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Background and Aims: Climate change is a global phenomenon affecting species, which in arid regions will translate into more frequent and intense periods of drought. The Sonoran Desert is becoming hotter and drier, and many organisms are rapidly changing in abundance and distribution. These population attributes depend directly on the dynamics of the population, which in turn depends on the vital rates of its individuals; yet few studies have documented the effects of climate change on the population dynamics of keystone species such as the saguaro cactus (Carnegiea gigantea). Although saguaros have traits that enable them to withstand present environmental conditions, climate change could make them vulnerable if forced beyond their tolerance limits., Methods: We evaluated the effect of climate change on 13 saguaro populations spanning most of the species' distribution range. Using field data from 2014 to 2016, we built an integral projection model (IPM) describing the environmentally explicit dynamics of the populations. We used this IPM, along with projections of two climate change scenarios and one no-change scenario, to predict population sizes (N) and growth rates (λ) from 2017 to 2099 and compared these scenarios to demonstrate the effect of climate change on the future of saguaro cactuses., Key Results: We found that all populations will decline, mainly due to future increases in drought, mostly hindering recruitment. However, the decline will be different across populations, since those located near the coast will be affected by harsher drought events than those located further inland., Conclusions: Our study demonstrates that climate change and its associated increase in drought pose a significant threat to the saguaro cactus populations in the Sonoran Desert. Our findings indicate that the recruitment of saguaros, vital for establishing new individuals, is particularly vulnerable to intensifying drought conditions. Importantly, regional climate trends will have different impacts on saguaro populations across their distribution range., (© The Author(s) 2024. Published by Oxford University Press on behalf of the Annals of Botany Company.)

Background: Gymnocarpos przewalskii Bunge ex Maxim. (G. przewalskii) is an endangered xerophytic shrub that plays a crucial role as a source of forage in the Alxa Desert. However, there is limited understanding regarding the forage quality of G. przewalskii and its response to drought. This study aimed to evaluate the forage quality of G. przewalskii and investigate the physiological and transcriptomic changes in G. przewalskii response to drought stress., Results: The ash, fat, crude protein, lignin, crude fiber, acid detergent fiber, and neutral detergent fiber contents in G. przewalskii twigs were 10.61%, 1.85%, 5.68%, 7.08%, 21.23%, 42.16%, and 58.42%, respectively. In contrast, these ingredients in its leaves were 20.39%, 0.92%, 11.96%, 2.40%, 17.51%, 14.29% and 20.26%, respectively. Osmotic stress led to a reduction in chlorophyll levels and an increase in malondialdehyde content. Levels of hydrogen peroxide and oxygen free radicals remained relatively stable under osmotic stress. The proline content, SOD and CAT activities, and ·OH scavenging capacity were enhanced in G. przewalskii under osmotic stress. RNA-sequencing of G. przewalskii generated 44.51 Gb clean reads, which were assembled into 102,191 Unigenes and 30,809 Unigenes were successfully annotated. Comparative analysis identified 3,015 differentially expressed genes under osmotic stress. There were 2,134 and 1,739 DEGs enriched in 47 GO secondary categories and 129 KEGG pathways, respectively. 2 up-regulated DEGs were annotated to P5CS, a key enzyme in the biosynthesis of proline. 32 DEGs were annotated to various antioxidases and antioxidants. 81 DEGs were annotated to 8 plant hormone signaling pathways, in which the auxin and ABA signaling pathways exhibited dominant enrichment. 150 DEGs were annotated to 35 transcription factor families with the abundant enrichment of TF families containing WRKY, bZIP, ERF, bHLH, MYB, and NAC., Conclusions: High forage quality and drought stress tolerance were observed in G. przewalskii. In response to drought stress, G. przewalskii orchestrates reactive oxygen species scavenging, proline biosynthesis, and other intricate physiological processes, with substantial contributions from plant hormones and transcription factors. This study provides new insights into the forage quality and the mechanisms involved in drought adaptation of G. przewalskii, offering a foundation for its conservation and sustainable utilization., Competing Interests: Declarations. Ethics approval and consent to participate: The permission for collecting twigs, leaves and seeds of Gymnocarpos przewalskii in Yabalai Mountain were obtained from the local government. Consent for publication: Not applicable. Competing interests: The authors declare no competing interests., (© 2025. The Author(s).)

Night-blooming cacti, primarily pollinated by bats and hawkmoths, also attract beetles seeking food and safe shelter for mating and brooding their offspring. The influence of flower density on beetle visitation rates remains unclear, with responses varying by species and environmental factors. In the Caatinga Seasonally Dry Tropical Forest, we studied the flower occupancy distribution of two beetle species, Cyclocephala paraguayensis and Nitops aff. pilosocerei, in Pilosocereus pachycladus cacti. Our findings indicate that both beetle species act as commensals with minimal impact on effective pollination. They forage for nectar and pollen without causing damage to pistils or ovaries. N. aff. pilosocerei was more abundant than C. paraguayensis, and their distributions significantly differed, with N. aff. pilosocerei displaying a more uniform spread. Instances of both species occupying the same flower were more frequent than exclusive occupation. Nitops aff. pilosocerei abundance exhibited spatial autocorrelation. Flower height and beetle species influenced the total number of beetles within flowers. Future studies should explore the impact of cactus flower distribution on beetle abundance with other species, conduct selective pollination experiments to determine their role as pollinators, and investigate how flower-beetle interaction systems are affected by flower and individual distribution in processes like florivory and pollination.

Pitaya, a desert plant, has an underexplored flowering mechanism due to a lack of functional validation assays. This study reveals that the transition from vegetative to generative growth in pitaya is regulated by significant metabolic shift, underscoring the importance of understanding and address the challenging issue pitaya's phase change. Lateral buds from 6-years-old 'Guanhuahong' pitaya (Hylocereus monacanthus) plants were collected on April 8th, 18th, and 28th 2023, representing early, middle, and late stages of phase transition, respectively. Results showed diminished nitrogen levels concurrent with increased carbon levels and carbon-to-nitrogen (C/N) ratios during pitaya phase transition. Transcriptomic analysis identified batches of differentially expressed genes (DEGs) involved in downregulating nitrogen metabolism and upregulating carbon metabolism. These batches of genes play a central role in the metabolic shifts that predominantly regulate the transition to the generative phase in pitaya. This study unveils the intricate regulatory network involving 6 sugar synthesis and transport, 11 photoperiod (e.g., PHY, CRY, PIF) and 6 vernalization (e.g., VIN3) pathways, alongside 11 structural flowering genes (FCA, FLK, LFY, AGL) out of a vast array of potential candidates in pitaya phase change. These findings provide insights into the metabolic pathways involved in pitaya's phase transition, offering a theoretical framework for managing flowering, guiding breeding strategies to optimize flowering timing and improve crop yields under varied nitrogen conditions., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2024 Elsevier B.V. All rights reserved.)

In this era of a global biodiversity crisis, vascular plants are facing unprecedented extinction rates. We conducted an assessment of the extinction risk of 32 species and 7 subspecies of Copiapoa, a genus endemic to Chile's fog-dependent coastal Atacama Desert. We applied the International Union for Conservation of Nature Red List Categories and Criteria enhanced by expert insights and knowledge. Our primary aim was to analyze the impact of trade and poaching on their extinction risk. We employed machine learning models, including multinomial logistic regression (MLR), decision tree (DT), and random forest (RF), to analyze the relationships between conservation status and various factors. These factors encompassed trade and poaching activities, landscape condition, human footprint, monthly cloud frequency, and biological traits such as evolutionary distinctiveness and maximum diameter. Seven taxa had an area of occupancy (AOO) of <10 km 2 , 10 additional taxa had an AOO of <20 km 2 , and 16 taxa had an AOO of ≤100 km 2 . This reassessment exposed a critical level of extinction risk for the genus; 92% of the taxa were classified as threatened, 41% as critically endangered, 41% as endangered, and 10% as vulnerable. MLR, DT, and RF exhibited accuracies of 0.784, 0.730, and 0.598, respectively, and identified trade and poaching pressure and landscape condition as the primary drivers of extinction risk. Our assessment of Copiapoa showed trade, poaching, habitat degradation, and their synergic impacts as the main drivers of the genus' extinction risk. Our results highlight the urgent need for nations to develop and enforce strategies to monitor and control trade and poaching pressure because these factors are crucial for the long-term persistence of desert plants., (© 2024 The Author(s). Conservation Biology published by Wiley Periodicals LLC on behalf of Society for Conservation Biology.)

Reproductive isolation is one of the mechanisms of speciation. The two currently accepted subspecies of Parodia haselbergii (P. haselbergii subsp. haselbergii and P. haselbergii subsp. graessneri) were studied regarding flower traits, phenology, breeding systems and pollination. In addition, a principal component analysis with 18 floral characters and germination tests under controlled conditions were performed for both taxa. Pollination was studied in the field, in two localities of Southern Brazil. Pollinators were recorded through photos and film. Breeding system experiments were performed by applying controlled pollinations to plants excluded from pollinators. Both taxa mostly differ in asynchronous flowering periods, floral traits (including floral part measurements and nectar concentration) and pollinators. The flowers of both subspecies are functionally protogynous and perform remarkably long lifespans (≥ 15 days), both traits being novelties for Cactaceae. Whereas the reddish flowers of P. haselbergii subsp. haselbergii (nectar concentration: ca. 18%) are pollinated by hummingbirds of Thalurania glaucopis, the greenish flowers of P. haselbergii subsp. graessneri (nectar concentration: ca. 29%) are pollinated by Augochlora bees (Halictidae). Both subspecies are self-compatible, yet pollinator-dependent. The principal component analysis evidenced that both subspecies are separated, regarding flower traits. The seeds of both subspecies performed differently in the germination tests, but the best results were recovered at 20 °C and germination considerably decreased around 30 °C. In conclusion, all these results support that both taxa are in reproductive isolation, and can be treated as different species., (© 2024. The Author(s) under exclusive licence to The Botanical Society of Japan.)

Key Message: Through a multidisciplinary study we found that Gymnocalycium bruchii, an endemic cactus from central Argentina, acts as a dioecious species, which is the first record in this genus. Cactaceae species are typically hermaphroditic; however, about 2% have other different reproductive systems. These non-hermaphroditic species may develop sexual dimorphism in flowers or other reproductive, vegetative or ecological traits, besides a specific breeding system and floral ontogeny. Therefore, multidisciplinary research is necessary to fully understand reproduction in those species. For this purpose, we studied Gymnocalicium bruchii, a globose cactus endemic to central Argentina that is presumably dioecious or gynodioecious. We made observations in two natural and two cultivated populations. We made morphological observations of plants and flowers, and performed quantitative analyses to determine the sex ratio, size of plants and flowers, flower production, fruiting, among other variables. We performed hand-pollination, self-fertilization and free-pollination tests to determine the breeding system. Finally, we studied the anatomy and ontogeny of the reproductive organs using permanent histological slides of flower morphs at different stages. Our results confirm that Gymnocalicium bruchii is a dioecious species. Female flowers have atrophied anthers and a functional gynoecium that produces fruits and seeds. Male flowers are bigger and have a functional androecium but a sterile gynoecium. In the cultivated population, the sex ratio was 1/1, whereas the number of male individuals was higher in both natural populations. Pollination tests corroborated dioecy. Ontogenetic studies revealed that in female flowers the anthers collapse before microspore maturation, while in male flowers the gynoecium shows normal development of the ovary, style, stigma, and ovules; however, the latter are never fertilized., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Key Message: Contrasting morphologies in Disocactus are the result of differential development of the vegetative and floral tissue where intercalary growth is involved, resulting in a complex structure, the floral axis. Species from the Cactaceae bear adaptations related with their growth in environments under hydric stress. These adaptations have translated into the reduction and modification of various structures such as leaves, stems, lateral branches, roots and the structuring of flowers in a so-called flower-shoot. While cacti flowers and fruits have a consistent structure with showy hermaphrodite or unisexual flowers that produce a fruit called cactidium, the developmental dynamics of vegetative and reproductive tissues comprising the reproductive unit have only been inferred through the analysis of pre-anthetic buds. Here we present a comparative analysis of two developmental series covering the early stages of flower formation and organ differentiation in Disocactus speciosus and Disocactus eichlamii, which have contrasting floral morphologies. We observe that within the areole, a shoot apical meristem commences to grow upward, producing lateral leaves with a spiral arrangement, rapidly transitioning to a floral meristem. The floral meristem produces tepal primordia and a staminal ring meristem from which numerous or few stamens develop in a centrifugal manner in D. speciosus and D. eichlamii, respectively. Also, the inferior ovary derives from the floral meristem flattening and an upward growth of the surrounding tissue of the underlying stem, producing the pericarpel. This structure is novel to cacti and lacks a clear anatomical delimitation with the carpel wall. Here, we present a first study that documents the early processes taking place during initial meristem determination related to pericarpel development and early floral organ formation in cacti until the establishment of mature floral organs., (© 2024. The Author(s).)

Main Conclusion: Natural selection influenced adaptive divergence between Cereus fernambucensis and Cereus insularis, revealing key genes governing abiotic stress responses and supporting neoteny in C. insularis. Uncovering the molecular mechanisms driving adaptive divergence in traits related to habitat adaptation remains a central challenge. In this study, we focused on the cactus clade, which includes Cereus sericifer F.Ritter, Cereus fernambucensis Lem., and Cereus insularis Hemsley. These allopatric species inhabit distinct relatively drier regions within the Brazilian Atlantic Forest, each facing unique abiotic conditions. We leveraged whole transcriptome data and abiotic variables datasets to explore lineage-specific and environment-specific adaptations in these species. Employing comparative phylogenetic methods, we identified genes under positive selection (PSG) and examined their association with non-synonymous genetic variants and abiotic attributes through a PhyloGWAS approach. Our analysis unveiled signatures of selection in all studied lineages, with C. fernambucensis northern populations and C. insularis showing the most PSGs. These PSGs predominantly govern abiotic stress regulation, encompassing heat tolerance, UV stress response, and soil salinity adaptation. Our exclusive observation of gene expression tied to early developmental stages in C. insularis supports the hypothesis of neoteny in this species. We also identified genes associated with abiotic variables in independent lineages, suggesting their role as environmental filters on genetic diversity. Overall, our findings suggest that natural selection played a pivotal role in the geographic range of these species in response to environmental and biogeographic transitions., (© 2024. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The colonization and expansion of any plant species into a novel environment depend on its structural and functional characteristics. Therefore, developing better control measures for any invasive plant species requires examining and understanding the mechanisms underlying its reproduction and adaptation to the environment it invades. Recently, a novel exotic species Cylindropuntia rosea (DC.) Backeb. has been identified in Baljurashi, Al-Baha province, in southwestern Saudi Arabia. Reports suggest that this species may become invasive with the current rate of habitat expansion in Baljurashi. Although C. rosea is an important invasive species, most of its morpho-anatomical and physiological characteristics have not been examined. Therefore, the aim of this study was to investigate the morpho-anatomical and related physiological adaptations of C. rosea in its new habitats in the southwestern highlands of Saudi Arabia. We observed that the species is well-equipped for invasion with traits to handle semi-arid conditions, including some morphological and anatomical features, CAM photosynthetic pathway, high growth rate, and highly effective defense mechanisms against herbivores and insects. These morpho-anatomical and physiological characteristics contribute to the high invasiveness of this species in Saudi Arabia.

Cactaceae (cacti), a New World plant family, is one of the most endangered groups of organisms on the planet. Conservation planning is uncertain as it is unclear whether climate and land-use change will positively or negatively impact global cactus diversity. On the one hand, a common perception is that future climates will be favourable to cacti as they have multiple adaptations and specialized physiologies and morphologies for increased heat and drought. On the other hand, the wide diversity of the more than 1,500 cactus species, many of which occur in more mesic and cooler ecosystems, questions the view that most cacti can tolerate warmer and drought conditions. Here we assess the hypothesis that cacti will benefit and expand in potential distribution in a warmer and more drought-prone world. We quantified exposure to climate change through range forecasts and associated diversity maps for 408 cactus species under three Representative Concentration Pathways (2.6, 4.5 and 8.5) for 2050 and 2070. Our analyses show that 60% of species will experience a reduction in favourable climate, with about a quarter of species exposed to environmental conditions outside of the current realized niche in over 25% of their current distribution. These results show low sensitivity to many uncertainties in forecasting, mostly deriving from dispersal ability and model complexity rather than climate scenarios. While current range size and the International Union for Conservation of Nature's Red List category were not statistically significant predictors of predicted future changes in suitable climate area, epiphytes had the greatest exposure to novel climates. Overall, the number of cactus species at risk is projected to increase sharply in the future, especially in current richness hotspots. Land-use change has previously been identified as the second-most-common driver of threat among cacti, affecting many of the ~31% of cacti that are currently threatened. Our results suggest that climate change will become a primary driver of cactus extinction risk with 60-90% of species assessed negatively impacted by climate change and/or other anthropogenic processes, depending on how these threat processes are distributed across cactus species., (© 2022. The Author(s), under exclusive licence to Springer Nature Limited.)

Background: In certain unisexual flowers, non-functional sexual organs remain vestigial and unisexuality can be overlooked leading to the ambiguous description of the sexual systems. Therefore, to accurately describe the sexual system, detailed morphological and developmental analyses along with experimental crosses must be performed. Cylindropuntia wolfii is a rare cactus endemic to the Sonoran Desert in southern California and northern Baja California that was described as gynodioecious by morphological analysis. The aims of our project include accurately identifying the sexual system of C. wolfii using histological and functional studies and characterizing the developmental mechanisms that underlie its floral development., Methods: Histological analyses were carried out on different stages of C. wolfii flowers and controlled crosses were performed in the field., Result: Our results identified C. wolfii to be functionally dioecious. The ovule and anther development differed between staminate and pistillate flowers. In vivo pollen germination tests showed that the pollen of staminate and pistillate flowers were viable and the stigma and style of both staminate and pistillate flowers were receptive. This suggests that there are no genetic or developmental barriers in the earlier stages of pollen recognition and pollen germination., Conclusions: Despite being functionally dioecious, we observed that functionally pistillate individuals produced fruits with a large number of aborted seeds. This implies that not only does this species have low reproductive success, but its small population sizes may lead to low genetic diversity., (© 2022. The Author(s).)

Background: A puzzle in evolution is the understanding of how the environment might drive subtle phenotypic variation, and whether this variation is adaptive. Under the neutral evolutionary theory, subtle phenotypes are almost neutral with little adaptive value. To test this idea, we studied the infraspecific variation in flower shape and color in Mammillaria haageana, a species with a wide geographical distribution and phenotypic variation, which populations are often recognized as infraspecific taxa., Results: We collected samples from wild populations, kept them in the greenhouse for at least one reproductive year, and collected newly formed flowers. Our first objective was to characterize tepal natural variation in M. haageana through geometric morphometric and multivariate pigmentation analyses. We used landmark-based morphometrics to quantify the trends of shape variation and tepal color-patterns in 20 M. haageana accessions, belonging to five subspecies, plus 8 M. albilanata accessions for comparison as the sister species. We obtained eight geometric morphometric traits for tepal shape and color-patterns. We found broad variation in these traits between accessions belonging to the same subspecies, without taxonomic congruence with those infraspecific units. Also the phenetic cluster analysis showed different grouping patterns among accessions. When we correlated these phenotypes to the environment, we also found that solar radiation might explain the variation in tepal shape and color, suggesting that subtle variation in flower phenotypes might be adaptive. Finally we present anatomical sections in M. haageana subsp. san-angelensis to propose some of the underlying tepal structural features that may give rise to tepal variation., Conclusions: Our geometric morphometric approach of flower shape and color allowed us to identify the main trends of variation in each accession and putative subspecies, but also allowed us to correlate these variation to the environment, and propose anatomical mechanisms underlying this diversity of flower phenotypes., (© 2022. The Author(s).)

When two or more parasitoid species, particularly candidates for biocontrol, share the same target in the same temporal window, a complex of behaviors can occur among them. We studied the type of interactions (competition and intraguild predation) that existed between the nymphal parasitoids Anagyrus cachamai and A. lapachosus (Hymenoptera: Encyrtidae), two candidate neoclassical biocontrol agents against the Puerto Rican cactus pest mealybug, Hypogeococcus sp. (Hemiptera: Pseudococcidae). The surrogate native congener host in Argentina, the cactus mealybug Hypogeococcus sp., was studied to predict which species should be released; in the case that both should be released, in which order, and their potential impact on host suppression. In the laboratory we conducted experiments where different densities of the host mealybug were exposed to naive females of A. cachamai and A. lapachosus sequentially in both directions. Experiments were analyzed by combining a series of competitive behavioral and functional response models. A fully Bayesian approach was used to select the best explaining models and calculate their parameters. Intraguild predation existed between A. cachamai, the species that had the greatest ability to exploit the resource, and A. lapachosus, the strongest species in the interference competition. The role that intraguild predation played in suppression of Hypogeococcus sp. indicated that a multiple release strategy for the two biocontrol agents would produce better control than a single release; as for the release order, A. lapachosus should be released first.

The Hylocereus species that are grown as exotic fruit crops are very often farmed under marginal agronomic conditions, which may include exposure to high temperatures. Here we present a pioneering investigation of grafting as an agro-technique to improve heat tolerance in Hylocereus. To this end, we studied the diploid species H. undatus, the tetraploid H. megalanthus and its di-haploid gamete-derived line 2719, and the interspecific-interploid tetraploid Z-10, all grafted onto H. undatus as the rootstock. Self-grafted, grafted and non-grafted plants were acclimated for one week (to obtain baseline values) and then exposed to heat stress (45/35 °C day/night) for three days, followed by a one-week recovery period under optimal temperatures (30/22 °C). A comparison of the physiological, biochemical and molecular performances of the grafted and self-grafted plants under heat stress and during the recovery period vs those of non-stressed plants (control; 30/22 °C) showed that the grafted and self-grafted plants performed better in most of the assessments: grafted and self-grafted plants recovered more rapidly from the heat stress and suffered far less stem damage. An unexpected - but important - finding that may have implications for other crop was that the self-grafted plants showed better performance than non-grafted plants throughout the trial. Our findings provide support for grafting as a strategy for coping with the stress induced by extremely high temperatures. This study thus paves the way for further investigations of grafting in Hylocereus as a valuable technique that will maintain crop productivity in the face of increasing worldwide temperatures., (Copyright © 2021 Elsevier Masson SAS. All rights reserved.)

High temperatures limit the successful cultivation of the Hylocereus species on a global basis. We aimed to investigate the degree of heat tolerance in three species, namely, the diploids Hylocereus undatus and H. monacanthus, and the tetraploid H. megalanthus, and nine of their interspecific-interploid hybrids. Rooted cuttings were exposed to heat stress (45/35 °C) or control conditions (25/20 °C) for eight days. Initially, the plants were screened for their tolerance to heat stress and ranked into four heat tolerance categories: good tolerance, moderate tolerance, low tolerance, or sensitive, according to the decrease in the maximum quantum efficiency of photosystem II (F v /F m ) and visual stem damage. The physiological and biochemical performances of the parental species and of three hybrids representing three different heat-tolerance categories were further analyzed in depth. H. megalanthus (classified as heat sensitive) showed a 65% decrease in F v /F m and severe visual stem damage, along with a marked reduction in total chlorophyll content, a large increase in malondialdehyde, and inhibition of catalase activity. H. undatus and H. monacanthus, (classified as low-tolerance species) exhibited slight stem "liquification." The good-tolerance hybrid Z-16 exhibited the best performance under heat stress (21% decrease in F v /F m ) and the absence of stem damage, coupled with a small decrease in total chlorophyll content, a slight increase in malondialdehyde, high antioxidant activity, and proline accumulation progressing with time. Our findings revealed that most of the hybrids performed better than their parental species, indicating that our breeding programs can provide Hylocereus cultivars suitable for cultivation in heat-challenging regions., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier Masson SAS. All rights reserved.)

The climate change projections for the Caatinga biome this century are for an increase in temperature and reduction in rainfall, leading to aridization and plant cover dominated by Cactaceae. The objective of this study was to model the potential distribution of Epiphyllum phyllanthus (L.) Haw., a cactus that is native to the Caatinga biome, considering two possible future climate scenarios, to assess this species' spatio-temporal response to these climate change, and thus to evaluate the need or not for conservation measures. For this purpose, we obtained biogeographic information on the target species from biodiversity databases, choosing nine environmental variables and applying the MaxEnt algorithm. We considered the time intervals 2041-2060 and 2061-2080, centered on 2050 and 2070, respectively, and the greenhouse gas scenarios RCP4.5 and 8.5. For all the scenarios considered, the models generated for 2050 and 2070 projected drastic contraction (greater than 80%) for the areas of potential occurrence of the species in relation to the present potential. The remaining areas were found to be concentrated in the northern portion of the biome, specifically in the northern part of the state of Ceará, which has particular characteristics.

The plant family Cactaceae is considered among the most threatened groups of organisms on the planet. The threatened status of the cacti family has created a renewed interest in the highly evolved physiological and morphological traits that underpin their persistence in some of the harshest subtropical environments in the Americas. Among the most important anatomical features of cacti is the modification of leaves into spines, and previous work has shown that the stable isotope chemistry of cacti spines records potential variations in stem water balance, stress, and Crassulacean acid metabolism (CAM). We review the opportunities, challenges, and pitfalls in measuring δ 13C, δ 2H, and δ 18O ratios captured in spine tissues that potentially reflect temporal and spatial patterns of stomatal conductance, internal to atmospheric CO2 partial pressures, and subsequent patterns of photosynthetic gas exchange. We then evaluate the challenges in stable isotope analysis in spine tissues related to variation in CAM expression, stem water compartmentalization, and spine whole-tissue composition among other factors. Finally, we describe how the analysis of all three isotopes can be used in combination to provide potentially robust analysis of photosynthetic function in cacti, and other succulent-stemmed taxa across broad spatio-temporal environmental gradients., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

To prevent plants from being damaged due to extreme temperature and sunlight, the pitaya orchards in Hainan Province, China, are increasingly adopting living and black fabric cloth mulching. In this study, an Illumina Hiseq sequencer was employed to compare the soil microbial communities of two pitaya orchards, one covered by living mulching (LM) and the other covered by black fabric cloth (FC). Bacterial abundance was higher in the LM orchard than in the FC orchard (1.19 × 10 4 versus 4.49 × 10 4  g -1 soil). In contrast, fungal abundance was higher in the FC orchard than in the LM orchard (2.71 × 10 6 versus 2.97 × 10 5  g -1 soil). We also found that the most dominant species in the FC orchard were from the genus Neoscytalidium, which included species that could cause infection in a large variety of plant hosts. However, the LM orchard mainly harbored useful fungal species, such as Trichoderma and Chaetomium. Soil nutrients were positively correlated in the FC orchard, which potentially indicated that the FC orchard could demonstrate better fertilizer utilization efficiency. However, the LM and FC strategies have both advantages and disadvantages with regards to the cultivation management of pitaya orchards.

MicroRNA396 ( miR396 ) is a conserved microRNA family that targets growth-regulating factors (GRFs), which play significant roles in plant growth and stress responses. Available evidence justifies the idea that miR396 -targeted GRFs have important functions in many plant species; however, no genome-wide analysis of the pitaya ( Hylocereus polyrhizus ) miR396 gene has yet been reported. Further, its biological functions remain elusive. To uncover the regulatory roles of miR396 and its targets, the hairpin sequence of pitaya miR396b and the open reading frame (ORF) of its target, HpGRF6 , were isolated from pitaya. Phylogenetic analysis showed that the precursor miR396b ( MIR396b ) gene of plants might be clustered into three major groups, and, generally, a more recent evolutionary relationship in the intra-family has been demonstrated. The sequence analysis indicated that the binding site of hpo-miR396b in HpGRF6 is located at the conserved motif which codes the conserved "RSRKPVE" amino acid in the Trp-Arg-Cys (WRC) region. In addition, degradome sequencing analysis confirmed that four GRFs ( GRF1 , c56908.graph&#95;c0; GRF4 , c52862.graph&#95;c0; GRF6 , c39378.graph&#95;c0 and GRF9 , c54658.graph&#95;c0) are hpo-miR396b targets that are regulated by specific cleavage at the binding site between the 10th and 11th nucleotides from the 5' terminus of hpo-miR396b . Furthermore, quantitative real-time polymerase chain reaction (qRT-PCR) analysis showed that hpo-miR396b is down-regulated when confronted with drought stress (15% polyethylene glycol, PEG), and its expression fluctuates under other abiotic stresses, i.e., low temperature (4 ± 1 °C), high temperature (42 ± 1 °C), NaCl (100 mM), and abscisic acid (ABA; 0.38 mM). Conversely, the expression of HpGRF6 showed the opposite trend to exposure to these abiotic stresses. Taken together, hpo-miR396b plays a regulatory role in the control of HpGRF6 , which might influence the abiotic stress response of pitaya. This is the first documentation of this role in pitaya and improves the understanding of the molecular mechanisms underlying the tolerance to drought stress in this fruit.

Frequent somatic variations exist in pitaya (Hylocereus undatus) plants grown under abiotic stress conditions. Long terminal repeat (LTR) retrotransposons can be activated under stressful conditions and play key roles in plant genetic variation and evolution. However, whether LTR retrotransposons promotes pitaya somatic variations by regulating abiotic stress responses is still uncertain. In this study, transcriptionally active LTR retrotransposons were identified in pitaya after exposure to a number of stress factors, including in vitro culturing, osmotic changes, extreme temperatures and hormone treatments. In total, 26 LTR retrotransposon reverse transcriptase (RT) cDNA sequences were isolated and identified as belonging to 9 Ty1-copia and 4 Ty3-gypsy families. Several RT cDNA sequences had differing similarity levels with RTs from pitaya genomic DNA and other plant species, and were differentially expressed in pitaya under various stress conditions. LTR retrotransposons accounted for at least 13.07% of the pitaya genome. HuTy1P4 had a high copy number and low expression level in young stems of pitaya, and its expression level increased after exposure to hormones and abiotic stresses, including in vitro culturing, osmotic changes, cold and heat. HuTy1P4 may have been subjected to diverse transposon events in 13 pitaya plantlets successively subcultured for four cycles. Thus, the expression levels of these retrotransposons in pitaya were associated with stress responses and may be involved in the occurrence of the somaclonal variation in pitaya., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

Premise of the Study: Phenology is the study of biological life cycle events, such as flowering and migration. Climate patterns can alter these life history events, having ecosystem-wide ramifications. For example, warmer springs are associated with earlier leaf-out for many species, impacting species interactions and growing-season carbon dynamics. While phenological research has been conducted extensively in temperate regions, relatively little is known about the phenological responses in arid and semi-arid regions., Methods: In this study we looked at the flowering phenology of a keystone species in the Sonoran Desert, the saguaro cactus (Carnegiea gigantea). The timing and abundance of flowering was observed on 151 individuals for 10 years at a site near Tucson, Arizona, USA. Using six phenological traits, we explored the relationship between saguaro size and flowering and the climatic drivers of flowering., Key Results: Our analyses demonstrated how the calculation of phenological traits at the individual versus the population level can yield differing responses to climate variability, suggesting that not all studies examining the same trait (e.g., first day of bloom) are directly comparable. We found that larger cacti began flowering earlier, flowered for longer, and produced more flowers. Warmer temperatures were correlated with advanced onset and higher bloom yields, while increased precipitation appeared to delay onset and reduce bloom yields., Conclusions: Given that climate models predict that the Southwestern USA will become increasingly warmer with more variable precipitation, saguaros may begin flowering earlier in the season and flower more intensely, which could impact pollen availability and population dynamics., (© 2019 Botanical Society of America.)

Background: Insights into the genetic capacities of species to adapt to future climate change can be gained by using comparative genomic and transcriptomic data to reconstruct the genetic changes associated with such adaptations in the past. Here we investigate the genetic changes associated with adaptation to arid environments, specifically climatic extremes and new cactus hosts, through such an analysis of five repleta group Drosophila species., Results: We find disproportionately high rates of gene gains in internal branches in the species' phylogeny where cactus use and subsequently cactus specialisation and high heat and desiccation tolerance evolved. The terminal branch leading to the most heat and desiccation resistant species, Drosophila aldrichi, also shows disproportionately high rates of both gene gains and positive selection. Several Gene Ontology terms related to metabolism were enriched in gene gain events in lineages where cactus use was evolving, while some regulatory and developmental genes were strongly selected in the Drosophila aldrichi branch. Transcriptomic analysis of flies subjected to sublethal heat shocks showed many more downregulation responses to the stress in a heat sensitive versus heat resistant species, confirming the existence of widespread regulatory as well as structural changes in the species' differing adaptations. Gene Ontology terms related to metabolism were enriched in the differentially expressed genes in the resistant species while terms related to stress response were over-represented in the sensitive one., Conclusion: Adaptations to new cactus hosts and hot desiccating environments were associated with periods of accelerated evolutionary change in diverse biochemistries. The hundreds of genes involved suggest adaptations of this sort would be difficult to achieve in the timeframes projected for anthropogenic climate change.

Host plant shifts in herbivorous insects often involve facing new environments that may speed up the evolution of oviposition behavior, performance-related traits, morphology, and, incidentally, reproductive isolation. In the genus Drosophila, cactophilic species of the repleta group include emblematic species in the study of the evolution of host plant utilization. The South American D. buzzatii and its sibling D. koepferae are a model system for the study of differential host plant use. Although these species exhibit a certain degree of niche overlap, the former breeds primarily on decaying cladodes of Opuntia cacti while D. koepferae main hosts are columnar cacti of the genus Trichocereus. Opuntia sulphurea and Trichocereus terscheckii are among the main hosts in nature. These cacti differ in ecological (spatial and temporal predictability) and chemical characteristics. Particularly relevant is the presence of toxic alkaloids in T. terscheckii. Studies of the effects of these cacti and alkaloids revealed the remarkable impact on oviposition behavior, viability, developmental time, wing morphology, mating success, and developmental stability in both species. Recent whole-genome expression studies showed that expression profiles are massively affected by the rearing cactus, and that the presence of alkaloids is the main factor modulating gene expression in D. buzzatii. Functional enrichment analysis indicated that differentially expressed genes are related to detoxification processes and stress response-though genes involved in development are an important part of the transcriptomic response. The implications of our studies in the evolution of host plant use in the repleta group are discussed.

Spines are common morphological features found in almost all major biological groups offering an opportunity to explore large-scale evolutionary convergence across disparate clades. As an example, opuntioid cacti have spines with barbed ornamentation that is remarkably similar in form and scale to that found on porcupine quills, suggesting specific biomechanical convergence across the animal and plant kingdoms. While the mechanics of porcupine quills as defensive mechanisms has been previously tested, the mechanics of cactus spines (which have evolved to fill a number of functions including defence, climbing and dispersal) has not been characterized. Here we study the puncturing and anchoring ability of six species of cactus, including both barbed and non-barbed spines. We found that barbed spines require less work to puncture a variety of targets than non-barbed spines. Barbed spines also require more work than non-barbed spines to withdraw from biological materials, owing to their barbs engaging with tissue fibres. These results closely match those found previously for barbed versus non-barbed porcupine quills, implying biomechanical convergence. The variation in performance of barbed versus non-barbed spines, as well as between barbed spines from different species, is probably tied to the diversity of ecological functions of cactus spines., (© 2018 The Author(s).)

There is controversy over whether cactus species form soil seed banks. Although it is commonly assumed that cacti do not form seed banks, very few studies have evaluated them. In this work, we analysed whether cactus species form soil seed banks, studying seed distribution, seed density and seed longevity in the Southern Central Andes. Soil samples were collected in two microhabitats (under nurse plants and in bare areas) at 12 selected sites. We determined seed presence-absence, density and distribution for 32 native cactus species. Seed longevity for six of these species was determined through a burial experiment. We recorded viable seeds for 62.5% of the 32 evaluated species, finding variation in seed density between microenvironments and among populations. In some species, the highest seed density was found under potential nurse plants. Seed germination and seed viability decreased with burial time, with seed longevity always being <24 months after burial. Our results show strong evidence that cactus species do form seed banks. Seed density can vary between microenvironments and among populations, suggesting that cactus-nurse plant associations can also be explained by differential seed dispersal and not only by differential establishment. We found that Echinopsis and Gymnocalycium species form short-term seed banks. Our results will help to better understand the population dynamics of cactus species, a focal species group for conservation actions because many of them are threatened by human activities., (© 2018 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Intra-individual variation in the production and size of reproductive traits has been documented in columnar cacti, being higher in equator-facing flowers. Such variation is attributed to the high amount of PAR intercepted by stems oriented towards the equator. Most studies focused on this phenomenon have documented the existence of intra-individual variation on traits associated with the female function; however, its impact on traits associated with the male function has been neglected. We tested the hypothesis that equator-facing flowers of Myrtillocactus geometrizans exhibit higher values on traits associated with both male and female functions than flowers facing against it. Number and size of anthers and ovaries, pollen:ovule ratio and number and quality of pollen grains (diameter, germinability, viability and pollen tube length) were estimated from reproductive structures facing north and south, and compared with t-tests between orientations. Number of anthers per flower, number of pollen grains per anther and per floral bud; pollen size, viability and germinability; pollen tube length; ovary length and pollen:ovule ratio were significantly higher in reproductive structures oriented towards the south (i.e. equator). These findings suggest that intra-individual variation in floral traits of M. geometrizans might be associated with different availability of resources in branches with contrasting orientation. Our results provide new evidence of the existence of a response to an orientation-dependent extrinsic gradient. To our knowledge, this is the first study documenting the existence of intra-individual variation on pollen quality and P:O ratio in Cactaceae species., (© 2018 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Naturally dark nighttime environments are being widely eroded by the introduction of artificial light at night (ALAN). The biological impacts vary with the intensity and spectrum of ALAN, but have been documented from molecules to ecosystems. How globally severe these impacts are likely to be depends in large part on the relationship between the spatio-temporal distribution of ALAN and that of the geographic ranges of species. Here, we determine this relationship for the Cactaceae family. Using maps of the geographic ranges of cacti and nighttime stable light composite images for the period 1992 to 2012, we found that a high percentage of cactus species were experiencing ALAN within their ranges in 1992, and that this percentage had increased by 2012. For almost all cactus species (89.7%) the percentage of their geographic range that was lit increased from 1992-1996 to 2008-2012, often markedly. There was a significant negative relationship between the species richness of an area, and that of threatened species, and the level of ALAN. Cacti could be particularly sensitive to this widespread and ongoing intrusion of ALAN into their geographic ranges, especially when considering the potential for additive and synergistic interactions with the impacts of other anthropogenic pressures.

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 geographic patterns, which are synthesised in the Geographic 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 differences between latitude parameters with a Student t-test. Overall, we found more specialised 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 generalised. Our results support the geographic dichotomy hypothesis in the cacti of South America, suggesting that environmental factors underlying the latitudinal patterns can help to explain differences in the pollination syndrome between tropical and extratropical regions., (© 2017 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Three synchronopatric Cactaceae species, Echinopsis rhodotricha, Harrisia balansae and Praecereus saxicola, have mostly nocturnal anthesis and similar flowers, characteristics that motivated us to perform a comparative study of reproductive ecology. Reproductive phenology was sampled monthly from December 2014 to November 2015. 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. 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 h. H. balansae seems to be self-incompatible, while E. rhodotricha presented self-compatibility. P. 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. It can be concluded that the studied species share nocturnal and diurnal pollinators, suggesting interspecific pollen flow, which, however, could not be detected with 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., (© 2017 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Recruitment from seeds is among the most vulnerable stage for plants as global temperatures change. While germination is the means by which the vast majority of the world's flora regenerate naturally, a framework for accurately predicting which species are at greatest risk of germination failure during environmental perturbation is lacking. Taking a physiological approach, we assess how one family, the Cactaceae, may respond to global temperature change based on the thermal buffering capacity of the germination phenotype. We selected 55 cactus species from the Americas, all geo-referenced seed collections, reflecting the broad environmental envelope of the family across 70° of latitude and 3700 m of altitude. We then generated empirical data of the thermal germination response from which we estimated the minimum (T b ), optimum (T o ) and ceiling (T c ) temperature for germination and the thermal time (θ 50 ) for each species based on the linearity of germination rate with temperature. Species with the highest T b and lowest T c germinated fastest, and the interspecific sensitivity of the germination rate to temperature, as assessed through θ 50 , varied tenfold. A left-skewed asymmetry in the germination rate with temperature was relatively common but the unimodal pattern typical of crop species failed for nearly half of the species due to insensitivity to temperature change at T o . For 32 fully characterized species, seed thermal parameters correlated strongly with the mean temperature of the wettest quarter of the seed collection sites. By projecting the mean temperature of the wettest quarter under two climate change scenarios, we predict under the least conservative scenario (+3.7°C) that 25% of cactus species will have reduced germination performance, whilst the remainder will have an efficiency gain, by the end of the 21st century., (© 2017 John Wiley & Sons Ltd.)

Soil seed banks are essential elements of plant population dynamics, enabling species to maintain genetic variability, withstand periods of adversity and persist over time, including for cactus species. However knowledge of the soil seed bank in cacti is scanty. In this study, over a 5-year period we studied the seed bank dynamics, seedling emergence and nurse plant facilitation of Polaskia chende, an endemic columnar cactus of central Mexico. P. chende seeds were collected for a wild population in Puebla, Mexico. Freshly collected seeds were sown at 25 °C and 12-h photoperiod under white light, far-red light and darkness. The collected seeds were divided in two lots, the first was stored in the laboratory and the second was use to bury seeds in open areas and beneath a shrub canopy. Seeds were exhumed periodically over 5 years. At the same time seeds were sown in open areas and beneath shrub canopies; seedling emergence and survival were recorded over different periods of time for 5 years. The species forms long-term persistent soil seed banks. The timing of seedling emergence via germination in the field was regulated by interaction between light, temperature and soil moisture. Seeds entered secondary dormancy at specific times according to the expression of environmental factors, demonstrating irregular dormancy cycling. Seedling survival of P. chende was improved under Acacia constricta nurse plants. Finally, plant facilitation affected the soil seed bank dynamics as it promoted the formation of a soil seed bank, but not its persistence., (© 2017 German Society for Plant Sciences and The Royal Botanical Society of the Netherlands.)

Mexico has one of the highest diversities of barrel cacti species worldwide; however, all are threatened and require conservation policies. Information on their reproductive biology is crucial, but few studies are available. Ferocactus recurvus subsp. recurvus is a barrel cactus endemic to the Tehuacán-Cuicatlán Valley. Our research aimed to characterise its floral and pollination biology. We hypothesised bee pollination, as suggested by its floral morphology and behaviour, and self-incompatibility, like most barrel cacti studied. Three study sites were selected in the semiarid Zapotitlán Valley, Mexico. We examined 190 flowers from 180 plants to determine: morphometry and behaviour of flowers, flower visitors and probable pollinators, and breeding system. Flowers showed diurnal anthesis, lasting 2-5 days, the stigma being receptive on day 2 or 3 after the start of anthesis. Flowers produced scarce/no nectar and main visitors were bees (Apidae), followed by flies (Muscidae), ants (Formicidae), thrips (Thripidae) and hummingbirds (Throchilidae); however, only native bees and occasionally wasps contacted the stigma and anthers. Pollination experiments revealed that this species is self-incompatible and xenogamous. In natural conditions, fruit set was 60% and cross-pollination fruit set was 100%. Percentage seed germination resulting from cross-pollination was higher than in the control treatment. Our results provide ecological information for conservation programmes to ensure a high probability of breeding and seed production in natural populations of F. recurvus., (© 2017 German Botanical Society and The Royal Botanical Society of the Netherlands.)

To determine seed removal influence on seed populations, we need to quantify pre- and post-dispersal seed removal. Several studies have quantified seed removal in temperate American deserts, but few studies have been performed in tropical deserts. These studies have only quantified pre- or post-dispersal seed removal, thus underestimating the influence of seed removal. We evaluated pre- and post-dispersal seed removal in the columnar cactus Stenocereus stellatus in a Mexican tropical desert. We performed selective exclosure experiments to estimate percentage of seeds removed by ants, birds and rodents during the pre- and post-dispersal phases. We also conducted field samplings to estimate abundance of the most common seed removers. Birds (10-28%) removed a higher percentage of seeds than ants (2%) and rodents (1-4%) during pre-dispersal seed removal. Melanerpes hypopolius was probably the main bird removing seeds from fruits. Ants (62-64%) removed a higher percentage of seeds than birds (34-38%) and rodents (16-30%) during post-dispersal seed removal. Pogonomyrmex barbatus was probably the main ant removing seeds from soil. Birds and ants are the main pre- and post-dispersal seed removers in S. stellatus, respectively. Further studies in other S. stellatus populations and plants with different life forms and fruit types will contribute to evaluate seed removal in tropical American deserts., (© 2016 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Historic demography changes of plant species adapted to New World arid environments could be consistent with either the Glacial Refugium Hypothesis (GRH), which posits that populations contracted to refuges during the cold-dry glacial and expanded in warm-humid interglacial periods, or with the Interglacial Refugium Hypothesis (IRH), which suggests that populations contracted during interglacials and expanded in glacial times. These contrasting hypotheses are developed in the present study for the giant columnar cactus Cephalocereus columna-trajani in the intertropical Mexican drylands where the effects of Late Quaternary climatic changes on phylogeography of cacti remain largely unknown. In order to determine if the historic demography and phylogeographic structure of the species are consistent with either hypothesis, sequences of the chloroplast regions psbA-trnH and trnT-trnL from 110 individuals from 10 populations comprising the full distribution range of this species were analysed. Standard estimators of genetic diversity and structure were calculated. The historic demography was analysed using a Bayesian approach and the palaeodistribution was derived from ecological niche modelling to determine if, in the arid environments of south-central Mexico, glacial-interglacial cycles drove the genetic divergence and diversification of this species. Results reveal low but statistically significant population differentiation (FST = 0.124, P < 0.001), although very clear geographic clusters are not formed. Genetic diversity, haplotype network and Approximate Bayesian Computation (ABC) demographic analyses suggest a population expansion estimated to have taken place in the Last Interglacial (123.04 kya, 95% CI 115.3-130.03). The species palaeodistribution is consistent with the ABC analyses and indicates that the potential area of palaedistribution and climatic suitability were larger during the Last Interglacial and Holocene than in the Last Glacial Maximum. Overall, these results suggest that C. columna-trajani experienced an expansion following the warm conditions of interglacials, in accordance with the GRH.

The equatorial orientation of reproductive structures is known in some columnar cacti from extratropical deserts. It has been hypothesised that photosynthetically active radiation (PAR) interception is the main reason for this orientation, because of its key effect on nocturnal CO 2 uptake. However, there are no studies addressing both the effect of PAR and its consequence, carbon gain, on fruit orientation. Accordingly, we tested whether PAR and carbon gain could explain the southern fruit orientation of Myrtillocactus geometrizans, an inter-tropical columnar cactus. We studied three populations of M. geometrizans in Mexico. For each population, azimuth of fruits, total daily PAR, nocturnal acid accumulation (NAA) and fruit production were measured. The relationships between rib orientation and number of fruits, as well as total daily PAR, were evaluated using periodic regressions. The effect of total daily PAR and NAA on number of fruits was assessed using generalised linear models. During spring, mean fruit orientation had a south azimuth for three populations. Likewise, rib orientation had a significant effect on fruit production, with the south-facing ribs having the maximum number of fruits. Total daily PAR was highest in the south-facing ribs, at least for those in the northern and central populations. Furthermore, during spring, there was a significant positive effect of total daily PAR and NAA on fruit production. Our results provide strong evidence that the higher carbon gain in equatorial ribs, through a highest interception of PAR, would be the responsible factor for equatorial orientation of fruits in an inter-tropical columnar cactus., (© 2016 German Botanical Society and The Royal Botanical Society of the Netherlands.)

The aim of this study is the biomimetic optimisation of branched fibre-reinforced composites based on the detailed analysis of biological concept generators. The methods include analyses of the functional morphology and biomechanics of arborescent monocotyledons and columnar cacti as well as measurements and modelling of mechanical properties of biomimetic fibre-reinforced composites. The key results show evidence of notch stress reduction by optimised stem-branch-attachment morphology in monocotyledons and columnar cacti. It could be shown that some of these highly interesting properties can be transferred into biomimetic fibre-reinforced composites.

Annual census data spanning seventy-five years document mortality and regeneration in a population of saguaro cactus (Carnegiea gigantea) in the Cactus Forest of the Rincon Mountain District of Saguaro National Park near Tucson, AZ. On 6 four-hectare plots, each saguaro was censused and a methodical search for new saguaros was conducted annually each year from 1942 through 2016, with the exception of 1955. Regeneration has been episodic with 828 plants established from 1959 through 1993 compared with 34 plants established between 1942 and 1958 and only three plants established after 1993. The years preceding 1959 and following 1993, include some of the driest decades in centuries in southern Arizona. While woodcutting and cattle grazing are believed to be among the causes of decades of failed regeneration prior to 1958, neither of these factors contributed to the failed regeneration following 1993. The height structure of the population from 1942 to 2016 shifted dramatically from a population dominated by large saguaros (> 5.4 m tall) in the first three decades of the study to a population dominated by small saguaros (< 1.8 m tall) in the most recent two decades. Mortality is shown to be strongly age dependent. In the year following the 2011 catastrophic freeze, 21 of 59 plants older than 80 years died compared with zero deaths in 270 plants between the ages of 29 and 80 years. Saguaros under 40 years old, growing under small shrubs or in the open, have a lower probability of survival than better protected saguaros. Long-term population monitoring is essential to understanding the complex impacts of human and environmental factors on the population dynamics of long-lived species.

The germination characteristics of the native cactus species are poorly known, being the temperature and the light the factors that the most interferes in that process. Thus, the objective of the present work was to characterize the fruits and evaluate the influence of the temperature and the light in the seed germination of Rhipsalis floccosa, Rhipsalis pilocarpa and Rhipsalis teres. The tested constant temperatures were 15, 20, 25, 30 and 35 °C and the alternate of 20-30 °C and 25-35 °C in a photoperiod of 10 hours, and with determination of the most appropriate temperature, the germination was tested in light absence. The germination percentage, the index of germination speed and medium time of germination were evaluated. For R. floccosa, the highest germination percentage was at 20 °C. For R. pilocarpa and R. teres, the highest germination percentages occurred in 15 °C and 20 °C. There was correlation to germination percentage between the three species, indicating that they had similar germination behavior. Total absence of germination was verified for the three species in condition of light absence. In conclusion, the temperature of 20 °C is the most suitable for the seed germination of R. floccosa. For the species R. pilocarpa and R. teres, the temperatures of 15 and 20 °C are the most suitable.

Pereskia aculeata and Pereskia grandifolia have been studied widely due to their high nutritional and therapeutic values. However, little is known about the biological requirements of their seeds for the various germination factors. Thus, this experiment aimed to evaluate the thermal effects on the germination of these species at the temperatures of 24°C, 27°C, 30°C, 33°C and 36°C. After verification of the existence of differences in the performance of germination, a non-linear regression was carried out, relating the germination to temperature and identifying its point of maximum efficiency. We found that the lowest synchronization indexes of germination were observed close to 30°C. The best germination response of the P. aculeata and P. grandifolia was observed at 30°C and 33°C, respectively, with greater germination strength and fewer days to attain 63.21% of germinations. The results obtained from the germination of P. aculeata and P. grandifolia can be described by the Weindull distribution model with three parameters, as proposed by Carneiro and Guedes (1992)., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Cereus peruvianus (Peruvian apple cactus) is a large erect and thorny succulent cactus characterized by column-like (cereus [L]: column), that is, candle-shaped, appendages. For three successive years (1100 days), between early April and late November, we studied the flowering patterns of eight cacti growing in public gardens and rural areas of north and central Tunisia, far from nighttime artificial illumination, in relation to natural environmental light, temperature, relative humidity and precipitation parameters. Flower blooming was assessed nightly between 23:00 h and until at least 02:00 h, and additionally around-the-clock at ~1 h intervals for 30 consecutive days during the late summer of each year of study to quantify both nyctohemeral (day-night) and lunar patterns. During the summer months of prolonged daytime photoperiod, flower blooming of C. peruvianus exhibited predictable-in-time variation as "waves" with average period of 29.5 days synchronized by the light of the full moon. The large-sized flower (~16 cm diameter) opens almost exclusively at night, between sunset and sunrise, as a 24 h rhythm during a specific 3-4-day span of the lunar cycle (full moon), with a strong correlation between moon phase and number and proportion of flowers in bloom (ranging from r = +0.59 to +0.91). Black, blue and red cotton sheets were used to filter specific spectral bands of nighttime moonlight from illuminating randomly selected plant appendages as a means to test the hypothesis of a "gating" 24 h rhythm phenomenon of photoreceptors at the bud level. Relative to control conditions (no light filtering), black sheet covering inhibited flower bud induction by 87.5%, red sheet covering by 46.6% and blue sheet covering by 34%, and the respective inhibiting effects on number of flowers in bloom were essentially 100%, ~81% and ~44%. C. peruvianus is a unique example of a terrestrial plant that exhibits a circadian flowering rhythm (peak ~00:00 h) "gated" by 24 h, lunar 29.5-day (bright light of full moon) and annual 365.25-day (prolonged summertime day length) environmental photoperiod cycles.

Many cactus species produce 'bird' flowers; however, the reproductive biology of the majority of these species has not been studied. Here, we report on a study of the pollination of two species from the Cleistocactus genus, cited as an ornithophilous genus, in the context of the different ways in which they are specialised to bird pollination. In addition, we re-evaluate the level of specialisation of previous studies of cacti with bird pollination and evaluate how common phenotypic specialisation to birds is in this family. Both Cleistocactus species exhibited ornithophilous floral traits. Cleistocactus baumannii was pollinated by hummingbirds, whereas Cleistocactus smaragdiflorus was pollinated by hummingbirds and bees. Pollination by birds has been recorded in 27 cactus species, many of which exhibit ornithophilous traits; however, they show generalised pollination systems with bees, bats or moths in addition to birds being their floral visitors. Of all cactus species, 27% have reddish flowers. This trait is associated with diurnal anthesis and a tubular shape. Phenotypic specialisation to bird pollination is recognised in many cactus species; however, it is not predictive of functional and ecological specialisation in this family., (© 2014 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Premise of the Study: Species of the endemic Chilean cactus genus Copiapoa have cylindrical or (sub)globose stems that are solitary or form (large) clusters and typically yellow flowers. Many species are threatened with extinction. Despite being icons of the Atacama Desert and well loved by cactus enthusiasts, the evolution and diversity of Copiapoa has not yet been studied using a molecular approach., Methods: Sequence data of three plastid DNA markers (rpl32-trnL, trnH-psbA, ycf1) of 39 Copiapoa taxa were analyzed using maximum likelihood and Bayesian inference approaches. Species distributions were modeled based on geo-referenced localities and climatic data. Evolution of character states of four characters (root morphology, stem branching, stem shape, and stem diameter) as well as ancestral areas were reconstructed using a Bayesian and maximum likelihood framework, respectively., Key Results: Clades of species are revealed. Though 32 morphologically defined species can be recognized, genetic diversity between some species and infraspecific taxa is too low to delimit their boundaries using plastid DNA markers. Recovered relationships are often supported by morphological and biogeographical patterns. The origin of Copiapoa likely lies between southern Peru and the extreme north of Chile. The Copiapó Valley limited colonization between two biogeographical areas., Conclusions: Copiapoa is here defined to include 32 species and five heterotypic subspecies. Thirty species are classified into four sections and two subsections, while two species remain unplaced. A better understanding of evolution and diversity of Copiapoa will allow allocating conservation resources to the most threatened lineages and focusing conservation action on real biodiversity., (© 2015 Botanical Society of America.)

Abiotic stresses usually cause H2O2 accumulation, with harmful effects, in plants. Catalase may play a key protective role in plant cells by detoxifying this excess H2O2. Pitaya (Hylocereus undatus) shows broad ecological adaptation due to its high tolerance to abiotic stresses, e.g. drought, heat and poor soil. However, involvement of the pitaya catalase gene (HuCAT) in tolerance to abiotic stresses is unknown. In the present study, a full-length HuCAT3 cDNA (1870 bp) was isolated from pitaya based on our previous microarray data and RACE method. The cDNA sequence and deduced amino acid sequence shared 73-77% and 75-80% identity with other plant catalases, respectively. HuCAT3 contains conserved catalase family domain and catalytic sites. Pairwise comparison and phylogenetic analysis indicated that HuCAT3 is most similar to Eriobotrya japonica CAT, followed by Dimocarpus longan CAT and Nicotiana tabacum CAT1. Expression profile analysis demonstrated that HuCAT3 is mainly expressed in green cotyledons and mature stems, and was regulated by H2O2, drought, cold and salt stress, whereas, its expression patterns and maximum expression levels varied with stress types. HuCAT activity increased as exposure to the tested stresses, and the fluctuation of HuCAT activity was consistent with HuCAT3 mRNA abundance (except for 0.5 days upon drought stress). HuCAT3 mRNA elevations and HuCAT activities changes under cold stress were also in conformity with the cold tolerances among the four genotypes. The obtained results confirmed a major role of HuCAT3 in abiotic stress response of pitaya. This may prove useful in understanding pitaya's high tolerance to abiotic stresses at molecular level., (Copyright © 2015 Elsevier B.V. All rights reserved.)

Premise of the Study: A key question concerns the vulnerability of desert species adapted to harsh, variable climates to future climate change. Evaluating this requires coupling long-term demographic models with information on past and projected future climates. We investigated climatic drivers of population growth using a 22-yr demographic model for Pediocactus bradyi, an endangered cactus in northern Arizona., Methods: We used a matrix model to calculate stochastic population growth rates (λs) and the relative influences of life-cycle transitions on population growth. Regression models linked population growth with climatic variability, while stochastic simulations were used to (1) understand how predicted increases in drought frequency and extreme precipitation would affect λs, and (2) quantify variability in λs based on temporal replication of data., Key Results: Overall λs was below unity (0.961). Population growth was equally influenced by fecundity and survival and significantly correlated with increased annual precipitation and higher winter temperatures. Stochastic simulations increasing the probability of drought and extreme precipitation reduced λs, but less than simulations increasing the probability of drought alone. Simulations varying the temporal replication of data suggested 14 yr were required for accurate λs estimates., Conclusions: Pediocactus bradyi may be vulnerable to increases in the frequency and intensity of extreme climatic events, particularly drought. Biotic interactions resulting in low survival during drought years outweighed increased seedling establishment following heavy precipitation. Climatic extremes beyond historical ranges of variability may threaten rare desert species with low population growth rates and therefore high susceptibility to stochastic events., (© 2014 Botanical Society of America, Inc.)

Mutualism is an often complex interaction among multiple species, each of which may respond differently to abiotic conditions. The effects of temperature on the formation, dissolution, and success of these and other species interactions remain poorly understood. We studied the thermal ecology of the mutualism between the cactus Ferocactus wislizeni and its ant defenders (Forelius pruinosus, Crematogaster opuntiae, Solenopsis aurea, and Solenopsis xyloni) in the Sonoran Desert, USA. The ants are attracted to extrafloral nectar produced by the plants and, in exchange, protect the plants from herbivores; there is a hierarchy of mutualist effectiveness based on aggression toward herbivores. We determined the relationship between temperature and ant activity on plants, the thermal tolerance of each ant species, and ant activity in relation to the thermal environment of plants. Temperature played a role in determining which species interact as mutualists. Three of the four ant species abandoned the plants during the hottest part of the day (up to 40 °C), returning when surface temperature began to decrease in the afternoon. The least effective ant mutualist, F. pruinosus, had a significantly higher critical thermal maximum than the other three species, was active across the entire range of plant surface temperatures observed (13.8-57.0 °C), and visited plants that reached the highest temperatures. F. pruinosus occupied some plants full-time and invaded plants occupied by more dominant species when those species were thermally excluded. Combining data on thermal tolerance and mutualist effectiveness provides a potentially powerful tool for predicting the effects of temperature on mutualisms and mutualistic species.

Columnar cacti occur naturally in many habitats and environments in the Americas but are conspicuously dominant in very dry desert regions. These majestic plants are widely regarded for their cultural, economic, and ecological value and, in many ecosystems, support highly diverse communities of pollinators, seed dispersers, and frugivores. Massive amounts of water and other resources stored in the succulent photosynthetic stems of these species confer a remarkable ability to grow and reproduce during intensely hot and dry periods. Yet many columnar cacti are potentially under severe threat from environmental global changes, including climate change and loss of habitat. Stems in columnar cacti and other cylindrical-stemmed cacti are morphologically diverse; stem volume-to-surface area ratio (V:S) across these taxa varies by almost two orders of magnitude. Intrinsic functional trade-offs are examined here across a broad range of V:S in species of columnar cacti. It is proposed that variation in photosynthetic gas exchange, growth, and response to stress is highly constrained by stem V:S, establishing a mechanistic framework for understanding the sensitivity of columnar cacti to climate change and drought. Specifically, species that develop stems with low V:S, and thus have little storage capacity, are expected to express high mass specific photosynthesis and growth rates under favourable conditions compared with species with high V:S. But the trade-off of having little storage capacity is that low V:S species are likely to be less tolerant of intense or long-duration drought compared with high V:S species. The application of stable isotope measurements of cactus spines as recorders of growth, water relations, and metabolic responses to the environment across species of columnar cacti that vary in V:S is also reviewed. Taken together, our approach provides a coherent theory and required set of observations needed for predicting the responses of columnar cacti to climate change., (© The Author 2014. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

In this work, motivated by an approach used in a cactus to collect fog, we have developed an artificial water-collection structure. This structure includes a large ZnO wire and an array of small ZnO wires that are branched on the large wire. All these wires have conical shapes, whose diameters gradually increase from the tip to the root of a wire. Accordingly, a water drop that is condensed on the tip of each wire is driven to the root by a capillary force induced by this diameter gradient. The lengths of stem and branched wires in the synthesized structures are in the orders of 1 mm and 100 μm, respectively. These dimensions are, respectively, comparable to and larger than their counterparts in the case of a cactus. Two groups of tests were conducted at relative humidity of 100% to compare the amounts of water collected by artificial and cactus structures within specific time durations of 2 and 35 s, respectively. The amount of water collected by either type of structures was in the order of 0.01 μL. However, on average, what has been collected by the artificial structures was 1.4-5.0 times more than that harvested by the cactus ones. We further examined the mechanism that a cactus used to absorb a collected water drop into its stem. On the basis of the gained understanding, we developed a setup to successfully collect about 6 μL of water within 30 min.

Succulent plants are widely distributed, reaching their highest diversity in arid and semi-arid regions. Their origin and diversification is thought to be associated with a global expansion of aridity. We test this hypothesis by investigating the tempo and pattern of Cactaceae diversification. Our results contribute to the understanding of the evolution of New World Succulent Biomes. We use the most taxonomically complete dataset currently available for Cactaceae. We estimate divergence times and utilize Bayesian and maximum likelihood methods that account for nonrandom taxonomic sampling, possible extinction scenarios and phylogenetic uncertainty to analyze diversification rates, and evolution of growth form and pollination syndrome. Cactaceae originated shortly after the Eocene-Oligocene global drop in CO2 , and radiation of its richest genera coincided with the expansion of aridity in North America during the late Miocene. A significant correlation between growth form and pollination syndrome was found, as well as a clear state dependence between diversification rate, and pollination and growth-form evolution. This study suggests a complex picture underlying the diversification of Cactaceae. It not only responded to the availability of new niches resulting from aridification, but also to the correlated evolution of novel growth forms and reproductive strategies., (© 2014 The Authors. New Phytologist © 2014 New Phytologist Trust.)

While plants require radiation for photosynthesis, radiation in warm deserts can have detrimental effects from high temperatures. This dilemma may be solved through plant morphological attributes. In cold deserts, stem tilting keeps reproductive organs warm by increasing radiation interception at the cost of decreased annual light interception. Conversely, little is known about stem tilting in warm deserts. We hypothesised that stem tilting in Echinocactus platyacanthus prevents high temperatures near the apex, where reproduction occurs. The study was conducted in the warm, inter-tropical portion of the Chihuahuan Desert, Mexico. We found that cacti preferentially tilted towards the south, which reduced temperatures of reproductive organs during the hot season, but increased total annual near-apex PAR interception. Tilting also maximised reproduction, a likely consequence of temperature control but perhaps also of the difficulty in translocating photosynthates in cacti; therefore, annual energy acquisition near floral meristems may be largely allocated to reproduction. Unlike plants of higher latitudes, in inter-tropical deserts sunlight at noon comes either from the north or the south, depending on the season, and thus stem tilting may more strongly affect total annual radiation received in different portions of the stem. Inter-tropical cacti can synchronise reproduction with irradiance peaks if flowering occurs in a specific (north or south) portion of the stem; also, they effectively solve the conflict between maximising annual PAR interception and minimising temperature at the hottest time of day. Notably, the two inter-tropical cacti in which stem tilting has been studied successfully solve this conflict., (© 2013 German Botanical Society and The Royal Botanical Society of the Netherlands.)