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Chloroplast genomes (plastomes) are frequently treated as highly conserved among land plants. However, many lineages of vascular plants have experienced extensive structural rearrangements, including inversions and modifications to the size and content of genes. Cacti are one of these lineages, containing the smallest plastome known for an obligately photosynthetic angiosperm, including the loss of one copy of the inverted repeat (∼25 kb) and the gene suite, but only a few cacti from the subfamily Cactoideae have been sufficiently characterized. Here, we investigated the variation of plastome sequences across the second-major lineage of the Cactaceae, the subfamily Opuntioideae, to address (1) how variable is the content and arrangement of chloroplast genome sequences across the subfamily, and (2) how phylogenetically informative are the plastome sequences for resolving major relationships among the clades of Opuntioideae. Our assembly of the plastome recovered an organelle of 150,347 bp in length with both copies of the inverted repeat and the presence of all the gene suite. An expansion of the large single copy unit and a reduction of the small single copy unit was observed, including translocations and inversion of genes, as well as the putative pseudogenization of some loci. Comparative analyses among all clades within Opuntioideae suggested that plastome structure and content vary across taxa of this subfamily, with putative independent losses of the gene suite and pseudogenization of genes across disparate lineages, further demonstrating the dynamic nature of plastomes in Cactaceae. Our plastome dataset was robust in resolving three tribes with high support within Opuntioideae: Cylindropuntieae, Tephrocacteae and Opuntieae. However, conflicting topologies were recovered among major clades when exploring different assemblies of markers. A plastome-wide survey for highly informative phylogenetic markers revealed previously unused regions for future use in Sanger-based studies, presenting a valuable dataset with primers designed for continued evolutionary studies across Cactaceae. These results bring new insights into the evolution of plastomes in cacti, suggesting that further analyses should be carried out to address how ecological drivers, physiological constraints and morphological traits of cacti may be related with the common rearrangements in plastomes that have been reported across the family.

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.

Premise of the study: Cactaceae is one of the most charismatic plant families because of the extreme succulence and outstanding diversity of growth forms of its members. Although cacti are conspicuous elements of arid ecosystems in the New World and are model systems for ecological and anatomical studies, the high morphological convergence and scarcity of phenotypic synapomorphies make the evolutionary relationships and trends among lineages difficult to understand. METHODS: We performed phylogenetic analyses implementing parsimony ratchet and likelihood methods, using a concatenated matrix with 6148 bp of plastid and nuclear markers (trnK/matK, matK, trnL-trnF, rpl16, and ppc). We included 224 species representing approximately 85% of the family's genera. Likelihood methods were used to perform an ancestral character reconstruction within Cactoideae, the richest subfamily in terms of morphological diversity and species number, to evaluate possible growth form evolutionary trends. Key results: Our phylogenetic results support previous studies showing the paraphyly of subfamily Pereskioideae and the monophyly of subfamilies Opuntioideae and Cactoideae. After the early divergence of Blossfeldia, Cactoideae splits into two clades: Cacteae, including North American globose and barrel-shaped members, and core Cactoideae, including the largest diversity of growth forms distributed throughout the American continent. Para- or polyphyly is persistent in different parts of the phylogeny. Main Cactoideae clades were found to have different ancestral growth forms, and convergence toward globose, arborescent, or columnar forms occurred in different lineages. CONCLUSIONS: Our study enabled us to provide a detailed hypothesis of relationships among cacti lineages and represents the most complete general phylogenetic framework available to understand evolutionary trends within Cactaceae.

Species with various reproductive modes accompanied by different mechanical properties of their (lateral) branch-branch junctions have evolved in the cactus subfamily Opuntioideae. Older branches of Opuntia ficus-indica with fracture-resistant junctions often bear flowers and fruits for sexual reproduction, whereas younger branches break off easily and provide offshoots for vegetative propagation. Cylindropuntia bigelovii plants are known for their vegetative reproduction via easily detachable branches that can establish themselves as offshoots. We characterized the elastic and fracture behaviors of these lateral junctions by tensile testing and analyzed local strains during loading. Additionally, we carried out finite element analyses to quantify the influence of five relevant tissue layers on joint elastic behavior. Our fracture analysis revealed various fracture modes: (i) most young samples of Opuntia ficus-indica failed directly at the junction and had smooth fracture surfaces, and relative fracture strain was on median 4% of the total strain; (ii) most older samples of Opuntia ficus-indica failed at the adjacent branch and exhibited rough fracture surfaces, and relative fracture strain was on median 47%; (iii) most samples of Cylindropuntia bigelovii abscised directly at the junction and exhibited cup and cone surfaces, and relative fracture strain was on median 28%. Various geometric and mechanical properties such as junction area, fracture energy, and tensile strength were analyzed with respect to significant differences between species and age of sample. Interestingly, the abscission of lateral branches naturally triggered by wind, passing animals, or vibration showed the following differences in maximum force: 153 N (older Opuntia ficus-indica ), 51 N (young Opuntia ficus-indica ), and 14 N ( Cylindropuntia bigelovii ).

Damage-repair is particularly important for the maintenance of the water-storing abilities of succulent plants such as cacti. Comparative morphological, anatomical, and biomechanical analyses of self-repair were performed on artificially wounded branches of Opuntia ficus-indica and Cylindropuntia bigelovii. Macroscopic observations, contrast staining, and lignin-proof staining were used to investigate morphological and anatomical responses after wounding at various time intervals. Two-point bending tests were repeatedly performed on the same branches under unwounded, freshly wounded, and healed conditions by using customized 3D-printed clamping jaws. Morphologically, both species showed a rolling-in of the wound edges, but no mucilage discharge. Anatomically, ligno-suberized peridermal layers developed that covered the wound region, and new parenchyma cells formed, especially in O. ficus-indica. In all samples, the wounding effect directly after damage caused a decrease between 18% and 37% in all the tested mechanical parameters, whereas a positive healing effect after 21 days was only found for C. bigelovii. Based on our data, we hypothesize a high selection pressure on the restoration of structural integrity in the wound area, with a focus on the development of efficient water-retaining mechanisms, whereas the concept of “sufficient is good enough” seems to apply for the restoration of the mechanical properties.

The invasion of Opuntioideae (cacti) was assessed at national, European, and global levels >14 years ago and requires an urgent update. At present, only three species of this subfamily are retained in the official list of the Swiss Federal Bureau of Environment, two of them being potentially invasive. About 550 new observations of Opuntia Mill. and Cylindropuntia (Engelm.) F. M. Knuth in the Swiss Cantons Valais, Vaud, and Ticino have been entered in the Swiss Info Flora database. Our observations include the revision of previously registered and new stations, using a revised taxonomy presented here. Our study covering the last 5 years reveals 12 species, each with a specific ecology. Four species have an invasive potential at a local level in the xeric Central Valais. We characterized the habitat of each species by bedrock and soil type, exposure, and microclimate. Opuntia mesacantha (ex O. humifusa Complex) and O. phaeacantha are currently invasive in the municipality of Sion. Opuntia cymochila (hitherto lumped with O. phaeacantha) is dominant in the municipality of Fully, and Opuntia engelmannii is expanding east of Fully. In Ticino and the Jura foothills, adaptations to 2–3-fold annual precipitation and limestone bedrock are explained by the different soil properties and root systems of Opuntias. Methods of control, in particular, the uprooting currently practiced in Fully and Sion, are discussed.

Opuntioideae, composed of roughly 370 species, occur in almost every biome in the Americas, from seasonally dry tropical forests (SDTF) to high-elevation Andean grasslands, American deserts and temperate zones. The interrelationships among the three major clades of Opuntioideae (Cylindropuntieae, Opuntieae and Tephrocacteae) are not well resolved, and thus, the ancestral habitat, biogeographic history and evolution of morphological characters, such as large photosynthetic leaves and flattened stems, are poorly understood. To test their geographic origin and evolution of key morphological characters, we built the largest phylogenomic dataset for Cactaceae to date using 103 plastid genes of 107 taxa of Opuntioideae. The subfamily Opuntioideae likely evolved in South America in a combination of seasonally dry tropical forest (SDTF)/desert habitats. Opuntieae most likely evolved in South America in SDTF and, from there, moved into desert regions, Chaco and temperate/subtropical zones, while Tephrocacteae and Cylindropuntieae evolved in South America in desert regions and moved into SDTF, Chaco and temperate/subtropical zones. Analyses of morphological evolution suggest that, although large leaves are plesiomorphic in Opuntioideae, long-lived, photosynthetically active leaves in Cylindropuntieae and Tephrocacteae are homoplasious and do not represent retained plesiomorphy, as is often assumed. Flattened stems are synapomorphic for Opuntieae, possibly representing adaptation to competition for light resources in SDTF, their most likely ancestral area.

Cactaceae comprise a diverse and iconic group of flowering plants which are almost exclusively indigenous to the New World. The wide variety of growth forms found amongst the cacti have led to the trafficking of many species throughout the world as ornamentals. Despite the evolution and physiological properties of these plants having been extensively studied, little research has focused on cactus-associated viral communities. While only single-stranded RNA viruses had ever been reported in cacti, here we report the discovery of cactus-infecting single-stranded DNA viruses. These viruses all apparently belong to a single divergent species of the family Geminiviridae and have been tentatively named Opuntia virus 1 (OpV1). A total of 79 apparently complete OpV1 genomes were recovered from 31 different cactus plants (belonging to 20 different cactus species from both the Cactoideae and Opuntioideae clades) and from nine cactus-feeding cochineal insects (Dactylopius sp.) sampled in the USA and Mexico. These 79 OpV1 genomes all share > 78.4% nucleotide identity with one another and < 64.9% identity with previously characterized geminiviruses. Collectively, the OpV1 genomes display evidence of frequent recombination, with some genomes displaying up to five recombinant regions. In one case, recombinant regions span ~40% of the genome. We demonstrate that an infectious clone of an OpV1 genome can replicate in Nicotiana benthamiana and Opuntia microdasys. In addition to expanding the inventory of viruses that are known to infect cacti, the OpV1 group is so distantly related to other known geminiviruses that it likely represents a new geminivirus genus. It remains to be determined whether, like its cactus hosts, its geographical distribution spans the globe.

The Opuntioideae include iconic cacti whose lateral branch–branch junctions are intriguing objects from a mechanical viewpoint. We have compared Opuntia ficus-indica, which has stable branch connections, with Cylindropuntia bigelovii, whose side branches abscise under slight mechanical stress. To determine the underlying structures and mechanical characteristics of these stable versus shedding cacti junctions, we conducted magnetic resonance imaging, morphometric and anatomical analyses of the branches and tensile tests of individual tissues. The comparison revealed differences in geometry, shape and material properties as follows: (i) a more pronounced tapering of the cross-sectional area towards the junctions supports the abscission of young branches of C. bigelovii. (ii) Older branches of O. ficus-indica form, initially around the branch–branch junctions, collar-shaped periderm tissue. This secondary coverage mechanically stiffens the dermal tissue, giving a threefold increase in strength and a tenfold increase in the elastic modulus compared with the epidermis. (iii) An approximately 200-fold higher elastic modulus of the vascular bundles of O. ficus-indica is a prerequisite for the stable junction of its young branches. Our results provide, for both biological and engineered materials systems, important insights into the geometric characteristics and mechanical properties of branching joints that are either stable or easily detachable.

Botanical surveys undertaken in Central and Northern Tunisia (North Africa), mostly between 2015 and 2020, have revealed three new allochthonous cacti. The reported taxa belong to the genus Cylindropuntia (Cactaceae, subfamily Opuntioideae, tribe Cylindropuntieae). C. leptocaulis, C. prolifera and C. spinosior are here reported for the first time from North Africa. Updated nomenclature, descriptions, general and national distributions are pointed out for each species. An analytical key and field photographs are also provided.