Get the shovel: morphological and evolutionary complexities of belowground organs in geophytes.

Herbaceous plants collectively known as geophytes, which regrow from belowground buds, are distributed around the globe and throughout the land plant tree of life. The geophytic habit is an evolutionarily and ecologically important growth form in plants, permitting novel life history strategies, enabling the occupation of more seasonal climates, mediating interactions between plants and their water and nutrient resources, and influencing macroevolutionary patterns by enabling differential diversification and adaptation. These taxa are excellent study systems for understanding how convergence on a similar growth habit (i.e., geophytism) can occur via different morphological and developmental mechanisms. Despite the importance of belowground organs for characterizing whole-plant morphological diversity, the morphology and evolution of these organs have been vastly understudied with most research focusing on only a few crop systems. Here, we clarify the terminology commonly used (and sometimes misused) to describe geophytes and their underground organs and highlight key evolutionary patterns of the belowground morphology of geophytic plants. Additionally, we advocate for increasing resources for geophyte research and implementing standardized ontological definitions of geophytic organs to improve our understanding of the factors controlling, promoting, and maintaining geophyte diversity.

Spatio-temporal evolution of climbing habit in the Dahlia-Hidalgoa group (Coreopsidae, Asteraceae).

In this paper we focus on Hidalgoa, a small genus distributed in cloud forest from Mexico to Colombia with the uncommon climbing habit of enlarging its petioles. This genus belongs to tribe Coreopsideae (Asteraceae), a group mostly from the Neotropics with a few taxa in Oceania and Polynesia. The phylogenetic position of Hidalgoa has not yet been determined though it will most probably be closely related to Dahlia. The aim of this paper is to understand the ecological and evolutionary consequences of the shift to a climbing growth form using a spatio-temporal approach to phylogenetic analyses, and by testing evolutionary models for ecological niche. Our results identified that Hidalgoa formed a sub- clade in a grade with Dahlia, with divergence occurring in the Pliocene. The ancestral life form for this clade was cryptophytic (e.g. geophytes). Analyses using the climate variables most important to the elements of the Dahlia-Hidalgoa clade indicate that they share the same preferences. Moreover, visualization by traitgrams of the significant climate variables shows no departure among clades. These results suggest that no evolutionary or ecological consequences have yet manifested, and that Hidalgoa can be considered a climbing Dahlia.

Ups and downs: Genetic differentiation among populations of the Podocarpus (Podocarpaceae) species in Mesoamerica.

The biogeographical history of Mesoamerican cloud forests is complex, encompassing a diverse and heterogeneous mixture of species with temperate and tropical origins. The dynamic geological landscape and climate change from the Miocene to the Pleistocene affected the distributions and composition of cloud forests in the region, and contributed to divergence events at different time scales. We assessed genetic variation of 29 populations of P. matudae, and closely related P. guatemalensis and P. oleifolius (Podocarpaceae) by sequencing 255 samples of the psbA-trnH and trnL-F intergenic spacer regions across the species ranges. We conducted phylogenetic, population and spatial genetic analyses as well as divergence time estimation and ecological niche modelling (ENM) to test the generality of demographic and genetic scenarios for cloud forest-adapted species. The results revealed genetic differentiation among species, with some individuals of P. oleifolius and P. guatemalensis placed in the P. matudae group and some P. oleifolius in the P. guatemalensis group. Predictions of ENMs under past climatic conditions and a strong signal of spatial expansion suggest that the highland P. matudae and P. oleifolius populations experienced expansions into lower elevation during the Last Glacial Maximum (LGM). Contrary to predictions by the two precipitation models and elevational ups and downs for cloud forest taxa during the LGM, genetic differentiation and predicted distribution of suitable habitat support the hypotheses that P. matudae and P. oleifolius remained in situ during the LGM primarily within the current fragmented distribution of the cloud forest and spread into the lowlands during the LGM, whereas the distribution of suitable habitat for P. guatemalensis had no major changes upwards from the Last Inter Glacial (LIG) to current conditions.

Origin and evolution of fleshy fruit in woody bamboos.

Several hypotheses have been suggested to explain the origin of fleshy fruit in monocots. One is that they originated in the understory of tropical regions and another is that fleshy fruit originated in tropical rainforests where high year-round rainfall implies that seasonality is not a limiting factor. Here we identify the time of origin and ecological preferences of woody bamboos to understand the evolution of the fleshy fruit known as the bacoid caryopsis. Bayesian Inference, Maximum Likelihood and molecular dating analyses were run based on eight plastid and two nuclear regions for 68 bamboo species. Climate data and soil parameters were gathered for 464 localities for these species. The ancestral type of caryopsis was reconstructed by parsimony. According to these analyses the bacoid caryopsis may have evolved independently seven times from the Late Miocene to the Early Pliocene and Mid-Pliocene to Mid-Pleistocene via convergent evolution. Our results suggest that in bamboos neither current climatic variables nor soil parameters were significantly correlated with the appearance of this type of fruit, nor do they have a phylogenetic signal. It is remarkable, however, that the first appearance of the bacoid caryopsis in bamboos might be associated with historical preferences for warmer and wetter climate during the Miocene. Further research is needed to identify whether other factors, such as vivipary or dispersal by small animals, rather than climate, could be responsible for the evolution of this trait in woody bamboos.

Spatio-temporal evolution of Leucophyllum pringlei and allies (Scrophulariaceae): a group endemic to North American xeric regions.

The taxa of the Leucophyllum pringlei clade were used to understand the influence of the Neogene orogenesis and the Quaternary climate cycles on the diversification of the flora of the xeric regions of North America. This clade includes the five southernmost species of the genus: L. ambiguum, L. flyrii, L. pruinosum and L. ultramonticola, which are distributed throughout the Chihuahuan Desert north of the Trans-Mexican Volcanic Belt, and L. pringlei in the region of Tehuacán-Cuicatlán south of this mountain range. Here we test whether these species diverged during the pluvial periods of the Pleistocene, and whether L. pringlei diverged earlier from the other species during the uplift of the Trans-Mexican Volcanic Belt. Using three plastid regions (psbA-trnH, psbK-psbI, trnL-F) and a nuclear (ITS) marker, phylogenetic analyses were carried out, along with a reconstruction of their ancestral area. Trees retrieved the five species in a monophyletic group with the most recent common ancestor distributed in the Sinaloan dry forest during the Late Miocene (8.08Ma), from where it dispersed to the Chihuahuan Desert during the Late Miocene (6.35Ma). The secondary uplift of the Sierra Madre Occidental during the Late Miocene to Early Pliocene influenced a vicariance event. Divergence between L. pringlei and the species from north of the Trans-Mexican Volcanic Belt occurred during the second volcanic episode in the Late Miocene (7.5-3Ma). The most recent common ancestor of L. ambiguum, L. pruinosum and L. ultramonticola was widely distributed in the southern part of the Chihuahuan Desert during the Early to Late Pliocene (3.50Ma). The diversification of these three species occurred in the Middle Pleistocene (0.9Ma) during the pluvial and inter-pluvial cycles.

Origin and diversification of the Milla Clade (Brodiaeoideae, Asparagaceae): a Neotropical group of six geophytic genera.

The Milla clade currently comprises six genera of geophytic plants distributed from Arizona to Guatemala. Three genera (Behria, Jaimehintonia and Petronymphe) are monotypic while the remaining genera (Bessera, Dandya and Milla) contain from two to ten (Milla) species. Parsimony, Maximum Likelihood and Bayesian Inference analyses were conducted with plastid and nuclear DNA sequences from a total of 181 plants belonging to 15 species in all six genera. Molecular dating was performed under a relaxed clock model. We examined the phylogenetic relationships of the genera and species, estimated origin-divergence times for the clade and genera and determined the ancestral distribution area of the clade by optimizing ancestral areas given current biogeographic distributions. The phylogenetic results suggest that final decisions on limits of the six genera in the Milla clade will have to be established until further taxonomic work is completed for Milla, in particular for the group of populations included under the name M. biflora. The later genus is rendered polyphyletic by other genera of the family. The origin of the Milla clade is estimated at 15.8Ma. Ancestral area of the clade most likely was located in the California Floristic Province and dispersal occurred most likely to the Chihuahuan-Coahuila Plateaus and the Trans-Mexican Volcanic Belt and from there to Baja California and the Sierra Madre del Sur. Two hypotheses that need further testing are proposed to explain complex relationships of genera and polyphyly of Milla, one in relation to fragmentation of populations and pollinator shifts and another suggesting that populations remained in refugia in the Trans-Mexican Volcanic Belt.

Stem transcriptome screen for selection in wild and cultivated pitahaya (Selenicereus undatus): an epiphytic cactus with edible fruit.

Dragon fruit, pitahaya or pitaya are common names for the species in the Hylocereus group of Selenicereus that produce edible fruit. These Neotropical epiphytic cacti are considered promising underutilized crops and are currently cultivated around the world. The most important species, S. undatus, has been managed in the Maya domain for centuries and is the focus of this article. Transcriptome profiles from stems of wild and cultivated plants of this species were compared. We hypothesized that differences in transcriptomic signatures could be associated with genes related to drought stress. De novo transcriptome assembly and the analysis of differentially expressed genes (DEGs) allowed us to identify a total of 9,203 DEGs in the Hunucmá cultivar relative of wild Mozomboa plants. Of these, 4,883 represent up-regulated genes and 4,320, down-regulated genes. Additionally, 6,568 DEGs were identified from a comparison between the Umán cultivar and wild plants, revealing 3,286 up-regulated and 3,282 down-regulated genes. Approximately half of the DEGs are shared by the two cultivated plants. Differences between the two cultivars that were collected in the same region could be the result of differences in management. Metabolism was the most representative functional category in both cultivars. The up-regulated genes of both cultivars formed a network related to the hormone-mediated signaling pathway that includes cellular responses to auxin stimulus and to hormone stimulus. These cellular reactions have been documented in several cultivated plants in which drought-tolerant cultivars modify auxin transport and ethylene signaling, resulting in a better redistribution of assimilates.

Wild and cultivated comestible plant species in the Gulf of Mexico: phylogenetic patterns and convergence of type of use.

Cross-cultural research on edible plants might include ecological and evolutionary perspectives to understand processes behind species selection and management. With a database of approximately 500 comestible plants of the Province of the Gulf of Mexico in Mesoamerica, phylogenetic analyses are conducted to identify convergence and phylogenetic signal of type of use and significant clustering in the resulting phylogenetic trees. Analyses considered type of management (wild/managed vs. cultivated), type of use (edible, condiment, for wrapping food) and organ utilized. Elevated phylogenetic diversity and signal are expected for wild comestible taxa, indicating that people are using lineages across the angiosperm tree for food, resulting in broadness in diet and use of their regional resources. Main results are: (i) condiment species were identified in groups with an elevated phylogenetic signal; (ii) hot nodes for lineages utilized for wrapping food were found in many monocot groups as well as in epiphytes of cloud forests with leathery leaves; (iii) edible taxa were identified with the highest significant clustering restricted to certain branches in the phylogeny; (iv) wild and cultivated edible plants belong to identical lineages with replacement of species, implying that same plant groups known for their comestible benefits are substituted by species distributed in the Province and (v) wild versus cultivated lineages for condiment are different. Most food species in the Province belong to four families, namely Fabaceae, Cactaceae, Solanaceae and Asparagaceae. Analyses discovered underutilized wild species in identical clades to managed/cultivated taxa that can be studied further to identify cultivation practices. Results suggest that people are utilizing different lineages in the angiosperm tree available locally, for particular uses, like condiment or for wrapping food. Evidence can be used to study further undervalued edible species closely related to the most common food taxa as well as for bioprospection of their nutritional content.

Diversification in North American arid lands: niche conservatism, divergence and expansion of habitat explain speciation in the genus Ephedra.

A lineage of 12 arid land shrubby species in the gymnosperm genus Ephedra (Gnetales) from North America is used to evaluate the influence of climate on speciation. With a long evolutionary history, and a well documented fossil record this lineage is an ideal model for understanding the process of speciation under a niche conservatism scenario. Using seven DNA molecular markers, Bayesian inference is carried out to uncover sister species and to estimate time of divergence of the lineages. Ecological niche models are generated for four parapatric and sympatric sister species and two analyses of niche evolution are performed, one based on ecological niche models and another using raw data and multivariate analysis. As previous analyses suggest, the diversification of North America Ephedra species may be the result of a recent secondary radiation. Both parapatric and sympatric species diverged mostly in a scenario of climatic niche conservatism. However, we also found strong evidence for niche divergence for one of the sister species pairs (E. californica-E. trifurca). Moreover, the multivariate analysis found environmental differences for some variables between sister species. The estimated divergence time of three pairs of sister species distributed in southwestern North America (E. cutleri-E. aspera, E. californica-E. trifurca and E. torreyana-E. viridis) is inferred to have occurred in the Late Miocene to Pliocene and for the sister species pair E. antisyphilitica-E. coryi distributed in the southern United States and northeastern Mexico, it was inferred from the Pliocene to Pleistocene. The orogenetic and climatic changes documented for these regions related to expansion of arid lands, may have contributed to the diversification in North American Ephedra, rather than adaptations to new climatic conditions.

Phylogenetic endemism of the orchids of Megamexico reveals complementary areas for conservation.

Orchid diversity provides a unique opportunity to further our understanding of biotic and abiotic factors linked to patterns of richness, endemism, and phylogenetic endemism in many regions. However, orchid diversity is consistently threatened by illegal trade and habitat transformation. Here, we identified areas critical for orchid conservation in the biogeographic province of Megamexico. For this purpose, we evaluated orchid endemism, phylogenetic diversity, and phylogenetic endemism within Megamexico and characterized orchid life forms. Our results indicate that the majority of the regions with the highest estimates of endemism and phylogenetic endemism are in southern Mexico and northern Central America, mostly located on the Pacific side of Megamexico. Among the most important orchid lineages, several belong to epiphytic lineages such as Pleurothallidinae, Laeliinae and Oncidiinae. We also found that species from diverse and distantly related lineages converge in montane forests where suitable substrates for epiphytes abound. Furthermore, the southernmost areas of phylogenetic diversity and endemism of Megamexico are in unprotected areas. Thus, we conclude that the most critical areas for orchid conservation in Megamexico are located in southern Mexico and northern Central America. We recommend that these areas should be given priority by the Mexican system of natural protected areas as complementary conservation areas.

Pollen morphology of the genera Hidalgoa and Dahlia (Coreopsideae, Asteraceae): implications for taxonomy.

Hidalgoa and Dahlia are two closely related genera in Asteraceae, tribe Coreopsideae whose limits need to be clarified. Pollen morphology has been useful for delimitation at the genus level in this family. To better define these genera, the morphology of pollen grains was observed and measured using light and scanning electron microscopy. The pollen grains of 25 species of Dahlia and Hidalgoa were acetolyzed and analyzed. Pollen is tricorporate in most of the species studied, although in a few species in Dahlia, grains were found to be hexacolporate. The most outstanding differentiating characters among species of Dahlia and Hidalgoa are colpus length (greater in Hidalgoa) and shape of spines (conical in Hidalgoa). In addition, lalongate ora are larger in Hidalgoa than in Dahlia. A PCA analysis of thirteen pollen characters, identified species of Hidalgoa in a discrete group and Dahliacuspidata as an outlier. These distinctive attributes in pollen morphology support the idea that pollen morphology is useful for delimitation at the generic level in the Dahlia clade. Further evidence from other sources, genetic or anatomical, might contribute to demarcating Dahlia and Hidalgoa, and provide insight into the family's evolutionary history.

Edible native plants of the Gulf of Mexico Province.

Background: Currently, at the global level, human food is mainly based on a few crops extensively cultivated as monocultures. Climate change, changes in land to agriculture and cattle raising, as well as the scarcity of water all affect and reduce the possibility of cultivating alternative crops. One way to face this global problem is to promote the knowledge, production and consumption of native food species on a regional scale.For this study, two databases were constructed for the Gulf of Mexico Province: 1) edible plant species with their corresponding common name, category of use, plant organ(s) utilised as food and type of management; 2) distribution records of these edible species. These species, in addition to being part of the biological diversity of Mexico are of high nutritional, cultural and gastronomical value and have been present in the diet of the inhabitants and ethnic groups in the region since pre-Hispanic times. New information: This study presents the native edible plants of the Gulf of Mexico Province, an area inhabited by 15 ethnic groups. The main novelty of this contribution is the edible plant species database, which includes the records of 482 species that belong to 101 families and 268 genera. We also present information rarely reported in an ethnobotanical inventory: 1) category of food use, 2) category of plant organ used, 3) common name, 4) type of management and 5) the georeferenced distribution of species occurrence in the Gulf of Mexico Province.

Transcriptional Basis for Haustorium Formation and Host Establishment in Hemiparasitic Psittacanthus schiedeanus Mistletoes.

The mistletoe Psittacanthus schiedeanus, a keystone species in interaction networks between plants, pollinators, and seed dispersers, infects a wide range of native and non-native tree species of commercial interest. Here, using RNA-seq methodology we assembled the whole circularized quadripartite structure of P. schiedeanus chloroplast genome and described changes in the gene expression of the nuclear genomes across time of experimentally inoculated seeds. Of the 140,467 assembled and annotated uniGenes, 2,000 were identified as differentially expressed (DEGs) and were classified in six distinct clusters according to their expression profiles. DEGs were also classified in enriched functional categories related to synthesis, signaling, homoeostasis, and response to auxin and jasmonic acid. Since many orthologs are involved in lateral or adventitious root formation in other plant species, we propose that in P. schiedeanus (and perhaps in other rootless mistletoe species), these genes participate in haustorium formation by complex regulatory networks here described. Lastly, and according to the structural similarities of P. schiedeanus enzymes with those that are involved in host cell wall degradation in fungi, we suggest that a similar enzymatic arsenal is secreted extracellularly and used by mistletoes species to easily parasitize and break through tissues of the host.

A character-based approach in the Mexican cycads supports diverse multigene combinations for DNA barcoding.

A DNA barcoding study was conducted to determine the optimal combination of loci needed for successful species-level molecular identification in three extant cycad genera-Ceratozamia, Dioon, and Zamia-that occur in Mexico. Based on conclusions of a previous multigene study in representative species of all genera in the Cycadales, we tested the DNA barcoding performance of seven chloroplast coding (matK, rpoB, rpoC1, and rbcL) and non-coding (atpF/H, psbK/I, and trnH-psbA) regions, plus sequences of the nuclear internal transcribed spacer. We analysed data under the assumptions of the "character attributes organization system" (CAOS), a character-based approach in which species are identified through the presence of 'DNA diagnostics'. In Ceratozamia, four chloroplast regions and one nuclear region were needed to achieve > 70% unique species identification. In contrast, the two-gene combination atpF/H + psbK/I and the four-gene combination atpF/H + psbK/I + rpoC1 + ITS2 were needed to reach 79% and 75% unique species identification in Dioon and Zamia, respectively. The combinations atpF/H + psbK/I and atpF/H + psbK/I + rpoC1 + ITS2 include loci previously considered by the international DNA barcoding community. However, none of the three combinations of potential DNA barcoding loci found to be optimal with a character-based approach in the Mexican cycads coincides with the 'core barcode' of chloroplast markers (matK + rbcL) recently proposed for universal use in the plant kingdom.

Delimiting species boundaries within the Neotropical bamboo Otatea (Poaceae: Bambusoideae) using molecular, morphological and ecological data.

Species delimitation is a task that has engaged taxonomists for more than two centuries. Recently, it has been demonstrated that molecular data and ecological niche modeling are useful in species delimitation. In this paper multiple data sets (molecular, morphological, ecological) were utilized to set limits for the species belonging to the Neotropical bamboo Otatea, because there is disagreement about species circumscriptions and also because the genus has an interesting distribution, with most of its populations in Mexico and a single disjunct population in Colombia. Molecular and morphological phylogenetic analyses recovered trees with conflicting topologies. Tree-based morphological and character-based analyses recognized the same entities. Ecological niche models and PCA/MANOVAS agreed with the recognition of the same entities that resulted from the morphological analyses. Morphological analyses retrieved clades supported by diagnostic characters and coherent geographical distributions. Based on these results seven entities should be recognized in Otatea, instead of the three previously described species.

Phylogenetic analyses of nucleotide sequences confirm a unique plant intercontinental disjunction between tropical Africa, the Caribbean, and the Hawaiian Islands.

Phylogenetic analyses of nucleotide sequences of the internal transcribed spacers and 5.8 regions of the nuclear ribosomal DNA and of the trnH-psbA spacer of the chloroplast genome confirm that the three taxa of the Jacquemontia ovalifolia (Choicy) Hallier f. complex (Convolvulaceae) form a monophyletic group. Levels of nucleotide divergence and morphological differentiation among these taxa support the view that each should be recognized as distinct species. These three species display unique intercontinental disjunction, with one species endemic to Hawaii (Jacquemontia sandwicensis A. Gray.), another restricted to eastern Mexico and the Antilles [Jacquemontia obcordata (Millspaugh) House], and the third confined to East and West Africa (J. ovalifolia). The Caribbean and Hawaiian species are sister taxa and are another example of a biogeographical link between the Caribbean Basin and Polynesia. We provide a brief conservation review of the three taxa based on our collective field work and investigations; it is apparent that J. obcordata is highly threatened and declining in the Caribbean.

Molecular phylogeny of the Acre clade (Crassulaceae): dealing with the lack of definitions for Echeveria and Sedum.

The phylogenetic relationships within many clades of the Crassulaceae are still uncertain, therefore in this study attention was focused on the "Acre clade", a group comprised of approximately 526 species in eight genera that include many Asian and Mediterranean species of Sedum and the majority of the American genera (Echeveria, Graptopetalum, Lenophyllum, Pachyphytum, Villadia, and Thompsonella). Parsimony and Bayesian analyses were conducted with 133 species based on nuclear (ETS, ITS) and chloroplast DNA regions (rpS16, matK). Our analyses retrieved four major clades within the Acre clade. Two of these were in a grade and corresponded to Asian species of Sedum, the rest corresponded to a European-Macaronesian group and to an American group. The American group included all taxa that were formerly placed in the Echeverioideae and the majority of the American Sedoideae. Our analyses support the monophyly of three genera--Lenophyllum, Thompsonella, and Pachyphytum; however, the relationships among Echeveria, Sedum and the various segregates of Sedum are largely unresolved. Our analyses represents the first broad phylogenetic framework for Acre clade, but further studies are necessary on the groups poorly represented here, such as the European and Asian species of Sedum and the Central and South American species of Echeveria.

Systematic relevance of pollen morphology in tribe Hylocereeae (Cactaceae).

Hylocereeae is one of the nine tribes in the subfamily Cactoideae (Cactaceae), for which the limits and recognition of genera have been controversial. Essentially, this group comprises epiphytic and hemiepiphytic genera with stems modified as climbing structures. The aim of this paper is to examine pollen attributes in representative species of genera of Hylocereeae, focusing on Selenicereus whose current circumscription comprises Hylocereus and three Weberocereus species, to find whether significant potentially apomorphic and/or autapomorphic systematic characters can be discovered. Utilizing SEM and light microscopy, 25 pollen characters were observed and measured. Tribe Hylocereeae is stenopalynous, with pollen grains isopolar and radially symmetrical monads, mostly tricolpate, except in Kimnachia, Pseudoripsalis and Weberocereus, whose pollen grains are pantocolpate. Seven attributes (five qualitative and two continuous) exhibited useful variation and were coded. The character of brevicolpate pollen grains was shared by Kimnachia ramulosa and Pseudorhipsalis amazonica. Convex quadrangular outline in the polar view was shared by Weberocereus tunilla and S. glaber. The absence of spinules on the exine was shared by S. minutiflorus and S. stenopterus. The largest pollen grain, found in Selenicereus megalanthus, might be correlated with polyploidy. Selenicereus is the taxon with the highest variation in pollen attributes, including species with an exine with or without spinules and variable polar area index and shape (subprolate or oblate-spheroidal).

Climate change and conservation in a warm North American desert: effect in shrubby plants.

BACKGROUND: Deserts are biologically rich habitats with a vast array of animals and plants adapted to xeric conditions, and most deserts are among the planet's last remaining areas of total wilderness. Among North American deserts, the Chihuahuan Desert has the highest levels of diversity and endemism. To understand the effect of future climate change on plants distributed in this arid land and propose effective conservation planning, we focused on five endemic shrubby species that characterize the Chihuahuan Desert and used an integrative approach. METHODS: Ecological niche-based modeling, spatial genetics and ecological resistance analyses were carried out to identify the effect of global warming on the studied five shrubby species. Key areas that need to be preserved were identified taking into account the existing protected areas within the Chihuahuan Desert. RESULTS: The extent of future distribution will vary among these species, and on average expansion will occur in the western part of the Chihuahuan Desert. For most species low environmental resistance to gene flow was predicted, while higher future resistance was predicted for one species that would lead to increased population isolation. The highest haplotype diversity was identified in three hotspots. Based on future suitability of habitat and in the haplotype diversity we suggest preserving two hotspots of genetic diversity in the Sierra Madre Oriental, located in areas without protection. The third hotspot was detected in the well preserved Tehuacán-Cuicatlán Man and Biosphere Reserve. CONCLUSION: Global climate change will have an effect in arid adapted plants, favoring expansion in the western of the Chihuahuan Desert however negatively affecting others with high ecological resistance disrupting gene flow. Two hotspots of genetic diversity in the Sierra Madre Oriental should be protected.

Anther development in tribe Epidendreae: orchids with contrasting pollination syndromes.

BACKGROUND: Epidendreae is one of the most diverse tribes among the orchids with remarkable variation in life form, floral morphology and pollination syndromes. Its circumscription was recently revised and subtribes Agrostophyllinae and Calypsoinae were transferred into this tribe. One of the principal floral characters utilized in classification of orchids is the incumbency or bending of the column. This study records and compares late stages of anther, column and lip development, and discusses anther characters in fifteen representative taxa of five of the six subtribes in Epidendreae with respect to classification and pollination biology. METHODS: A series of late floral stages were sampled and fixed for examination under scanning electron microscope. RESULTS: Anther incumbency or bending in this group varies from 90° to almost 180°. Incumbency in the late stages of development is reached in Bletiinae, Ponerinae, Pleurothallidinae and Laeliinae whereas incumbency is reached early in its development in Corallorhiza and Govenia of Calypsoinae. DISCUSSION: Our observations indicate that the position of Chysis in subtribe Bletiinae needs revision based on differences in a number floral, and in particular of anther characters; and that Coelia only shares the early anther incumbency with Calypsoinae members, but not the rest of floral and anther characters. Anatomical characters such as crystals around the actinocytic stomata on the anther cap and sugar crystals in Laeliinae; lack of rostellum in Bletiinae; coalescent anther with the column, lack of trichomes and papillae on lip keels, and underdeveloped rostellum in Chysis; a mechanism by which the anther cap comes off (it is joined with the grooved lip by a claw) in Isochilus are all related to pollination syndromes and reproductive biology.