, xenogamia e controle. A observação dos visitantes florais foi realizada durante todo o período de longevidade floral (22 h noite+18 h dia= 40 horas de observação). Fluxo polínico foi verificado aplicando pó fluorescente sobre as anteras de seis flores, conferidas no fim da antese para registrar se houve e local onde este foi depositado sobre as flores e sua procedência. Resultados Echinopsis rhodotricha possui flores solitárias, actinomorfas, brancas, hipocrateriformes, com tubo longo (131±20mm) (Fig. 1a), onde o néctar é produzido (80±20µl, [%]= 10,1±4,5) e armazenado. As anteras são rimosas, produzem pólen com 87±9% de viabilidade e se dispõem ao redor e ligeiramente abaixo dos lobos estigmáticos. As flores duram ~15 h; abrem ~20h00 e fecham ~11h00 período em que emitem leve odor adocicado. As características acima enquadram as flores de E. rhodotricha na síndrome da esfingofilia [5]. Frutificação foi registrada somente após autopolinização manual (27,8%), geitonogamia (63,6%), xenogamia (60%) e controle (30%). Deposição de pó fluorescente ocorreu em 83% das flores, sempre na mesma flor, sobre os estigmas e/ou pétalas (Fig. 1b). Echinopsis rhodotricha é o primeiro registro de autocompatibilidade no gênero, o que possibilita autopolinizações, porém a espécie é polinizador dependente, por ser auto-estéril. Doze espécies visitaram as flores: abelhas (diurnas), Coleoptera (4 spp. cada), Araneae (diurna) (1 sp.), Orthoptera (2 spp.) (Fig. 1d), Hemipitera (noturnos, 1 sp.), para coletar pólen, néctar (abelhas), copular (Coleoptera), comer partes florais (Coleoptera e Orthoptera) e talvez predar (Araneae). As abelhas foram as mais frequentes (1,04), seguidas de Coleoptera (0,74) e demais visitantes (0,16). Ao coletar pólen e/ou néctar as abelhas Apis mellifera, Melipona cf. rufiventris e Augochloropsis sp contatam os anteras e estigmas (Fig. 1c), podendo atuar como polinizadores secundários como em outras espécies de Echinopsis [2,3]. Os florívoros comem principalmente estames (Fig. 1d), danificam as flores e podem comprometer o sucesso reprodutivo da espécie ao reduzir a quantidade de pólen, danificar os estigmas e diminuir a atratividade floral [6]. Com efeito, a frutificação em condições naturais foi relativamente baixa indicando serviço deficiente de polinização e/ou florivoria. Conclusões Echinopsis rhodotricha parece ser primeiro caso de autocompatibilidade no gênero. A antese estendida além do período noturno permitiu a ocorrência de visitantes diurnos, que podem atuar como polinizadores secundários (abelhas), sugerindo polinização generalista (ou mista) para esta espécie esfingófila que não recebeu visitas de mariposas.
... In fact, all previous reports of Trichocereus species in Catalonia have been made as Echinopsis. Both genera are separated based on the following evidences: i) Echinopsis is polyphyletic (Schlumpberger & Renner, 2012), ii) Trichocereus is monophyletic (Albesiano & Terrazas, 2012) and iii) the Trichocereus clade is supported by three synapomorphies: basitonic growth with prostrate branches, imbricate scales along the floral tube, and subglobose fruits (Albesiano & Terrazas, 2012). Without considering the species that were included wtihin Trichocereus, two species of Echinopsis were reported from Catalonia: E. eyriesii (Turpin) Pfeiff. ...
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New data on the identity and distribution of alien plants of the genera Cereus and Trichocereus in Catalonia are presented. Cereus jamacaru, C. repandus and C. peruvianus should be excluded from the flora of Catalonia. At present, the only species of Cereus regarded as casual in the studied area is C. hildmannianus. Four species of Trichocereus (T. macrogonus, T. schickendantzii subsp. schickendantzii, T. spachianus and T. taquimbalensis) are currently known the studied area.
... Among angiosperms, the species of the Cactaceae family present one of the most 30 impressive evolutionarily labile reproductive systems (Schlumpberger & Renner, 2012;31 Hernández-Hernández et al., 2014;Guerrero et al., 2019b). Cacti have evolved conspicuous 32 flowers that attract a wide range of animal pollinators, including vertebrates (e.g., bats, 33 hummingbirds, passerine birds, lizards) and insects, such as moths and bees (Guerrero et al., 34 2012). ...
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Background. Sympatric plant species that share pollinators may have similar mating systems because their floral traits are subject to comparable canalization imposed by pollinators. However, if each sympatric species bears specialized floral morphology, each species may attract different pollinators. Our study aims to describe the pollinator diversity and pollination systems of four taxa of Eriosyce that co-occur in an endangered coastal Mediterranean ecosystem in Central Chile. We took two approaches in our study: we assessed the composition and similarity of flower visitors among taxa, and we characterized the breeding systems to determine dependence on pollinators and self-compatibility. Methods . We performed field observations to characterized pollinators during two consecutive years (2016-2017). Additionally, we performed pollination experiments to elucidate reproductive modes using three treatments: manual cross-pollination, automatic self-pollination, and control (unmanipulated individuals). Results . We observed one bird species (Giant hummingbird Patagona gigas only visiting E. subgibbosa ) and 14 bee species (13 natives plus Apis mellifera ) visiting cacti of the genus Eriosyce . We observed variation in the similarity of intra-specific pollinator composition between years and among Eriosyce species within the same year. Individuals of E. subgibbosa were visited by less number of species (2016 = 4; 2017 = 2), while E. chilensis (2016 = 4; 2017 = 8) , E. chilensis var. albidiflora (2016 = 7; 2017 = 4) and E. curvispina var . mutabilis (2016 = 7; 2017 = 6) were visited by a richest guild of visitors (up to 10 bee species each).Autonomous pollination was unfeasible in E. chilensis , which depend on bees to achieve their reproductive success. Eriosyce subgibbosa , visited mainly by the Giant hummingbird, depends on pollinators to achieve reproductive success. Both E. chilensis var. albidiflora and E. curvispina var. mutabilis were visited by a diverse assemblage of non-social native bees, showing some degree of autonomous pollination and self-compatibility. Discussion . Pollinator diversity analyses showed considerable pollinator differences between the species with ornithophilous flowers ( E. subgibbosa ) and remain taxa which solely dependent on Apoidea species for pollen transfer. The high diversity of native bees among sympatric Eriosyce may be a caused by their microclimatic differences at spatial (differences among cacti microhabitats) and temporal levels (differences of climatic conditions between August to December when different Eriosyce species bloom). Our study contributes to unveiling the evolutionary mechanisms for pollinator partitioning of sympatric close-related plant species. Furthermore, it improves understanding of threatened species reproductive system and ecological interactions, especially to E. chilensis and E. chilensis var. albidiflora , whose studied populations are the only known for these taxa.
... Among angiosperms, the species of the Cactaceae family present one of the most 30 impressive evolutionarily labile reproductive systems (Schlumpberger & Renner, 2012;31 Hernández-Hernández et al., 2014;Guerrero et al., 2019b). Cacti have evolved conspicuous 32 flowers that attract a wide range of animal pollinators, including vertebrates (e.g., bats, 33 hummingbirds, passerine birds, lizards) and insects, such as moths and bees (Guerrero et al., 34 2012). ...
Preprint
Full-text available
Background. Sympatric plant species that share pollinators may have similar mating systems because their floral traits are subject to comparable canalization imposed by pollinators. However, if each sympatric species bears specialized floral morphology, each species may attract different pollinators. Our study aims to describe the pollinator diversity and pollination systems of four taxa of Eriosyce that co-occur in an endangered coastal Mediterranean ecosystem in Central Chile. We took two approaches in our study: we assessed the composition and similarity of flower visitors among taxa, and we characterized the breeding systems to determine dependence on pollinators and self-compatibility. Methods . We performed field observations to characterized pollinators during two consecutive years (2016-2017). Additionally, we performed pollination experiments to elucidate reproductive modes using three treatments: manual cross-pollination, automatic self-pollination, and control (unmanipulated individuals). Results . We observed one bird species (Giant hummingbird Patagona gigas only visiting E. subgibbosa ) and 14 bee species (13 natives plus Apis mellifera ) visiting cacti of the genus Eriosyce . We observed variation in the similarity of intra-specific pollinator composition between years and among Eriosyce species within the same year. Individuals of E. subgibbosa were visited by less number of species (2016 = 4; 2017 = 2), while E. chilensis (2016 = 4; 2017 = 8) , E. chilensis var. albidiflora (2016 = 7; 2017 = 4) and E. curvispina var . mutabilis (2016 = 7; 2017 = 6) were visited by a richest guild of visitors (up to 10 bee species each).Autonomous pollination was unfeasible in E. chilensis , which depend on bees to achieve their reproductive success. Eriosyce subgibbosa , visited mainly by the Giant hummingbird, depends on pollinators to achieve reproductive success. Both E. chilensis var. albidiflora and E. curvispina var. mutabilis were visited by a diverse assemblage of non-social native bees, showing some degree of autonomous pollination and self-compatibility. Discussion . Pollinator diversity analyses showed considerable pollinator differences between the species with ornithophilous flowers ( E. subgibbosa ) and remain taxa which solely dependent on Apoidea species for pollen transfer. The high diversity of native bees among sympatric Eriosyce may be a caused by their microclimatic differences at spatial (differences among cacti microhabitats) and temporal levels (differences of climatic conditions between August to December when different Eriosyce species bloom). Our study contributes to unveiling the evolutionary mechanisms for pollinator partitioning of sympatric close-related plant species. Furthermore, it improves understanding of threatened species reproductive system and ecological interactions, especially to E. chilensis and E. chilensis var. albidiflora , whose studied populations are the only known for these taxa.
... S) at strictly inland habitats between 600 and 1300 m.a.s.l. Schlumpberger & Renner (2012) showed that the genus Echinopsis is polyphyletic and split it into various genera, one of which is Leucostele. Schlumpberger (2012: 29) proposed the new combination Leucostele coquimbana (Molina 1782: 170) Schlumpberger (2012. ...
Article
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The taxonomy of Chilean Cactaceae is notoriously difficult (see e.g., Duarte et al. 2014). Molecular data, and phylogenetic approaches, provided answer to better understand the natural classification of cacti (see e.g., Nyffeler 2002, Edwards 2005, Nyffeler & Eggli 2010, Ritz et al. 2012, Schlumpberger & Renner 2012, Larridon et al. 2015, 2018). As part of an ongoing revision of the Chilean Cacti, which is part of the “Modern Flora of Chile” initiative, nomenclatural novelties are needed in Eulychnia Philippi (1860: 23), Leucostele Backeberg (1953: 36) and Maihueniopsis Spegazzini (1925: 86) (Cactaceae Juss.).
... The cytogenetics of Cactaceae has been studied extensively over the previous century (Arakaki et al. 2007;Assis et al. 2003;Baker 2006Baker , 2016Baker & Pinkava 1987;Baker et al. 2009;Baker & Cloud-Hughes 2014;Bhattacharyya 1970, Cid & Palomino 1996Cohen & Tel-Zur 2012;Cota & Phibrick 1994;Das et al. 1997Das et al. , 1998aDas et al. , 1998bDonati 2011Donati , 2017aDonati , 2017bFenstermacher 2016;Gadella et al. 1979;Gutiérrez-Flores et al. 2018, Las Peñas et al. 2014, 2017Majure et al. 2012aMajure et al. , 2012bMajure et al. , 2012cMoreno et al. 2015;Negrón-Ortiz 2007;Palomino 2016;Parfitt 1987Parfitt , 1997Pinkava & McLeod 1971;Pinkava et al. 1973Pinkava et al. , 1977Pinkava et al. , 1985Pinkava et al. , 1992aPinkava et al. , 1992bPinkava et al. , 1998Pinkava & Parfitt 1982;Powell & Weedin 2001, 2005Ross 1981;Schlumpberger & Renner 2012;Ward 1984;Weedin & Powell 1978a, 1978b, 1980Weedin et al. 1989, Wellard 2016. The general degree of polyploidy has been estimated for most genera (Pinkava et al. 1998). ...
Article
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Chromosome ploidy and the evolution and distribution of the Cactaceae, especially in the Genera Cylindropuntia, Echinocereus, Grusonia, and Micropountia.
... Mescaline occurs naturally in some members of the Cactaceae plant family ( Fig. 1) [28][29][30][31][32][33][34][35]. It is also found in small amounts in certain members of the Fabaceae (bean) family, including Acacia berlandieri [36]. ...
Article
Background: Mescaline (3,4,5-trimethoxyphenethylamine), mainly found in the Peyote cactus (Lophophora williamsii), is one of the oldest known hallucinogenic agents that influence human and animal behavior, but its psychoactive mechanisms remain poorly understood. Objectives: This article aims to fully review pharmacokinetics and pharmacodynamics of mescaline, focusing on the in vivo and in vitro metabolic profile of the drug and its implications for the variability of response. Methods: Mescaline pharmacokinetic and pharmacodynamic aspects were searched in books and in PubMed (U.S. National Library of Medicine) without a limiting period. Biological effects of other compounds found in peyote were also reviewed. Results: Although its illicit administration is less common, in comparison with cocaine and Cannabis, it has been extensively described in adolescents and young adults, and licit consumption often occurs in religious and therapeutic rituals practiced by the Native American Church. Its pharmacodynamic mechanisms of action are primarily attributed to the interaction with the serotonergic 5-HT2A-C receptors, and therefore clinical effects are similar to those elicited by other psychoactive substances, such as lysergic acid diethylamide (LSD) and psilocybin, which include euphoria, hallucinations, depersonalization and psychoses. Moreover, as a phenethylamine derivative, signs and symptoms are consistent with a sympathomimetic effect. Mescaline is mainly metabolized into trimethoxyphenylacetic acid by oxidative deamination but several minor metabolites with possible clinical and forensic repercussions have also been reported. Conclusion: Most reports concerning mescaline were described in a complete absence of exposure confirmation, since toxicological analysis is not widely available. Addiction and dependence are practically absent and it is clear that most intoxications appear to be mild and are unlikely to produce life-threatening symptoms, which favors the contemporary interest in the therapeutic potential of the drugs of the class.
... To carry out a character state reconstruction we scored four morphological characters relevant for Eulychnia at species level: (1) habit: arborescent (>3 m) vs. shrubby (<3 m); (2) branching architecture: ascending to erect vs. procumbent; (3) rib height and width: steep (2-3 cm) and narrow ( 1 cm) vs. low (<1 cm) and wide (1.5-3 cm); and (4) indumentum of pericarpel and hypanthium: long dense wool vs. short scant wool vs. spiniferous. The characters 'habit' and 'rib height and width' were selected based on Hern andez-Hern andez et al. (2011), Schlumpberger and Renner (2012), and Gibson and Nobel (1986). Variations in fractional rib heights have an important impact on Photosynthetically Active Radiation (PAR) interception and daily CO 2 uptake in Crassulacean Acid Metabolism (CAM) plants (Gibson & Nobel, 1986). ...
Article
Populations of the columnar cactus genus Eulychnia (Cactaceae) are an iconic sight in the Chilean Atacama Desert. The most recent taxonomic treatment of the genus suggested to accept up to seven taxa at species level based on morphological data. To date, species boundaries and infrageneric relationships in Eulychnia have not been investigated using a molecular approach. In this study, sequence data were generated for six chloroplast markers (rpl32-trnL, trnH-psbA, trnL-trnF, trnQ-rps16, trnS-trnG, and ycf1) for the seven species. Where possible, samples were collected from the south and north of the distribution range of widely distributed species, as well as plants from two morphologically distinct populations in the Atacama and Coquimbo Regions. Evolutionary trends of morphological characters were investigated using ancestral state reconstruction, and the habitat of the Chilean taxa was taken into account based on latitudinal and altitudinal distribution, precipitation regime, and vegetation zones. Two major clades were retrieved in the molecular phylogenetic hypotheses, a northern clade and a southern clade, which can easily be distinguished morphologically by differences in rib shape and type of the indumentum of the pericarpel and the hypanthium. The only Eulychnia taxon found outside Chile is most commonly accepted as Eulychnia iquiquensis subsp. ritteri. However, its isolated geographic distribution and the fact that this taxon is not most closely related to E. iquiquensis but was retrieved as sister to the rest of the northern clade in our molecular phylogenetic results support the recognition of E. ritteri at species level. Our results also provide some support for the two recently published species, E. chorosensis (previously placed in E. acida s.l.) and E. taltalensis (previously considered to be part of E. breviflora s.l.). The relationships in the southern clade need further study.
... In Rio Grande do Sul (BR), the populations of P. graessneri appear to be the northeast continuation of the populations of P. haselbergii. Furthermore, while it is certainly noticeable, as shown in Hunt et al., that there is a diversity in the structure/habit of the tepals, we also know from the recent molecular analysis (Nyffeler and Eggli, 2010;Schlumpberger & Renner, 2012: 1347-1348, that the characters of the flowers and the different pollination syndromes, are not indicators of proximity or distance of two evolutionary lines. In this regard, the growth forms of the two taxa are practically indistinguishable. ...
Article
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The history of the ex-segregates of the genus Parodia Spegazzini s.l. (Cactaceae), since 1819 has been reviewed. A rather conservative approach is adopted to sum up the large number of the proposed names within 62 natural species. For each accepted species, a complete synonymy is given down to the subspecies level (varieties are included only in the case of basionyms). An enlarged description is taken from the field data, along with etymology, information about distribution, biome, ecological region and habitat, maps and, where necessary, an update on the conservation status expressed by the IUCN assessors.
... It has been proposed that Trichocereus be included in the related Echinopsis, but this change is not supported by DNA analysis. 20,21 As a consequence, T. pachanoi sometimes appears in the literature as E. pachanoi, T. peruvianus as E. peruviana and, more confusingly, T. bridgesii as E. lageniformis. Moreover, T. pachanoi and T. peruvianus have recently been combined into a single species as T. macrogonus var. ...
Article
Archeological studies in the USA, Mexico and Peru suggest that mescaline, as a cactus constituent, has been used for more than 6000 years. Although it is a widespread cactus alkaloid, it is present in high concentrations in few species, notably the North American peyote (Lophophora williamsii) and the South American wachuma (Trichocereus pachanoi, T. peruvianus and T. bridgesii). Spanish 16th century chroniclers considered these cacti “diabolic”, leading to their prohibition, but their use persisted to our days and has been spreading for the last 150 years. In the late 1800s peyote attracted scientific attention, mescaline was isolated, and its role in the psychedelic effects of peyote tops or “mescal buttons” was demonstrated. Its structure was established by synthesis in 1929, and alternative routes were developed providing larger amounts for pharmacological and biosynthetic research. Although its effects are attributed mainly to its action as a 5-HT2A serotonin receptor agonist, mescaline binds in a similar concentration range to 5-HT1A and 2A receptors. It is largely excreted unchanged in human urine, and its metabolic products are apparently unrelated to its psychedelic properties. Its low potency is probably responsible for its relative neglect by recreational substance users, as the successful search for structure-activity relationships in the hallucinogen field focused largely on finding more potent analogs. Renewed interest in the possible therapeutic applications of psychedelic drugs may hopefully lead to novel insights regarding the commonalities and differences between the actions of individual classic hallucinogens.
... Some authorities maintain that Trichocereus and Echinopsis should be recognized as separate genera while others maintain they should be merged under Echinopsis. In this article the genera are recognized as separate following Hunt et al (2013) based on the molecular research by Schlumpberger & Renner (2012) and personal communication with Roberto Kiesling (Kiesling 1978). The reader should, however, refer to Hunt (2016) in the CITES Cactaceae checklist where 'alternative names' are provided. ...
... Hunt & al. 2006) include Acanthocalycium in Echinopsis s.l. Schlumpberger and Renner (2012) found that Echinopsis s.l. is both polyphyletic and paraphyletic relative to other genera of the subtribe, and accept Acanthocalycium in expanded form, together with numerous further segregates. ...
Article
Sequential flowering of sympatric plant species is a commonly observed phenomenon and is thought to contribute towards maintaining the pollinator community. Sequential flowering is also interpreted as a potential way to reduce competition for pollinators and to reduce interspecific pollen transport. Within-season sequential flowering is well-known for many species assemblages and usually shows a staggering over weeks and months. Here, we report on the flowering behaviour of two unrelated sympatric cacti (both Cactaceae — Cactoideae) from N Argentina, Acanthocalycium thionanthum (Cereeae — Trichocereinae) and Parodia microsperma (Notocacteae). At the start of the flowering season, the two taxa exhibit a temporally stable sequential mass-flowering that is triggered by the first rainfall event after the dry season: On the 6th or 7th day after the rainfall, Parodia microsperma starts a 3-day “big-bang” flowering period, and on the 8th or 9th day after the rainfall, Acanthocalycium thionanthum also starts a 3-day “big-bang” flowering period, with 1 day of overlap with that of Parodia. The sequence and duration, including the 1-day overlap, has been found to be temporally stable in all study years, with c. 90% of all flowers buds of a population opening synchronously in years with average spring rainfall. Later in the season, after the fruits derived from these initial flowerings have ripened, opportunistic flowering (including sporadic local small-scale mass flowerings) is exhibited by both species with no apparent synchronization or sequence. Only once was a second “big bang” flowering observed, likely caused by above-average dry conditions. The species pair is notable for the massive “big-bang” flowering triggered by rainfall with a tightly staggered synchronization, as well as for the change in flowering strategy later in the season.
... Weber) Britton & Rose) (Walters et al., 2011). Molecular studies by Schlumpberger & Renner (2012) suggest including Trichocereus (A. Berger) Riccob. in Echinopsis Zucc. ...
Article
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Recent field work in Gran Canaria (Canary Islands, Spain), mostly in March and April 2017, yielded first records of 10 species of Cactaceae, all originally introduced as ornamentals. Cylindropuntia bigelovii, C. imbricata, C. prolifera, Disocactus speciosus, Opuntia lindheimeri and O. microdasys are locally naturalized or likely to become so, whereas Ferocactus herrerae, Harrisia tetracantha, Lophocereus schottii and Trichocereus spachianus are considered casuals. All these species are illustrated, georeferenced localities provided and other useful information, especially with regard to their identification and invasion status (in Gran Canaria as well as elsewhere in the world) are also presented. The presence of two further species, Opuntia monacantha and O. robusta, both often considered doubtful in the Canary Islands, is confirmed from Gran Canaria. New data are also provided for Opuntia leucotricha and O. pilifera, two species that were recently recorded for the first time for Gran Canaria, that seem to have increased.
... Columnar cacti with large flowers that were accommodated in Trichocereus (including T. huascha) have been transferred to Echinopsis based on molecular data (Schlumpberger & Renner, 2012). However, pending additional data, Ritter's concept of Trichocereus is here followed (Ritter, 1980). ...
Article
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Summary: Recent field work in Gran Canaria and Tenerife (Canary Islands, Spain) yielded records for 30 alien taxa of Cactaceae that had not been reported before, either from the whole area, or from one of the islands. Out of these, 17 are considered locally naturalised and/or potentially invasive: Cylindropuntia bigelovii, C. fulgida, C. pallida, C. prolifera, C. tunicata, Echinocereus rigidissimus, Haageocereus kagenekii, Hylocereus triangularis, Opuntia basilaris, O. elatior, O. ficus-indica × O. tomentosa, O. macrocentra, O. microdasys, O. pilifera, Oreocereus pseudofossulatus, Tephrocactus articulatus and Trichocereus huascha. The same applies to a rather characteristic form of O. ficus-indica that sometimes is referred to as f. amyclaea (syn.: O. megacantha). The presence of Opuntia monacantha and O. robusta, two species with a dubious status in the Canary Islands, is confirmed. Ten further taxa are considered casuals, often relics of cultivation. All taxa are illustrated, and for the naturalised and/or potentially invasive taxa additional information is provided. Two new combinations are proposed for Cylindropuntia fulgida f. mamillata and Tephrocactus articulatus f. papyracanthus.
... These recurrent hybridization processes could result in complex phylogenetic placement, which in turn increased incongruences among Ilex species. In addition, it is believed that plants in similar habitats located at distant regions may exhibit similar morphologies due to adaptation to similar environments, which influences the consistency between molecular and morphological phylogenies (Clark et al., 2012;Loizeau et al., 2005;Qiu et al., 1995;Schlumpberger & Renner, 2012). Thus, it was not surprising that discordances between phylogenetic and morphological patterns were frequently found in the Ilex species because of their worldwide distribution. ...
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To investigate the phylogenetic relationships among Ilex species distributed in China, we analyzed two alignments including 4,698 characters corresponding to six plastid sequences (matK, rbcL, atpB-rbcL, trnL-F, psbA-trnH, and rpl32-trnL) and 1,748 characters corresponding to two nuclear sequences (ITS and nepGS). Using different partitioning strategies and approaches (i.e., Bayesian inference, maximum likelihood, and maximum parsimony) for phylogeny reconstruction, different topologies and clade supports were determined. A total of 18 Ilex species was divided into two major groups (group I and II) in both plastid and nuclear phylogenies with some incongruences. Potential hybridization events may account, in part, for those phylogenetic uncertainties. The analyses, together with previously identified sequences, indicated that all 18 species were recovered within Eurasia or Asia/North America groups based on plastid data. Meanwhile, the species in group II in the nuclear phylogeny were placed in the Aquifolium clade, as inferred from traditional classification, whereas the species in group I belonged to several other clades. The divergence time of most of the 18 Ilex species was estimated to be not more than 10 million years ago. Based on the results of this study, we concluded that paleogeographical events and past climate changes during the same period might have played important roles in these diversifications.
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The competition between floristic catalogues and the nomenclatural issues of the treated taxa, is a problem for the botanical knowledge of countries. Consequently, it seems to be necessary to merge former taxonomical proposals into a unified list based on phylogenetic hypotheses, the rules of nomenclature and dichotomous keys to the Chilean subfamilies, tribes and genera. With this approach we here propose an updated catalogue of the Chilean cacti. It would be necessary to merge the various taxonomic proposals into a unified list based on both phylogenetic hypotheses and the rules of nomenclature. With this approach, we here propose to updated the catalogue of Chilean cactus. A neotype was designated for Echinocactus jussieui. In addition, we present a dichotomous taxonomic key to the Chilean subfamilies, tribes, and genera.
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Abstract Floral nectaries are essential for plant reproduction but little is known about the relationship between these secretory structures and pollination system in cacti. To test phenotypic patterns in nectaries associated with pollination syndromes and/or with its pollinators, we selected from evolutionarily related genera Cleistocactus, Denmoza, and Echinopsis, a set of species with bird-pollinated flowers and floral traits that may fit with ornithophily or with sphingophily, and other set of sphingophilous species with moths as effective pollinator. Observations were made under light microscope and scanning and transmission electron microscopes. Nectaries are located at the base of the filaments welded to the tube, forming a chamber. The nectary consists of the epidermis with distinctive features in each genus, a secretory parenchyma which may be vascularized and a non-secretory vascularized parenchyma. Anatomical variants observed in nectaries of different species are not consistent with the floral pollination syndromes neither with groups of pollinators. The basic structure of the nectar chamber is relatively conserved, a fact that may be explained by phylogenetic conservatism among the genera investigated. Our results revalue the role of anatomical traits for the systematics of Cactaceae.
Article
In this study, the effects of heat stress (30 °C, 35 °C, 40 °C) on pollen performance of Echinopsis chamaecereus Friedrich & Glaetzle were analyzed. Heat-treated pollen grains showed a insignificant reduction in germination rate. Pollen tubes treated by temperatures above 30 °C exhibited a decrease in tube length and a negative effect on pollen performance according to cumulative stress response index data. Pollen tube abnormality was enhanced by increased temperature, and bifurcated pollen tubes were very dominant than the other abnormalities. Callose accumulation on tube tips was enhanced by increased temperature. In bifurcated pollen tubes, one of the tubes blocked by callose plugs or dense callose accumulation was observed on tips. Actin organization showed disruption, and anisotropy of actin filaments increased at temperatures above 30 °C. The findings show that high temperatures cause destructive effects on male reproductive biology even in species such as cacti known to be temperature tolerant.
Chapter
Phylogenies based on molecular characters has dominated publications rather than those based on morphological characters. Some authors have defended the use of morphology in phylogenetic reconstruction. In Cactaceae few studies have been made combining molecular and morphological characters. A good example about the use of morphology in phylogenetic analysis has been addressed in Echinocereus. Echinocereus is a morphologically diverse genus including 67 species that have been grouped into eight taxonomic sections based on morphological traits. Previous molecular phylogenetic analyses did not show entirely the relationships in Echinocereus species, and did not provide useful characters to recognize clades. Therefore, we performed a combined phylogenetic analysis with a set of 44 morphological characters and six chloroplast DNA sequences. Topologies from parsimony and Bayesian analyses resulted mostly congruent. However, relationships of E. poselgeri did not agree between analyses. A second bayesian analysis using long-branch extraction test resulted in a topology with a morphologically congruent position of E. poselgeri. Parsimony and Bayesian analyses corroborated the monophyly of Echinocereus, which included eight monophyletic groups. The clades did not recover the recent infrageneric classification. As a consequence, a new sectional classification for Echinocereus is proposed based on the eight recovered clades, which are supported by a combination of morphological and molecular characters. An identification key for sections in the genus is included.
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The genus Agave sensu lato contains ca. 211 described species, many of which are considered keystone species because of their ecological dominance and the quantity of resources they provide with their massive, nectar-rich inflorescences. The large diversity of Agave species has been hypothesized as being related to their reproductive strategy (predominantly monocarpic) and diverse pollinators (e.g., bats, hummingbirds, hawkmoths). In particular, Agave species provide resources that a few genera of nectar feeding bats from the subfamily Glosophaginae are dependent upon. To explore a possible coevolutionary relationship between Agave and the bat species that pollinate them, we calibrated molecular phylogenies of both groups and looked for a correlation in their dates of divergence. One coding and two non-coding regions of the chloroplast genome were sequenced from 49 species of the Agavoideae (Asparagaceae), and the mitochondrial gene Cyt-b and nuclear coding gene RAG2 were either sequenced or obtained from gene bank for 120 Phyllostomid bats. Results from the analyses indicate that Agave sensu lato is a young genus (estimated crown age 2.7–8.5/stem age 4.6–12.3 Ma), with an increasing diversification rate, and the highest speciation rate among Agavoideae’s clades. The origin of the Glossophaginae bats (stem age 20.3–23.5 Ma) occurred prior to the stem age of Agave sensu lato, while the origin of the current pollinators of Agave species, members of the genera Glossophaga, Leptonycteris, Anoura, Choeronyscus, Musonycteris and Choeronycteris, was estimated to be around 6.3–16.2 Ma, overlapping with the stem age of Agave sensu lato, supporting the hypothesis of diffuse coevolution. Link to the full-text: https://authors.elsevier.com/c/1YRKb3m3nMuJlM
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Pollinator-mediated selection on plants can favor transitions to a new pollinator depending on the relative abundances and efficiencies of pollinators present in the community. A frequently observed example is the transition from bee pollination to hummingbird pollination. We present a population genetic model that examines whether the ability to inbreed can influence evolutionary change in traits that underlie pollinator attraction. We find that a transition to a more efficient but less abundant pollinator is favored under a broadened set of ecological conditions if plants are capable of delayed selfing rather than obligately outcrossing. Delayed selfing allows plants carrying an allele that attracts the novel pollinator to reproduce even when this pollinator is rare, providing reproductive assurance. In addition, delayed selfing weakens the effects of Haldane’s sieve by increasing the fixation probability for recessive alleles that confer adaptation to the new pollinator. Our model provides novel insight into the paradoxical abundance of recessive mutations in adaptation to hummingbird attraction. It further predicts that transitions to efficient but less abundant pollinators (such as hummingbirds in certain communities) should disproportionately occur in self-compatible lineages. Currently available mating system data sets are consistent with this prediction, and we suggest future areas of research that will enable a rigorous test of this theory.
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We present the catalogue of the Cactaceae family of the department of Arequipa, Peru; for which we made field trips between years 2010 and 2016, from 0 to above 4500 m of elevation, we obtained data from national and foreign herbaria, reviewed specialized literature, observed images of the types (HSP, F, B, K, ZZS, HEID, US and U) and consulted specialists. It consists of 56 non-cultivated taxa, grouped into 2 subfamilies (Opuntioideae and Cactoideae), 7 tribes and 20 genera. The most representative taxa correspond to Cumulopuntia (11 spp.), Corryocactus (7 spp.) and Loxanthocereus (6 spp.); 21 of these taxa are endemic and the most diverse of them are Cumulopuntia with 6 spp., Loxanthocereus with 3 spp. and Corryocactus with 3 spp. The taxa are listed in alphabetical order and for each taxon is indicated the scientific name, synonyms and basionyms, bibliographic references, habit, ecology, altitudinal distribution, specimen voucher, departmental distribution, provincial distribution and conservation status. In addition, a dichotomous key is presented for the genera of Arequipa and the taxonomy and distribution are discussed.
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The Cactaceae is one of the most conspicuous and ecologically important plant families in the world; its species may have specialist or generalist pollination systems that show geographical patterns, which are synthesized in the Geographical Dichotomy Hypothesis. Here, we assess this hypothesis in five countries in both tropical and extratropical regions, evaluating the pollinator visitation rate and pollinator identity and abundance. We calculate the Shannon diversity index (H') and evenness (J) and evaluate the differences between latitude parameters by a t-student test. Overall, we found more specialized pollination systems in all tropical sites; the richness, diversity and evenness of pollinators was reduced in comparison to extratropical regions where the pollination system was generalized. Our results support the geographical dichotomy hypothesis in the cacti of South America, suggesting that environmental factors behind the latitudinal patterns can help to explain the difference in the pollination syndrome between tropical and extratropical regions. This article is protected by copyright. All rights reserved.
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1 Three synchronopatric Cactaceae species, Echinopsis rhodotricha, Harrisia balansae and Praecereus saxicola, have mostly nocturnal anthesis and similar flowers, characteristics which motivated us to perform a comparative study of reproductive ecology. 2 Reproductive phenology was sampled monthly from December 2014 to November 2016. We describe floral biology, breeding system via pollination treatments and evaluate floral visitors from focal and filming observations. Pollen grains found on moth proboscis were compared among cactus species under light microscopy. We used fluorescent dye particles to test intra- and interspecific pollen flow. 3 These three species have extended flowering with greater intensity in the wet season, causing high overlap. They have white and hypocrateriformis flowers that open at twilight or nightfall and last about 15 hours. Harrisia balansae seems to be self-incompatible (SI), while E. rhodotricha presented self-compatibility (SC). Praecereus saxicola presented self-fertility, but most of the population seems to be self-incompatible. We suggest sphingophily for the three species, but only P. saxicola was visited by Manduca rustica (Sphingidae). However, we observed pollen grains of all three species on the proboscis of moths, especially M. rustica and M. sexta. Prolonged anthesis allowed bees (herein considered as secondary pollinators) to visit flowers of E. rhodotricha and P. saxicola. 4 It can be concluded that the studied species share nocturnal and diurnal pollinators, suggesting interspecific pollen flow, which, however, could not be detected by fluorescent dye particles. In view of the low frequency of primary pollinators, it appears that these three species have different reproductive strategies, ensuring the fruiting and production of viable seeds. This article is protected by copyright. All rights reserved.
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Echinocereus is a morphologically diverse genus that includes 64 species grouped into eight taxonomic sections based on morphological traits. In previous molecular phylogenetic analyses, the relationships amongst Echinocereus species were not entirely revealed and useful characters to recognize clades were not provided. The inclusion of several sources of evidence in a phylogenetic analysis is likely to produce more supported hypotheses. Therefore, we performed a combined phylogenetic analysis with a set of 44 morphological characters and six chloroplast DNA sequences. Topologies from parsimony and Bayesian analyses were mostly congruent. However, the relationships of E. poselgeri were not consistent between analyses. A second Bayesian analysis using a long-branch extraction test resulted in a topology with the morphological position of E. poselgeri congruent with that in parsimony analysis. Parsimony and Bayesian analyses corroborated the monophyly of Echinocereus, which included eight monophyletic groups. The combined phylogeny integrated into different clades those taxa that were not determined in previous analyses and changed the relationships of some recognized clades. The clades did not recover the recent infrageneric classification. In the present study, a new sectional classification for Echinocereus is proposed based on the eight recovered clades, which is supported by a combination of morphological and molecular characters. An identification key for sections in the genus is included.
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Analyses of meiotic and mitotic chromosomes were undertaken in 16 taxa of Echinocereus belonging to 12 species and all seven taxonomic sections (sensu Taylor). Chromosome numbers are reported for the first time for eight taxa, and previously published chromosome counts are confirmed for the remaining eight. Both diploid and polyploid counts were obtained. Eleven (69%) of the taxa surveyed were diploid (2n = 22); the five varieties of E. engelmannii were polyploid (2n = 44). Overall, chromosome counts are available for 23 of the 48 proposed species (sensu Taylor). Of these, 19 (82%) are diploid, and four (18%) are polyploid. Polyploid cytotypes are most common in the primitive sections, e.g., sections Erecti and Triglochidiatus, which suggests that polyploidy is probably a derived condition in Echinocereus. Polyploid taxa range from medium to high latitudes and elevations relative to the overall distribution of the genus. Polyploidy, hybridization, and cryptic chromosomal rearrangements are thought to be the major causes of the speciation events of the genus.
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I present data on flower morphology, pollination, breeding system, and genetic diversity of the columnar cactus Weber-bauerocereus weberbaueri at 2500 m elevation in southwestern Peru. Weberbauerocereus weberbaueri is a self-compatible columnar cactus that is visited and pollinated by one species of rare endemic bat, Platalina genovensium, and two species of hummingbirds, Patagona gigas and Rhodopis vesper. W. weberbaueri exhibits pronounced interplant variation in floral color and size, and flowers exhibit traits corresponding to both bat and hummingbird pollination syndromes. Starch-gel electrophoresis of flower bud tissue indicated that W. weberbaueri is an autotetraploid and that genetic diversity (Hep = 0.257) of the study population was high relative to diploid plants but similar to other tetraploid species. Initial fruit set from pollinator exclusion experiments conducted in 1991 and 1993, at the onset of and after a drought associated with the El Niño event of 1991–1992, revealed that bats were the most important pollinators in 1991, but that hummingbirds and diurnal insects were most important in 1993. In both years, however, autogamy and lepidopteran larval infestation of fruits reduced differences in mature fruit production among pollinator exclusion treatments so that differences in mature fruit set were not statistically significant. Reduced bat pollination in 1993 is attributed to the reduced abundance of bats at the study site during a drought caused by El Niño. I hypothesize that interaction among several factors, including tetraploidy, autogamy, larval infestation of developing fruits, and variation in pollinator abundance, may not result in strong selection for a bat vs. hummingbird floral morph, thus allowing the persistence of floral variation in this cactus.
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Natural hybridization has been frequently reported in Cactaceae, but there are few studies on the impact of hybridization in the evolution and diversification of this family. The majority of the studies that have addressed this subject have been restricted to the subfamily Opuntioideae. Little is known about the role of hybridization on the more diverse subfamily Cactoideae. This paper discusses putative examples of hybrid lineages in subfamily Cactoideae, focussing on examples of lineages for which hybrid or introgressive origin have been confirmed or deduced from morphological and molecular studies. These examples suggest that natural hybridization may have played an important role in the evolution and diversification of taxa in Cactaceae, and further studies of hybrid lineages may provide us with important insights on the evolution of this family.
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First-time meiotic chromosome determinations were made for some South American Cactaceae: Armatocereus godingianus (2n = 22), Cleistocactus leonensis (2n = 22), C. sepium var. morleyanus (2n = 22), C. sepium var. sepium (2n = 22), C. sepium var. ventimigliae (2n = 22), Opuntia xaequatorialis (2n = 66), O. bakeri (2n = 99), O. pubescens (2n = 44), O. pubescens x O. soederstromiana (2n = 88), O. quitensis (2n = 22), O. soederstromiana (2n = 88), and Pilosocereus lanuginosus (2n = 22). New chromosome determinations were made for Austrocylindropuntia cylindrica (2n = 88) and A. subulata (2n = 110). A chromosome determination for O. ficus-indica (2n = 88) agreed with previous work. In Ecuador, the hexaploid hybrid O. Xaequatorialis, the nanoploid O. bakeri, and two unnamed octoploids are morphologically intermediate between the widespread O. pubescens and the endemic O. soederstromiana and may represent hybrids between these two species.
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Understanding evolutionary responses of plants to desert environments depends upon phylogenetic knowledge of desert plants. The diverse American desert family Cactaceae has been presumed, on the basis of distinctiveness, to be phylogenetically isolated and relatively ancient (> 65 million years old). Using maximum likelihood and parsimony analyses of the rapidly evolving internal transcribed spacer (ITS) sequences of nuclear ribosomal DNA (nrDNA), we show that the cacti are phylogenetically nested among other aridity-adapted lineages of the angiosperm family Portulacaceae. The ITS divergence between pereskioid cacti and the genus Talinum (Portulacaceae) is less than that between many Portulacaceae genera. Synthesis of the ITS data with morphological and chloroplast DNA evidence suggests an origin of cacti in mid-Tertiary, c 30 million years ago, and a later Tertiary diversification coincident with development of the American desert. This, in turn, implies that the diversification rate in cacti was much higher than in their nearest relatives. The present results illustrate the central role of phylogenetic reconstruction in ecological and evolutionary theory.
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Mitotic chromosomes of 11 diploid (2n = 22) species of Echinocereus and five varieties of the polyploid (2n = 4x = 44) E. engelmannii included in the seven taxonomic sections of the genus were studied. The genus exhibits relatively symmetric karyotypes, having mostly metacentric chromosomes. Likewise, interspecific and intraspecific variability was observed in terms of genome length, chromosome length, and karyotipic formula. There was also variability in the number of satellites, and they were observed on the short arms of the chromosomes. There is an apparent correlation in decreasing chromosome and genome length in species with more derived morphological characters. In addition, karyotype asymmetry increases as the chromosome and genome length increases. Polyploid taxa investigated had larger chromosomes and genomes than diploid taxa. Robertsonian changes and centric fusion of telocentric chromosomes, are thought to be involved in the existence of homogeneous karyotypes. No evidence of structural chromosomal rearrangements was detected. Overall, the high morphological plasticity that characterizes the genus Echinocereus parallels the relatively large karyotypic variability found in the species investigated.
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Documented meiotic and mitotic chromosome numbers and behavior are reported for 62 taxa representing 11 genera of Cactaceae from the southwestern United States and northern Mexico. Chromosome numbers for 12 species and six natural interspecific hybrids are first counts. New numbers were determined for four additional species. Chromosome counts of diploid and polyploid taxa, including triploid and pentaploid hybrids, are all consistent with the base number, x = 11. Translocations are reported for the first time in the Cactaceae.
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I present data on flower morphology, pollination, breeding system, and genetic diversity of the columnar cactus Weberbauerocereus weberbaueri at 2 500 m elevation in southwestern Peru. Weberbauerocereus weberbaueri is a self-compatible columnar cactus that is visited and pollinated by one species of rare endemic bat, Platalina genovensium, and two species of hummingbirds, Patagona gigas and Rhodopis vesper. W. weberbaueri exhibits pronounced interplant variation in floral color and size, and flowers exhibit traits corresponding to both bat and hummingbird pollination syndromes. Starch-gel electrophoresis of flower bud tissue indicated that W. weberbaueri is an autotetraploid and that genetic diversity (H-ep = 0.257) of the study population was high relative to diploid plants but similar to other tetraploid species. Initial fruit set from pollinator exclusion experiments conducted in 1991 and 1993, at the onset of and after a drought associated with the Fl Nino event of 1991-1992, revealed that bats were the most important pollinators in 1991, but that hummingbirds and diurnal insects were most important in 1993. In both years, however, autogamy and lepidopteran larval infestation of fruits reduced differences in mature fruit production among pollinator exclusion treatments so that differences in mature fruit set were not statistically significant. Reduced bat pollination in 1993 is attributed to the reduced abundance of bats at the study site during a drought caused by El Nino. I hypothesize that interaction among several factors, including tetraploidy, autogamy, larval infestation of developing fruits, and variation in pollinator abundance, may not result in strong selection for a bat vs. hummingbird floral morph, thus allowing the persistence of floral variation in this cactus.
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The phylogenetic relationships and chromosomal evolution of the diverse tropical genus Passiflora (Passifloraceae) are explored using data from two chloroplast markers: the rpoC1 intron and the trnL/trnT spacer region. A survey of the presence or absence of the rpoC1 intron in 136 species representing 17 of Killip's (1938) 22 subgenera of Passiflora and four other genera in the Passifloraceae revealed intron losses in 46 taxa. A minimum of two losses were confirmed by a parametric bootstrap approach on sequence data from the trnL/trnT chloroplast non-coding region for 61 taxa. The results of phylogenetic analyses of the trnL/trnT sequence data support the reduction of Killip's 22 subgenera to four as proposed in a new classification system by Feuillet and MacDougal (2004). The monophyly of the ‘n=6’ and ‘n=9’ chromosomal and morphological groups is strongly supported. In addition, these data indicate that Passiflora biflora, or closely related species, is the likely continental sister to the red-flowered Caribbean taxa, while P. auriculata is weakly supported as the New World sister to the Old World Passifloras. Finally, character optimization of chromosome numbers on the phylogenetic tree supports x=12 as the base chromosome number for Passiflora.
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The genus Aristolochia sensu lato contains over 400 species from warm temperate to tropical regions worldwide. Taxonomic treatments of Aristolochia have been ambiguous and controversial. In a recent cladistic analysis based on morphological characters, it was proposed that the genus should be divided into four genera in two subtribes. To reconsider the systematics of Aristolochia sensu lato, we reconstructed its phylogeny based on nucleotide sequences of the chloroplast rbcL gene and the nuclear-encoded phytochrome A (phyA) gene for 19 representative species and the chloroplast matK gene of over 80 species. All phylogenetic trees produced with the three genes indicate that Aristolochia sensu lato is a monophyletic group, consisting of two lineages that correspond to the subtribes Aristolochiinae and Isotrematinae. The matK phylogeny shows that each of the lineages includes two sublineages. The Aristolochiinae clade is composed of the Aristolochia sensu stricto and Pararistolochia clades, and the Isotrematinae clade of the Isotrema and Endodeca clades. Chromosome numbers, including newly reported counts for 30 species, are predominantly congruent with the phylogeny: the Aristolochiinae clade shows chromosome numbers of 2n = 6, 12, 14, or 16, while the Isotrematinae clade is characterized by 2n = 32. In the Isotrematinae clade, the paralogous relationships of the phyA gene suggest that polyploidization might have occurred.
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Phylogenetic analysis of amplified fragment length polymorphisms (AFLP) was used to infer pat-terns of morphologic and chromosomal evolution in an eastern North American group of sedges (ENA clade I of Carex sect. Ovales). Distance analyses of AFLP data recover a tree that is topologically congruent with previous phylogenetic estimates based on nuclear ribosomal DNA (nrDNA) sequences and provide support for four species groups within ENA clade I. A maximum likelihood method designed for analysis of restriction site data is used to evaluate the strength of support for alternative topologies. While there is little support for the precise placement of the root, the likelihood of topol-ogies in which any of the four clades identified within the ENA clade I is forced to be paraphyletic is much lower than the likelihood of the optimal tree. Chromosome counts for a sampling of species from throughout sect. Ovales are mapped onto the tree, as well as counts for all species in ENA clade I. Parsimony reconstruction of ancestral character states suggest that: (1) Heilborn's hypothesis that more highly derived species in Carex have higher chromosome counts does not apply within sect. Ovales, (2) the migration to eastern North America involved a decrease in average chromosome count within sect. Ovales, and (3) intermediate chromosome counts are ancestral within ENA clade I. A more precise understanding of chromosomal evolution in Carex should be possible using likelihood analyses that take into account the intraspecific polymorphism and wide range of chromosome counts that characterize the genus.
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Chromosome numbers for a total of 54 individuals representing 13 genera and 40 species of Cactaceae, mostly in tribe Trichocereeae, are reported. Five additional taxa examined belong to subfamily Opuntioideae and other tribes of Cactoideae (Browningieae, Pachycereeae, Notocacteae, and Cereeae). Among Trichocereeae, counts for 35 taxa in eight genera are reported, with half of these (17 species) for the genus Haageocereus. These are the first chromosome numbers reported for 36 of the 40 taxa examined, as well as the first counts for the genus Haageocereus. Both diploid and polyploid counts were obtained. Twenty nine species were diploid with 2n=2x=22. Polyploid counts were obtained from the genera Espostoa, Cleistocactus, Haageocereus, and Weberbauerocereus; we detected one triploid (2n=3x=33), nine tetraploids (2n=4x=44), one hexaploid (2n=6x=66), and three octoploids (2n=8x=88). In two cases, different counts were recorded for different individuals of the same species (Espostoa lanata, with 2n=22, 44, and 66; and Weberbauerocereus rauhii, with 2n=44 and 88). These are the first reported polyploid counts for Haageocereus, Cleistocactus, and Espostoa. Our counts support the hypothesis that polyploidy and hybridization have played prominent roles in the evolution of Haageocereus, Weberbauerocereus, and other Trichocereeae.
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Cacti are a large and diverse group of stem succulents predominantly occurring in warm and arid North and South America. Chloroplast DNA sequences of the trnK intron, including the matK gene, were sequenced for 70 ingroup taxa and two outgroups from the Portulacaceae. In order to improve resolution in three major groups of Cactoideae, trnL-trnF sequences from members of these clades were added to a combined analysis. The three exemplars of Pereskia did not form a monophyletic group but a basal grade. The well-supported subfamilies Cactoideae and Opuntioideae and the genus Maihuenia formed a weakly supported clade sister to Pereskia. The parsimony analysis supported a sister group relationship of Maihuenia and Opuntioideae, although the likelihood analysis did not. Blossfeldia, a monotypic genus of morphologically modified and ecologically specialized cacti, was identified as the sister group to all other Cactoideae. The tribe Cacteae was found to be sister to a largely unresolved clade comprising the genera Calymmanthium, Copiapoa, and Frailea, as well as two large and well-supported clades. Browningia sensu stricto (excluding Castellanosia), the two tribes Cereeae and Trichocereeae, and parts of the tribes Notocacteae and Rhipsalideae formed one clade. The distribution of this group is largely restricted to South America. The other clade consists of the columnar cacti of Notocacteae, various genera of Browningieae, Echinocereeae, and Leptocereeae, the tribes Hylocereeae and Pachycereeae, and Pfeiffera. A large portion of this latter group occurs in Central and North America and the Caribbean.
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Polyploidy, an important mechanism of plant evolution, was investigated in Consolea, an endemic Caribbean opuntioid genus represented by nine subdioecious species with very narrow distributions, including species classified as rare or threatened. Standard chromosome counting and flow cytometric analyses were used to determine chromosome numbers and ploidy of each taxon. Compared to the base number (x = 11), the mitotic and meiotic counts indicated that there are seven hexaploid (2n = 66) and two octoploid species (2n = 88); no diploids were found. Histograms of intact nuclei confirmed that all species are polyploid, with C-DNA values ranging from 4.88-9.50 pg. The variation of DNA content was significantly higher for the octoploids than for the hexaploids. Male and female sexual morphs had similar DNA content, suggesting that there are no sex chromosomes. Cytomixis between cells and microsporocytes with no chromatin were observed. This provides a mechanism whereby gametes with variable chromosome numbers are produced, influencing reproduction and promoting speciation. In conclusion, C-DNA content and chromosome number separated Consolea species into two groups, which may correspond to two phylogenetic lineages or indicate that polyploidization occurred independently, with comparable effects on C-DNA content.
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The sunflower genus, Helianthus, is recognized widely for the cultivated sunflower H. annuus and scientifically as a model organism for studying diploid and polyploid hybrid speciation, introgression, and genetic architecture. A resolved phylogeny for the genus is essential for the advancement of these scientific areas. In the past, phylogenetic relationships of the perennial species and polyploid hybrids have been particularly difficult to resolve. Using the external transcribed spacer region of the nuclear 18S-26S rDNA region, we reveal for the first time a highly resolved gene tree for Helianthus. Phylogenetic analysis allowed the determination of a monophyletic annual H. sect. Helianthus, a two-lineage polyphyletic H. sect. Ciliares, and the monotypic H. sect. Agrestis, all of which were nested within a large perennial and polyphyletic H. sect. Divaricati. The distribution of perennial polyploids and known annual diploid hybrids on this phylogeny suggested multiple independent hybrid speciation events that gave rise to at least four polyploids and three diploid hybrids. Also provided by this phylogeny was evidence for homoploid hybrid speciation outside H. sect. Helianthus. Finally, previous hypotheses about the secondary chemistry in the genus were tested in a phylogenetic framework to obtain a better understanding of the evolution of these compounds in Helianthus.
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The cacti are one of the most celebrated radiations of succulent plants. There has been much speculation about their age, but progress in dating cactus origins has been hindered by the lack of fossil data for cacti or their close relatives. Using a hybrid phylogenomic approach, we estimated that the cactus lineage diverged from its closest relatives ≈35 million years ago (Ma). However, major diversification events in cacti were more recent, with most species-rich clades originating in the late Miocene, ≈10-5 Ma. Diversification rates of several cactus lineages rival other estimates of extremely rapid speciation in plants. Major cactus radiations were contemporaneous with those of South African ice plants and North American agaves, revealing a simultaneous diversification of several of the world's major succulent plant lineages across multiple continents. This short geological time period also harbored the majority of origins of C(4) photosynthesis and the global rise of C(4) grasslands. A global expansion of arid environments during this time could have provided new ecological opportunity for both succulent and C(4) plant syndromes. Alternatively, recent work has identified a substantial decline in atmospheric CO(2) ≈15-8 Ma, which would have strongly favored C(4) evolution and expansion of C(4)-dominated grasslands. Lowered atmospheric CO(2) would also substantially exacerbate plant water stress in marginally arid environments, providing preadapted succulent plants with a sharp advantage in a broader set of ecological conditions and promoting their rapid diversification across the landscape.
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Karyotype analyses in members of the four Cactaceae subfamilies were performed. Numbers and karyotype formula obtained were: Pereskioideae = Pereskiaaculeata(2n = 22; 10 m + 1 sm), Maihuenioideae = Maihuenia patagonica (2n = 22, 9 m + 2 sm; 2n = 44, 18 m + 4 sm), Opuntioideae = Cumulopuntia recurvata(2n = 44; 20 m + 2 sm), Cactoideae = Acanthocalycium spiniflorum (2n = 22; 10 m + 1 sm),Echinopsis tubiflora (2n = 22; 10 m + 1 sm), Trichocereus candicans (2n = 22, 22 m). Chromosomes were small, the average chromosome length was 2.3 mum. Diploid species and the tetraploid C. recurvata had one terminal satellite, whereas the remaining tetraploid species showed four satellited chromosomes. Karyotypes were symmetrical. No CMA(-)/DAPI(+) bands were detected, but CMA(+)/DAPI(-) bands associated with NOR were always found. Pericentromeric heterochromatin was found in C. recurvata, A. spiniflorum, and the tetraploid cytotype of M. patagonica. The locations of the 18S-26S rDNA sites in all species coincided with CMA(+)/DAPI(-) bands; the same occurred with the sizes and numbers of signals for each species. This technique was applied for the first time in metaphase chromosomes in cacti. NOR-bearing pair no.1 may be homeologous in all species examined. In Cactaceae, the 18S-26S loci seem to be highly conserved.
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A South American cactus species, Echinopsis ancistrophora (Cactaceae), with dramatic among-population variation in floral traits is presented. Eleven populations of E. ancistrophora were studied in their habitats in northern Argentina, and comparisons were made of relevant floral traits such as depth, stigma position, nectar volume and sugar concentration, and anthesis time. Diurnal and nocturnal pollinator assemblages were evaluated for populations with different floral trait combinations. Remarkable geographical variations in floral traits were recorded among the 11 populations throughout the distribution range of E. ancistrophora, with flower lengths ranging from 4.5 to 24.1 cm. Other floral traits associated with pollinator attraction also varied in a population-specific manner, in concert with floral depth. Populations with the shortest flowers showed morning anthesis and those with the longest flowers opened at dusk, whereas those with flowers of intermediate length opened at unusual times (2300-0600 h). Nectar production varied non-linearly with floral length; it was absent to low (population means up to 15 microL) in short- to intermediate-length flowers, but was high (population means up to 170 microL) in the longest tubed flowers. Evidence from light-trapping of moths, pollen carriage on their bodies and moth scale deposition on stigmas suggests that sphingid pollination is prevalent only in the four populations with the longest flowers, in which floral morphological traits and nectar volumes match the classic expectations for the hawkmoth pollination syndrome. All other populations, with flowers 4.5-15 cm long, were pollinated exclusively by solitary bees. The results suggest incipient differentiation at the population level and local adaptation to either bee or hawkmoth (potentially plus bee) pollination.
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Despite recent advances achieved by application of high-performance computing methods and novel algorithmic techniques to maximum likelihood (ML)-based inference programs, the major computational bottleneck still consists in the computation of bootstrap support values. Conducting a probably insufficient number of 100 bootstrap (BS) analyses with current ML programs on large datasets-either with respect to the number of taxa or base pairs-can easily require a month of run time. Therefore, we have developed, implemented, and thoroughly tested rapid bootstrap heuristics in RAxML (Randomized Axelerated Maximum Likelihood) that are more than an order of magnitude faster than current algorithms. These new heuristics can contribute to resolving the computational bottleneck and improve current methodology in phylogenetic analyses. Computational experiments to assess the performance and relative accuracy of these heuristics were conducted on 22 diverse DNA and AA (amino acid), single gene as well as multigene, real-world alignments containing 125 up to 7764 sequences. The standard BS (SBS) and rapid BS (RBS) values drawn on the best-scoring ML tree are highly correlated and show almost identical average support values. The weighted RF (Robinson-Foulds) distance between SBS- and RBS-based consensus trees was smaller than 6% in all cases (average 4%). More importantly, RBS inferences are between 8 and 20 times faster (average 14.73) than SBS analyses with RAxML and between 18 and 495 times faster than BS analyses with competing programs, such as PHYML or GARLI. Moreover, this performance improvement increases with alignment size. Finally, we have set up two freely accessible Web servers for this significantly improved version of RAxML that provide access to the 200-CPU cluster of the Vital-IT unit at the Swiss Institute of Bioinformatics and the 128-CPU cluster of the CIPRES project at the San Diego Supercomputer Center. These Web servers offer the possibility to conduct large-scale phylogenetic inferences to a large part of the community that does not have access to, or the expertise to use, high-performance computing resources.
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Six primers for the amplification of three non-coding regions of chloroplast DNA via the polymerase chain reaction (PCR) have been designed. In order to find out whether these primers were universal, we used them in an attempt to amplify DNA from various plant species. The primers worked for most species tested including algae, bryophytes, pteridophytes, gymnosperms and angiosperms. The fact that they amplify chloroplast DNA non-coding regions over a wide taxonomic range means that these primers may be used to study the population biology (in supplying markers) and evolution (inter- and probably intraspecific phylogenies) of plants.
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Chromosome counts and observations of reproduction for 55 taxa of Cactaceae indicate that polyploidy is correlated with self-fertility, adventive embryony, profuse branching, and vegetative reproduction. Six genera (Blossfeldia. Cleistocactus, Frailea, Pelecyphora, Rebutia, and Strombocactus) and 35 species or varieties are reported here for the first time. Preliminary observations of pachytene and diplotene indicate that these stages may be more useful in chromosome recognition than mitotic stages. Secondary association at metaphase I and II is interpreted as a retention of homologue association at interphase I and II (interkinesis). During meiosis of certain species, Feulgen negative bodies are present. The production of an abnormal premeiotic division is suggested as a mechanism for polyploid origin.
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Flower–pollinator interactions are among the most fascinating research areas in ecology. The ways in which such interactions evolve are influenced by a number of biotic and abiotic factors, such as habitat and climate, plant and animal morphology, phylogenetic background etc. The increasing amount of data on pollination ecology combined with more and more detailed and precise phylogenetic reconstructions based on (not only) molecular evidence, allows improvement of our understanding of how flower–pollinator relationships evolve. Plant lineages especially well-suited for the study of factors influencing the evolution of flower–pollinator interactions would contain pollination by various groups of pollinators, preferably both diurnal and nocturnal, including both vertebrates and invertebrates, and sufficient evolutionary shifts among pollination modes. Cactaceae are ideal for this type of study for a number of reasons: manageable number of species great diversity of habitats diverse growth forms adaptation to several pollinator guilds, including bimodal and generalized pollination an increasingly solid phylogenetic base of data.
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The chromosome number of 74 species and varieties in Cactaoeae were observed. 1 . The polyploidy of the. each group are as follows., 1 : 2x(2), II :2x(5) 3x(1) 4x(4) 6x(3) 6x(3) 10x( I ) 12x( I ), III : 2x(13) 4x(4). IV : 2x(36) 4x( 1), In these: polyploids 3x, 8x. 10x and 12x have not been known. 2 . The basic number of chr'omosorile in Cactaoeae including Pereskieae is n=11. 3 . No heteroploid was found in this study. 4. The pollen rtLother cells divid, ed at about noon, and the fact ls in accord, with the observation reported by Miss. BEARD. 5. The division of cells in root tip occurred from 7 to 8 o'clock a.m. or from 4 t, o 6 o'clock p. nl. and the observation is somewhat differ from Miss. i3EARD's report. For ascertain the diffefence above mentioned, the root tips of two species-Trichocereus spachianus and Notocactus sudmammulosus-were fixed at intervals of one half or tWo hours duririg 7 o'clock a. m. to 6 o'clock p. m: In the root tips fixed at noon the ', division of cells were scarcely found, but there were many dividing cells fixed at morning Or evening. e . The chromosomes of cacti are very small, and.: there are some difference in size among each species . The size of chromosomes in tropical or desert plants are small in many cases, but not absolutely. So it may not be said that the smaller chromosomes. ls not related to the tropical or succulent plants: 7. There was not found real relation between polyploidy and cold reslstance 8. There was not recognized ariy relation between polyploidy and other characters .
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Columnar cacti can be pollinated by various insect and vertebrate species, often showing either close adaptations to a single pollinator group, or a mixed pollination syndrome, depending on a variety of different animals for pollination. As pollination data for columnar cacti in South America are scarce, we investigated the spectrum, frequency, and behavior of floral visitors in Echinopsis atacamensis subsp. pasacana. We compared two habitats with differences in mean annual precipitation and human impact, finding considerably higher activity and diversity of flower visitors in the population with higher humidity. Bees, wasps, and the giant hummingbird (Patagona gigas) were shown to visit cactus flowers during the day to gather nectar or pollen. We found marked differences in the spectrum of floral visitors between both populations. In one study site, the introduced honeybee Apis mellifera removed large quantities of pollen, but seemed to be a comparatively inefficient pollinator. Additionally, first evidence for visits by nocturnal hawkmoths is provided. These results demonstrate an unspecialized floral syndrome for Echinopsis atacamensis subsp. pasacana, with both diurnal and nocturnal pollinators. Such a mixed floral syndrome is fairly widespread among columnar cacti in the northern hemisphere but was hitherto only shown for one South American species, Weberbauerocereus weberbaueri.
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Chromosome counts and observations of reproduction for 55 taxa of Cactaceae indicate that polyploidy is correlated with self-fertility, adventive embryony, profuse branching, and vegetative reproduction. Six genera (Blossfeldia, Cleistocactus, Frailea, Pelecyphora, Rebutia, and Strombocactus) and 35 species or varieties are reported here for the first time. Preliminary observations of pachytene and diplotene indicate that these stages may be more useful in chromosome recognition than mitotic stages. Secondary association at metaphase I and II is interpreted as a retention of homologue association at interphase I and II (interkinesis). During meiosis of certain species, Feulgen negative bodies are present. The production of an abnormal premeiotic division is suggested as a mechanism for polyploid origin.