Alfred James Ewart

ABSTRACTAimFerns are globally distributed, yet the number of studies examining the historical evolution of African taxa is relatively low. Investigation of the evolution of African fern diversity is critical in order to understand patterns and processes that have global relevance (e.g., the pantropical diversity disparity [PDD] pattern). This study aims to examine when and from where a globally distributed fern lineage arrived in sub‐Saharan Africa, to obtain a better understanding of potential processes contributing to patterns of diversity across the region.LocationGlobal, sub‐Saharan Africa.TaxonAsplenium (Aspleniaceae).MethodsWe analysed five loci from 537 Asplenium taxa using a maximum likelihood (IQ‐Tree) phylogenetic framework. For age estimation, we performed penalised likelihood as implemented in treePL, and executed a Bayesian analysis using BEAST. Biogeographical analyses were carried out using BioGeoBEARS.ResultsMost dispersals into Africa occurred within the last ~55 myr, with the highest diversity of sub‐Saharan African taxa concentrated in two clades, each of which descended from an Asian ancestor. Additional dispersals to sub‐Saharan Africa can be found throughout the phylogeny. Lastly, potential cryptic species diversity exists within Asplenium as evidenced by several polyphyletic taxa.Main ConclusionsWe recover multiple dispersals of Asplenium to sub‐Saharan Africa, with two major lineages likely diversifying after arrival.

Premise Cleomaceae is an important model clade for studies of evolutionary processes including genome evolution, floral form diversification, and photosynthetic pathway evolution. Diversification and divergence patterns in Cleomaceae remain tangled as research has been restricted by its worldwide distribution, limited genetic sampling and species coverage, and a lack of definitive fossil calibration points.MethodsWe used target sequence capture and the Angiosperms353 probe set to perform a phylogenetic study of Cleomaceae. We estimated divergence times and biogeographic analyses to explore the origin and diversification of the family. Seed morphology across extant taxa was documented with multifocal image‐stacking techniques and morphological characters were extracted, analyzed, and compared to fossil records.ResultsWe recovered a well‐supported and resolved phylogenetic tree of Cleomaceae generic relationships that includes 236 (~86%) species. We identified 11 principal clades and confidently placed Cleomella as sister to the rest of the family. Our analyses suggested that Cleomaceae and Brassicaceae diverged ~56 mya, and Cleomaceae began to diversify ~53 mya in the Palearctic and Africa. Multiple transatlantic disjunct distributions were identified. Seeds were imaged from 218 (~80%) species in the family and compared to all known fossil species.ConclusionsOur results represent the most comprehensive phylogenetic study of Cleomaceae to date. We identified transatlantic disjunctions and proposed explanations for these patterns, most likely either long‐distance dispersals or contractions in latitudinal distributions caused by climate change over geological timescales. We found that seed morphology varied considerably but mostly mirrored generic relationships.

Parasitic weeds in the genera Orobanche, Phelipanche (broomrapes) and Striga (witchweeds) have a devastating impact on food security across much of Africa, Asia and the Mediterranean Basin. Yet, how climatic factors might affect the range expansion of these weeds in the context of global environmental change remains unexplored. We examined satellite‐based environmental variables such as surface temperature, root zone soil moisture, and elevation, in relation to parasitic weed distribution and environmental conditions over time, in combination with observational data from the Global Biodiversity Information Facility (GBIF). Our analysis reveals contrasting environmental and altitude preferences in the genera Striga and Orobanche. Asiatic witchweed (Striga asiatica), which infests corn, rice, sorghum, and sugar cane crops, appears to be expanding its range in high elevation habitats. It also shows a significant association with heat‐moisture coupling events, the frequency of which is rising in such environments. These results point to geographical shifts in distribution and abundance in parasitic weeds due to climate change.

Myoporum is a genus of trees and shrubs native to the Northern Hemisphere that has been introduced to many parts of the world, mainly for ornamental purposes. We assessed the introduction history, distribution, and extent of naturalization/invasion for Myoporum species in South Africa.Information was collated to determine key events associated with the introduction, establishment, and naturalization of Myoporum in South Africa. Data were collated to determine the current distribution of the genus in South Africa. Twenty sites in the Western Cape were sampled to determine correlates of naturalization. Myoporum was first recorded in South Africa in 1934. Three species were confirmed to be present in South Africa: M. insulare, M. laetum and M. montanum (37 %, 25 % and 24 % of all iNaturalist records respectively). Most records are from the Western Cape (91 %) and small parts of the Eastern Cape; isolated populations occur in Gauteng and the Northern Cape. We could not confirm the presence M. petiolatum, M. tenuifolium or M. tetrandrum. Field surveys revealed widespread naturalization of M. insulare (46 % of all Research Grade observations in iNaturalist); this species was categorized code D1 in the introduction-naturalization-invasion continuum. Myoporum laetum (C3) and M. montanum (C2) are also widely naturalized but over smaller areas. Naturalized populations comprised predominantly juvenile M. insulare plants occurring in highly disturbed (transformed) habitats. Formal risk analyses for all Myoporum species in South Africa are needed as the basis for re-evaluation of their status in national legislation.

Present-day geographic and phylogenetic patterns often reflect the geological and climatic history of the planet. Neontological distribution data are often sufficient to unravel a lineage’s biogeographic history, yet ancestral range inferences can be at odds with fossil evidence. Here, I use the fossilized birth–death process and the dispersal–extinction cladogenesis model to jointly infer the dated phylogeny and range evolution of the tree fern order Cyatheales. I use data for 101 fossil and 442 extant tree ferns to reconstruct the biogeographic history of the group over the last 220 million years. Fossil-aware reconstructions evince a prolonged occupancy of Laurasia over the Triassic–Cretaceous by Cyathealean tree ferns, which is evident in the fossil record but hidden from analyses relying on neontological data alone. Nonetheless, fossil-aware reconstructions are affected by uncertainty in fossils’ phylogenetic placement, taphonomic biases, and specimen sampling and are sensitive to interpretation of paleodistributions and how these are scored. The present results highlight the need and challenges of incorporating fossils into joint inferences of phylogeny and biogeography to improve the reliability of ancestral geographic range estimation.

Dense branching and spines are common features of plant species in ecosystems with high mammalian herbivory pressure. While dense branching and spines can inhibit herbivory independently, when combined, they form a powerful defensive cage architecture. However, how cage architecture evolved under mammalian pressure has remained unexplored. Here we show how dense branching and spines emerged during the age of mammalian radiation in the Combretaceae family and diversified in herbivore-driven ecosystems in the tropics. Phylogenetic comparative methods revealed that modern plant architectural strategies defending against large mammals evolved via a stepwise process. First, dense branching emerged under intermediate herbivory pressure, followed by the acquisition of spines that supported higher speciation rates under high herbivory pressure. Our study highlights the adaptive value of dense branching as part of a herbivore defence strategy and identifies large mammal herbivory as a major selective force shaping the whole plant architecture of woody plants. This study explores the evolution of two traits, branching density and spine presence, in the globally distributed plant family Combretaceae. These traits were found to have appeared in a two-step process in response to mammalian herbivory pressure, revealing the importance of large mammals in the evolution of plant architecture diversity.

The number of naturalised and invasive woody plant species has increased rapidly in recent decades. Despite the increasing interest in tree and shrub invasions, little is known about the invasion ecology of most species. This paper explores the global movement of species in the genus Melaleuca (Myrtaceae; here including the genus Callistemon). We assess the global introduction history, distribution and biogeographic status of the genus. Various global species occurrence databases, citizen science (iNaturalist), and the literature were used.Seventy-two species [out of 386 Melaleuca species; 19%] have been introduced to at least 125 regions outside their native range. The main regions of global Melaleuca introductions are Southeast Asia, the southern parts of North America, south-eastern South America, southern Africa and Europe. The earliest record of a Melaleuca species outside of the native range of the genus is 1789. First records of Melaleuca species outside their native range were most commonly recorded in the 1960s, with records from all over the world. The main reasons for Melaleuca introductions were for use in the tea tree (pharmaceutical value) and ornamental horticulture industries. Melaleuca introductions, naturalizations and invasions are recent compared to many other woody plant taxa. Experiences in Florida and South Africa highlight the potential of Melaleuca species to spread rapidly and have significant ecological impacts. It is likely that the accumulating invasion debt will result in further naturalization and invasion of Melaleuca species in the future.

Paleogene flower-insect interactions and paleo-pollination processes are, in general, poorly understood and fossil evidence for such floral and faunal interactions are rarely reported. To shed light on angiosperm flower-insect interactions, we investigated several hundred fossil flowers and insects from the middle Eocene Fossil Lagerstätte of Eckfeld, Germany. During our work, we discovered a unique fossil Ludwigia flower (bud) with in situ pollen. The ecological preferences (climate, biome, habitat, etc.) of extant Ludwigia and the paleoecological configurations of the fossil plant assemblage support the taxonomic affiliation of the flower bud and an Eocene presence of Ludwigia in the vicinity of the former Lake Eckfeld. Today’s Ludwigia are mostly pollinated by Hymenoptera (bees). Therefore, we screened all currently known hymenopteran fossils from Eckfeld but found no Ludwigia pollen adhering to any of the specimens. On the contrary, we discovered Ludwigia pollen adhering to two different groups of Coleoptera (beetles). Our study suggests that during the Eocene of Europe, Ludwigia flowers were visited and probably pollinated by beetles and over time there was a shift in primary flower visitors/pollinators, from beetles to bees, sometime during the late Paleogene to Neogene.

Aim The aim of this study was to further advance our understanding of the species-rich, and ecologically important angiosperm family Combretaceae to provide new insights into their evolutionary history. We assessed phylogenetic relationships in the family using target capture data and produced a dated phylogenetic tree to assess fruit dispersal modes and patterns of distribution. Location Tropical and subtropical regions. Time Period Cretaceous to present. Major Taxa Studied Family Combretaceae is a member of the rosid clade and comprises 10 genera and more than 500 species, predominantly assigned to genera Combretum and Terminalia, and occurring on all continents and in a wide range of ecosystems. Methods We use a target capture approach and the Angiosperms353 universal probes to reconstruct a robust dated phylogenetic tree for the family. This phylogenetic framework, combined with seed dispersal traits, biome data and biogeographic ranges, allows the reconstruction of the biogeographical history of the group. Results Ancestral range reconstructions suggest a Gondwanan origin (Africa/South America), with several intercontinental dispersals within the family and few transitions between biomes. Relative abundance of fruit dispersal types differed by both continent and biome. However, intercontinental colonizations were only significantly enhanced by water dispersal (drift fruit), and there was no evidence that seed dispersal modes influenced biome shifts. Main Conclusions Our analysis reveals a paradox as drift fruit greatly enhanced dispersal distances at intercontinental scale but did not affect the strong biome conservatism observed.

Low temperature constitutes one of the main barriers to plant distributions, confining many clades to their ancestrally tropical biome. However, recent evidence suggests that transitions from tropical to temperate biomes may be more frequent than previously thought. Here, we study the evolution of cold and frost tolerance in the globally distributed and highly stress-tolerant Salicornieae (Salicornioideae, Amaranthaceae s.l.). We first generate a phylogenetic tree comprising almost all known species (85-90%), using newly generated (n = 106) and published nuclear-ribosomal and plastid sequences. Next, we use geographical occurrence data to document in which clades and geographical regions cold-tolerant species occur and reconstruct how cold tolerance evolved. Finally, we test for correlated evolution between frost tolerance and the annual life form. We find that frost tolerance has evolved independently in up to four Northern Hemisphere lineages but that annuals are no more likely to evolve frost tolerance than perennials, indicating the presence of different strategies for adapting to cold environments. Our findings add to mounting evidence for multiple independent out-of-the-tropics transitions among close relatives of flowering plants and raise new questions about the ecological and physiological mechanism(s) of adaptation to low temperatures in Salicornieae.