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The work was supported by a grant from the Spanish Ministerio de Ciencia e Innovación, PID2020-119163 GB-I00, and SG benefited from a Ramón y Cajal contract (RYC-2014–16608), both funded by MCIN/AEI/10.13039/501100011033., Introduction Materials and methods Results and discussion Data availability References Acknowledgements Funding Author information Ethics declarations Additional information Rights and permissions About this article

Summary While Sherwin J. Carlquist (1930–2021) did not originate the concept of long-distance dispersal and its role in evolution — a major pillar in Darwin’s theory (1859) — he almost single-handedly turned research on dispersal to insular habitats into an empirical and experimental research area. This contribution explains how and why this occurred based on Carlquist’s own papers and personal account, and provides a brief assessment of the historical context of his research on long-distance dispersal. I end on a personal note; in 1981, when I was a graduate student, Carlquist participated in a symposium on ‘Dispersal and Distribution’ in Hamburg, and the paper he gave there on intercontinental dispersal greatly influenced my own work.

The Cucurbitaceae (cucurbit) family consists of about 1,000 species in 95 genera, including many economically important and popular fruit and vegetable crops. During the past several years, reference genomes have been generated for >20 cucurbit species, and variome and transcriptome profiling data have been rapidly accumulated for cucurbits. To efficiently mine, analyze and disseminate these large-scale datasets, we have developed an updated version of Cucurbit Genomics Database. The updated database, CuGenDBv2 (http://cucurbitgenomics.org/v2), currently hosts 34 reference genomes from 27 cucurbit species/subspecies belonging to 10 different genera. Protein-coding genes from these genomes have been comprehensively annotated by comparing their protein sequences to various public protein and domain databases. A novel ‘Genotype’ module has been implemented to facilitate mining and analysis of the functionally annotated variome data including SNPs and small indels from large-scale genome sequencing projects. An updated ‘Expression’ module has been developed to provide a comprehensive gene expression atlas for cucurbits. Furthermore, synteny blocks between any two and within each of the 34 genomes, representing a total of 595 pair-wise genome comparisons, have been identified and can be explored and visualized in the database.

Microscopically dimorphic sex chromosomes in plants are rare, reducing our ability to study them. One difficulty has been the paucity of cultivatable species pairs for cytogenetic, genomic and experimental work. Here, we study the newly recognized sisters Coccinia grandis and Coccinia schimperi, both with large Y chromosomes as we here show for Co. schimperi. We built genetic maps for male and female Co. grandis using a full-sibling family, inferred gene sex-linkage, and, with Co. schimperi transcriptome data, tested whether X- and Y-alleles group by species or by sex. Most sex-linked genes for which we could include outgroups grouped the X- and Y-alleles by species, but some 10% instead grouped the two species' X-alleles. There was no relationship between XY synonymous-site divergences in these genes and gene position on the non-recombining part of the X, suggesting recombination arrest shortly before or after species divergence, here dated to about 3.6 Ma. Coccinia grandis and Co. schimperi are the species pair with the most heteromorphic sex chromosomes in vascular plants (the condition in their sister remains unknown), and future work could use them to study mechanisms of Y chromosome enlargement and parallel degeneration, or to test Haldane's rule about lower hybrid fitness in the heterogametic sex. This article is part of the theme issue ‘Sex determination and sex chromosome evolution in land plants’.

In lineages of allopolyploid origin, sets of homoeologous chromosomes may coexist that differ in gene content and syntenic structure. Presence or absence of genes and microsynteny along chromosomal blocks can serve to differentiate subgenomes and to infer phylogenies. We here apply genome-structural data to infer relationships in an ancient allopolyploid lineage, the walnut family (Juglandaceae), by using seven chromosome-level genomes, two of them newly assembled. Microsynteny and gene-content analyses yield identical topologies that place Platycarya with Engelhardia as did a 1980s morphological-cladistic study. DNA-alignment-based topologies here and in numerous earlier studies instead group Platycarya with Carya and Juglans, perhaps misled by past hybridization. All available data support a hybrid origin of Juglandaceae from extinct or unsampled progenitors nested within, or sister to, Myricaceae. Rhoiptelea chiliantha, sister to all other Juglandaceae, contains proportionally more DNA repair genes and appears to evolve at a rate 2.6- to 3.5-times slower than the remaining species.

PREMISE State‐sponsored weather stations became ubiquitous by the 1880s, yet many old climate data and phenological observations still need to be digitized and made accessible. METHODS We here make available flowering times for 450 species of herbs and shrubs gathered in 1844 by Carl Friedrich Philipp von Martius (1794–1868), director of the Munich Botanical Garden. The data formed part of the world’s third‐oldest phenological monitoring network as we explain in a brief overview of the history of such networks. Using data from one of the world’s oldest continuously functioning weather stations, Hohenpeißenberg, we relate temperature to flowering in three species with short flowering times and herbarium collections made since 1844 within the city’s perimeter, namely, Anemone patens, A. pulsatilla, and Arum maculatum. RESULTS Mean advances in flowering dates were 1.3–2.1 days/decade or 3.2–4.2 days/1°C warming. These advances are in keeping with similar advances in other European herbs during more recent periods. CONCLUSIONS Future studies might use the 1844 flowering data made available here as a source of information on the availability of particular flowers for specialized pollinators including insects looking for oviposition sites, such as the Psychoda flies that become trapped in Arum inflorescences. Another use of Martius’s 1844 data would be their incorporation into larger‐scale analyses of flowering in southern‐central Europe., American Journal of Botany, 108 (4), ISSN:1914-2016, ISSN:0002-9122

Hundreds of land plant lineages have independently evolved separate sexes in either gametophytes (dioicy) or sporophytes (dioecy), but 43% of all dioecious angiosperms are found in just 34 entirely dioecious clades, suggesting that their mode of sex determination evolved a long time ago. Here, we review recent insights on the molecular mechanisms that underlie the evolutionary change from individuals that each produce male and female gametes to individuals specializing in the production of just one type of gamete. The canonical model of sex chromosome evolution in plants predicts that two sex-determining genes will become linked in a sex-determining region (SDR), followed by expanding recombination suppression, chromosome differentiation and, ultimately, degeneration. Experimental work, however, is showing that single genes function as master regulators in model systems, such as the liverwort Marchantia and the angiosperms Diospyros and Populus. In Populus, this type of regulatory function has been demonstrated by genome editing. In other systems, including Actinidia, Asparagus and Vitis, two coinherited factors appear to independently regulate female and male function, yet sex chromosome differentiation has remained low. We discuss the best-understood systems and evolutionary pathways to dioecy, and present a meta-analysis of the sizes and ages of SDRs. We propose that limited sexual conflict explains why most SDRs are small and sex chromosomes remain homomorphic. It appears that models of increasing recombination suppression with age do not apply because selection favours mechanisms in which sex determination depends on minimal differences, keeping it surgically precise.

SummaryAnimal-pollinated plants have to get pollen to a conspecific stigma while protecting it from getting eaten. We provide experimental evidence that touch-sensitive stamens function in (i) enhancing pollen export and (ii) reducing pollen loss to thieves. Stamens of Berberis and Mahonia are inserted between paired nectar glands and when touched by an insect’s tongue rapidly snap forward so that their valvate anthers press pollen on the insect’s tongue or face. We immobilized the stamens in otherwise unmodified flowers and studied pollen transfer in the field and under enclosed conditions. On flowers with immobilized stamens, the commonest bee visitor stayed up to 3.6x longer, yet removed 1.3x fewer pollen grains and deposited 2.1x fewer grains on stigmas per visit. Self-pollen from a single stamen hitting the stigma amounted to 6% of the grains received from single bee visits. Bees discarded pollen passively placed on their bodies, likely because of its berberine content; nectar has no berberine. Syrphid flies fed on both nectar and pollen, taking more when stamens were immobilized. Pollen-tracking experiments in two species showed that mobile-stamen-flowers donate pollen to many more recipients. These results demonstrate another mechanism by which plants simultaneously meter out their pollen and reduce pollen theft.HighlightsStamens that snap forward when triggered by a flower visitor may serve to meter out pollen, scare away pollen thieves, or place pollen more accurately.We tested these hypotheses by experimentally immobilizing all six stamens in numerous flowers of Berberis and Mahonia species in the field and under enclosed conditions.In flowers with immobilized stamens, the commonest bee species stayed up to 3.6x longer, yet removed 1.3x fewer pollen grains and deposited 2.1x fewer grains on stigmas per visit. Mobile stamens exported their pollen to significantly more neighboring flowers.Graphic abstractBehaviour and pollen transfer after flower visitors received a beating on the tongue or in the face by the forward-snapping stamens of Berberis. Stamens only snap forward if their filament basis is touched by an insect tongue.

Animal-pollinated plants have to get pollen to a conspecific stigma while protecting it from getting eaten. Touch-sensitive stamens, which are found in hundreds of flowering plants, are thought to function in enhancing pollen export and reducing its loss, but experimental tests are scarce. Stamens of Berberis and Mahonia are inserted between paired nectar glands and when touched by an insect’s tongue rapidly snap forward so that their valvate anthers press pollen on the insect’s tongue or face. We immobilized the stamens in otherwise unmodified flowers and studied pollen transfer in the field and under enclosed conditions. On flowers with immobilized stamens, the most common bee visitor stayed up to 3.6× longer, yet removed 1.3× fewer pollen grains and deposited 2.1× fewer grains on stigmas per visit. Self-pollen from a single stamen hitting the stigma amounted to 6% of the grains received from single bee visits. Bees discarded pollen passively placed on their bodies, likely because of its berberine content; nectar has no berberine. Syrphid flies fed on both nectar and pollen, taking more when stamens were immobilized. Pollen-tracking experiments in two Berberis species showed that mobile-stamen-flowers donate pollen to many more recipients. These results demonstrate another mechanism by which plants simultaneously meter out their pollen and reduce pollen theft.

In 1917, sex chromosomes in plants were discovered in a liverwort with hetermorphic U and V chromosomes. Such heteromorphy is unexpected because, unlike the XY chromosomes in diploid-dominant plants, in haploid-dominant plants the female U and the male V chromosomes experience largely symmetrical potential recombination environments. Here we use molecular cytogenetics and super-resolution microscopy to study Frullania dilatata, a liverwort with one male and two female sex chromosomes. We applied a pipeline to Illumina sequences to detect abundant types of repetitive DNA and developed FISH probes to microscopically distinguish the sex chromosomes. We also determined the phenotypic population sex ratio because biased ratios have been reported from other liverworts with heteromorphic sex chromosomes. Populations had male-biased sex ratios. The sex chromosomes are monocentric, and of 14 probes studied (eight satellites, five transposable elements and one plastid region), four resulted in unique signals that differentiated the sex chromosomes from the autosomes and from each other. One FISH probe selectively marked the centromeres of both U chromosomes, so we could prove that during meiosis each U chromosome associates with one of the opposite telomeres of the V chromosome, resulting in a head-to-head trivalent. The similarity of the two U chromosomes to each other in size and in their centromere FISH signal positions points to their origin via a non-disjunction event (aneuploidy), which would fit with the general picture of sex chromosomes rarely crossing-over and being prone to suffer from non-disjunction.

Bees require suitably close foraging and nesting sites to minimize travel time and energy expenditure for brood provisioning. Knowing foraging distances in persistent (‘healthy’) populations is therefore crucial for assessing harmful levels of habitat fragmentation. For small bees, such distances are poorly known because of the difficulty of individual tagging and problems with mark-recapture approaches. Using apiarist’s number tags and colour codes, we marked 2689 males and females of four oligolectic and two polylectic species of Osmiini bees (Megachilidae, genera Chelostoma, Heriades, Hoplitis, Osmia) with body lengths of 6 to 15 mm. The work was carried out in 21 ha-large urban garden that harbours at least 106 species of wild bees. Based on 450 re-sightings, mean female flight distances ranged from 73 to 121 m and male distances from 59 to 100 m. These foraging distances suggest that as a rule of thumb, flower strips and nesting sites for supporting small solitary bees should be no further than 150 m apart.

Previous work has shown that among 428 species of bees occurring in Germany, decline or extinction over the past 40 years have been correlated with late-season emergence and restricted habitats, while other factors, such as pollen specialization, body size, nesting sites, and sociality, played no role in models that included a phylogeny of these bees. Doing best are spring-flying, city-dwelling species. Building on these results, we here investigate changes in bee diversity from the 1990s to 2018 at three protected sites within the city perimeter of Munich, focusing on the correlates of flight season (spring or summer), flight duration (in months), and number of habitats (one or two vs. three to six). Local species pools were assessed against the total species pool from 1795 onwards. Twenty years ago, 150 species were present at one or more of the sites, while in 2017/2018, this was true of 188 species, with the increase at two sites being of similar proportion. In two of the three areas, broad habitat use was positively correlated with persistence. Flight season or duration had no statistical effect. These results underscore the function of urban protected sites in bee conservation and imply that summer food shortages, which negatively affect bees in agricultural areas, play no role in urbanized regions so that late-season flight is not an extinction handicap.

AIM: Trees need to avoid frost damage to their young leaves by leafing out after the occurrence of the last frost, yet they also need to start photosynthesis early in the season to achieve sufficient growth. This trade‐off leads to the hypothesis that ‘safety margins’ against spring frost should become shorter, the longer the winter duration, perhaps reaching an asymptotic limit where frost damage would occur in most years. Physiologically, shorter safety margins in high‐latitude ecotypes might be achieved by lower degree‐day requirements for leaf‐out, compared to low‐latitude ecotypes. LOCATION: Europe. TIME PERIOD: 1902–2009. MAJOR TAXA STUDIED: Temperate trees. METHODS: Using herbarium collections of Acer platanoides, Carpinus betulus, Fagus sylvatica and Prunus spinosa made over 108 years at 40° to 60° N latitude, we related historic leaf‐out dates to winter and spring temperatures (chilling and degree‐days), winter duration, and date of last frost occurrence in the relevant years and locations. RESULTS: In all species, frost safety margins decreased towards high‐latitude regions with long winters, with each day increase in winter duration reducing frost safety margins by 0.48 days in Fagus and 0.32–0.21 days in Prunus, Acer and Carpinus. These latitudinal differences correlate with northern ecotypes’ shorter degree‐day requirements for leaf‐out. MAIN CONCLUSIONS: The decline in spring frost safety margins in regions with long winters supports the new hypothesis that species may reach their geographic range limit where they ‘bump up’ against experiencing regular frost injury to their young leaves. Larger datasets are necessary to further corroborate our hypothesis and future efforts should thus be directed toward increasing the latitudinal range of existing phenological databases.

Background and Aims Unlike other nuclear genes in eukaryotes, rDNA genes (5S and 35S loci) are present in numerous copies per cell and, when stained, can therefore provide basic information about genome organization. In tracheophytes (vascular plants), they are usually located on separate chromosomes, the so-called S-type organization. An analysis of 1791 species of land plants suggested that S-type arrays might be ancestral in land plants, while linked (L-type) organization may be derived. However, no outgroup and only a handful of ferns and bryophytes were included. Methods We analysed genome sizes and the distribution of telomere, 5S and 35S rDNA FISH signals in up to 12 monoicous or dioicous species of liverworts from throughout a phylogeny that includes 287 of the 386 currently recognized genera. We also used the phylogeny to plot chromosome numbers and the occurrence of visibly distinct sex chromosomes. Key Results Chromosome numbers are newly reported for the monoicous Lejeunea cavifolia and for females of the dioicous Scapania aequiloba. We detected sex-related differences in the number of rDNA signals in the dioicous Plagiochila asplenioides and Frullania dilatata. In the latter, the presence of two UU chromosomes in females and additional 5S-35S rDNA loci result in a haploid genome 0.2082 pg larger than the male genome; sex-specific genome differences in the other dioicous species were small. Four species have S-type rDNA, while five species have mixed L-S rDNA organization, and transitions may have occurred multiple times, as suggested by rDNA loci not being conserved among closely related species of Pellia. All species shared an Arabidopsis-like telomere motif, and its detection allowed verification of the chromosome number of Radula complanata and chromosome rearrangements in Aneura pinguis and P. asplenioides, the latter also showing sex-specific interstitial telomere repeats. Conclusions The S and L rDNA arrangements appear to have evolved repeatedly within liverworts, even in the same species. Evidence for differential accumulation of rDNA between the sexes so far is limited.

Coriariaceae comprise only Coriaria , a genus of shrubs with nine species in Australasia (but excluding Australia), five in the Himalayas, Taiwan, the Philippines, and Japan, one in the Mediterranean, and one ranging from Patagonia to Mexico. The sister family, Corynocarpaceae, comprises five species of evergreen trees from New Guinea to New Zealand and Australia. This distribution has long fascinated biogeographers as potential support for Wegener's theory of continental drift, with alternative scenarios invoking either Antarctic or Beringian range expansions. Here, we present the discovery of pollen grains from Early Campanian (ca. 82 Mya) deposits in Antarctica, which we describe as Coriaripites goodii sp. nov., and newly generated nuclear and plastid molecular data for most of the family's species and its outgroup. This greatly expands the family's fossil record and is the so far oldest fossil of the order Cucurbitales. We used the phylogeny, new fossil, and an Oligocene flowering branch assigned to a small subclade of Coriaria to generate a chronogram and to study changes in chromosome number, deciduousness, and andromonoecy. Coriaria comprises a Northern (NH) and a Southern Hemisphere (SH) clade that diverged from each other in the Paleocene (ca. 57 Mya), with the SH clade reaching the New World once, through Antarctica, as supported by the fossil pollen. While the SH clade retained perfect flowers and evergreen leaves, the NH clade evolved andromonoecy and deciduousness. Polyploidy occurs in both clades and points to hybridization, matching weak species boundaries throughout the genus.

Background Persian walnut, Juglans regia, occurs naturally from Greece to western China, while its closest relative, the iron walnut, Juglans sigillata, is endemic in southwest China; both species are cultivated for their nuts and wood. Here, we infer their demographic histories and the time and direction of possible hybridization and introgression between them. Results We use whole-genome resequencing data, different population-genetic approaches (PSMC and GONE), and isolation-with-migration models (IMa3) on individuals from Europe, Iran, Kazakhstan, Pakistan, and China. IMa3 analyses indicate that the two species diverged from each other by 0.85 million years ago, with unidirectional gene flow from eastern J. regia and its ancestor into J. sigillata, including the shell-thickness gene. Within J. regia, a western group, located from Europe to Iran, and an eastern group with individuals from northern China, experienced dramatically declining population sizes about 80 generations ago (roughly 2400 to 4000 years), followed by an expansion at about 40 generations, while J. sigillata had a constant population size from about 100 to 20 generations ago, followed by a rapid decline. Conclusions Both J. regia and J. sigillata appear to have suffered sudden population declines during their domestication, suggesting that the bottleneck scenario of plant domestication may well apply in at least some perennial crop species. Introgression from introduced J. regia appears to have played a role in the domestication of J. sigillata.

Predicting the timing of autumn leaf senescence in northern trees remains challenging because the seasonal interplay in the effects of day length, climate, and plant productivity is not well understood. This severely limits our ability to forecast vegetation activity and carbon uptake in temperate and boreal ecosystems. Here we present a new framework for predicting autumn senescence dates based on the idea that day length mediates the effects of climate on autumn phenology, with early-season (pre-solstice) growth and late-season temperatures constituting antagonistic forces. To test these predictions, we used a combination of satellite-derived vegetation productivity across Northern Hemisphere forests, ground-sourced European phenology observations of four widespread tree species, and a climate-manipulation experiment on European beech. Our results reveal important constraints on the late-season carbon uptake potential of northern trees, improving our understanding of vegetation dynamics in response to climate change.

Linnaeus's very first opus, written when he was 22 years old, dealt with the analogy that exists between plants and animals in how they ‘propagate their species’, and a revised version with a plate depicting the union of male and female Mercurialis annua plants became a foundational text on the sexuality of plants. The question how systems with separate males and females have evolved in sedentary organisms that appear ancestrally bisexual has fascinated biologists ever since. The phenomenon, termed dioecy, has important consequences for plant reproductive success and is of commercial interest since it affects seed quality and fruit production. This theme issue presents a series of articles that synthesize and challenge the current understanding of how plants achieve dioecy. The articles deal with a broad set of taxa, including Coccinia , Ginkgo , Mercurialis , Populus , Rumex and Silene , as well as overarching topics, such as the field's terminology, analogies with animal sex determination systems, evolutionary pathways to dioecy, dosage compensation, and the longevity of the two sexes. In this introduction, we focus on four topics, each addressed by several articles from different angles and with different conclusions. Our highlighting of unclear or controversial issues may help future studies to build on the current understanding and to ask new questions that will expand our knowledge of plant sexual systems. This article is part of the theme issue ‘Sex determination and sex chromosome evolution in land plants’.

Microscopically dimorphic sex chromosomes in plants are rare, reducing our ability to study them. One difficulty has been the paucity of cultivatable species pairs for cytogenetic, genomic and experimental work. Here, we study the newly recognized sistersCoccinia grandisandCoccinia schimperi, both with large Y chromosomes as we here show forCo. schimperi. We built genetic maps for male and femaleCo. grandisusing a full-sibling family, inferred gene sex-linkage, and, withCo. schimperitranscriptome data, tested whether X- and Y-alleles group by species or by sex. Most sex-linked genes for which we could include outgroups grouped the X- and Y-alleles by species, but some 10% instead grouped the two species' X-alleles. There was no relationship between XY synonymous-site divergences in these genes and gene position on the non-recombining part of the X, suggesting recombination arrest shortly before or after species divergence, here dated to about 3.6 Ma.Coccinia grandisandCo. schimperiare the species pair with the most heteromorphic sex chromosomes in vascular plants (the condition in their sister remains unknown), and future work could use them to study mechanisms of Y chromosome enlargement and parallel degeneration, or to test Haldane's rule about lower hybrid fitness in the heterogametic sex.This article is part of the theme issue ‘Sex determination and sex chromosome evolution in land plants’.

Motivation Phylogenetic and phylogenomic analyses require multi-gene input files in different formats, but there are few user-friendly programs facilitating the workflow of combining, concatenating or separating, aligning and exploring multi-gene datasets. Results We present Concatenator, a user-friendly GUI-driven program that accepts single-marker and multi-marker DNA sequences in different input formats, including Fasta, Phylip and Nexus, and that outputs concatenated sequences as single-marker or multi-marker Fasta, interleaved nexus or Phylip files, including command files for downstream model selection in IQ-TREE. It includes the option to (re)align markers with MAFFT and produces exploratory trees with FastTree. Although tailored for medium-sized phylogenetic projects, Concatenator is able to process phylogenomic datasets of up to 30 000 markers. Availability and implementation Concatenator is written in Python, with C extensions for MAFFT and FastTree. Compiled stand-alone executables of Concatenator for MS Windows and Mac OS along with a detailed manual can be downloaded from www.itaxotools.org; the source code is openly available on GitHub (https://github.com/iTaxoTools/ConcatenatorGui).

In a commentary on our paper (Renner et al., Oecologia 195:825–831, 2021), Harder and Miksha lay out why they think that our finding of higher honeybee abundances reducing wild bee abundances in an urban botanical garden is not statistically supported. Here, we explain the statistical test provided in our paper, which took advantage of a natural experiment offered by 2019 being a poorer year for bee keeping than 2020.

Although hybridization plays a large role in speciation, some unknown fraction of hybrid individuals never reproduces, instead remaining as genetic dead-ends. We investigated a morphologically distinct and culturally important Chinese walnut, Juglans hopeiensis, suspected to have arisen from hybridization of Persian walnut (J. regia) with Asian butternuts (J. cathayensis, J. mandshurica, and hybrids between J. cathayensis and J. mandshurica). Based on 151 whole-genome sequences of the relevant taxa, we discovered that all J. hopeiensis individuals are first-generation hybrids, with the time for the onset of gene flow estimated as 370,000 years, implying both strong postzygotic barriers and the presence of J. regia in China by that time. Six inversion regions enriched for genes associated with pollen germination and pollen tube growth may be involved in the postzygotic barriers that prevent sexual reproduction in the hybrids. Despite its long-recurrent origination and distinct traits, J. hopeiensis does not appear on the way to speciation.

When flowers provide nectar, they can count on visits, and potential pollen transport, by many kinds of nectar-drinking animals. Yet, some flowers instead offer fatty oils that certain specialized bees gather with their forelegs. A recent study reveals how such a switch occurred and may have contributed to the formation of a new species.

Squamellaria is a genus of epiphytic Rubiaceae which has been domesticated by ants on the Fiji islands. One species is on the verge of extinction.

Persian walnut (Juglans regia) is cultivated worldwide for its high-quality wood and nuts, but its origin has remained mysterious because in phylogenies it occupies an unresolved position between American black walnuts and Asian butternuts. Equally unclear is the origin of the only American butternut, J. cinerea. We resequenced the whole genome of 80 individuals from 19 of the 22 species of Juglans and assembled the genome of its relatives Pterocarya stenoptera and Platycarya strobilacea. Using phylogenetic-network analysis of single-copy nuclear genes, genome-wide site pattern probabilities, and Approximate Bayesian Computation, we discovered that J. regia (and its landrace J. sigillata) arose as a hybrid between the American and the Asian lineages and that J. cinerea resulted from massive introgression from an immigrating Asian butternut into the genome of an American black walnut. Approximate Bayesian Computation modeling placed the hybrid origin in the late Pliocene, ∼3.45 My, with both parental lineages since having gone extinct in Europe.

Cytogenetic studies in bryophytes have been limited by the difficulty of obtaining sufficient dividing nuclei and by the absence of modern protocols. The technical difficulties stem from the plants’ small size and lack of roots and pollen mother cells, the main sources of cells in division in vascular plants. In bryophytes instead, tiny sporophytes, antheridia, or phyllid meristems must be used to obtain meiotic or mitotic chromosome spreads. We here describe the preparation of such spreads from phyllids, antheridia, and sporophytes in several species of liverworts and compare available protocols with or without prefixation treatments. We also provide illustrated step-by-step instructions. The three prefixation agents (including colchicine) that we tested failed to improve synchronization of cell divisions. Young sporophytes were the best source of diploid synchronized cells, while antheridia were the best source of haploid cells. For meiotic nuclei, a short fixation of capsule tissue at the right developmental stage with 45% acetic acid sufficed to conserve the DNA for cytological investigations, while for mitotic nuclei, fixation in 3:1 ethanol/glacial acetic acid for a longer period (4–24 h) worked well.

Wild relatives or progenitors of crops are important resources for breeding and for understanding domestication. Identifying them, however, is difficult because of extinction, hybridization, and the challenge of distinguishing them from feral forms. Here, we use collection-based systematics, iconography, and resequenced accessions of Citrullus lanatus and other species of Citrullus to search for the potential progenitor of the domesticated watermelon. A Sudanese form with nonbitter whitish pulp, known as the Kordofan melon (C. lanatus subsp. cordophanus), appears to be the closest relative of domesticated watermelons and a possible progenitor, consistent with newly interpreted Egyptian tomb paintings that suggest that the watermelon may have been consumed in the Nile Valley as a dessert by 4360 BP. To gain insights into the genetic changes that occurred from the progenitor to the domesticated watermelon, we assembled and annotated the genome of a Kordofan melon at the chromosome level, using a combination of Pacific Biosciences and Illumina sequencing as well as Hi-C mapping technologies. The genetic signature of bitterness loss is present in the Kordofan melon genome, but the red fruit flesh color only became fixed in the domesticated watermelon. We detected 15,824 genome structural variants (SVs) between the Kordofan melon and a typical modern cultivar, "97103," and mapping the SVs in over 400 Citrullus accessions revealed shifts in allelic frequencies, suggesting that fruit sweetness has gradually increased over the course of watermelon domestication. That a likely progenitor of the watermelon still exists in Sudan has implications for targeted modern breeding efforts.

While powerful and user-friendly software suites exist for phylogenetics, and an impressive cybertaxomic infrastructure of online species databases has been set up in the past two decades, software targeted explicitly at facilitating alpha-taxonomic work, i.e., delimiting and diagnosing species, is still in its infancy. Here we present a project to develop a bioinformatic toolkit for taxonomy, based on open-source Python code, including tools focusing on species delimitation and diagnosis and centered around specimen identifiers. At the core of iTaxoTools is user-friendliness, with numerous autocorrect options for data files and with intuitive graphical user interfaces. Assembled standalone executables for all tools or a suite of tools with a launcher window will be distributed for Windows, Linux, and Mac OS systems, and in the future also implemented on a web server. The initial version (iTaxoTools 0.1) distributed with this paper (https://github.com/iTaxoTools/iTaxoTools-Executables) contains graphical user interface (GUI) versions of six species delimitation programs (ABGD, ASAP, DELINEATE, GMYC, PTP, tr2) and a simple threshold-clustering delimitation tool. There are also new Python implementations of existing algorithms, including tools to compute pairwise DNA distances, ultrametric time trees based on non-parametric rate smoothing, species-diagnostic nucleotide positions, and standard morphometric analyses. Other utilities convert among different formats of molecular sequences, geographical coordinates, and units; merge, split and prune sequence files, tables and species partition files; and perform simple statistical tests. As a future perspective, we envisage iTaxoTools to become part of a bioinformatic pipeline for next-generation taxonomy that accelerates the inventory of life while maintaining high-quality species hypotheses. The open source code and binaries of all tools are available from Github (https://github.com/iTaxoTools) and further information from the website (http://itaxotools.org).

Our study showed that increases in seasonal productivity drive earlier autumn senescence of temperate trees. Norby argues that this finding is contradicted by observations from free-air CO 2 enrichment (FACE) experiments, where elevated CO 2 has been found to delay senescence in some cases. We provide a detailed answer showing that the results from FACE studies are in agreement with our conclusions.

Limits to the growing season The length of the growing season in temperate forests has been increasing under recent climate change because of earlier leaf emergence and later leaf senescence. However, Zani et al. show that this trend might be reversed as increasing photosynthetic productivity begins to drive earlier autumn leaf senescence (see the Perspective by Rollinson). Using a combination of experimental, observational, and modeling studies based on European forest trees, the researchers conclude that leaf senescence will advance by 3 to 6 days by the end of the 21st century rather than lengthening by 1 to 3 weeks as current phenological models have predicted. In turn, this predicted phenological pattern will limit the capacity of temperate forests to mitigate climate change through carbon uptake. Science , this issue p. 1066 ; see also p. 1030

Natural history collections are leading successful large-scale projects of specimen digitization (images, metadata, DNA barcodes), thereby transforming taxonomy into a big data science. Yet, little effort has been directed towards safeguarding and subsequently mobilizing the considerable amount of original data generated during the process of naming 15,000–20,000 species every year. From the perspective of alpha-taxonomists, we provide a review of the properties and diversity of taxonomic data, assess their volume and use, and establish criteria for optimizing data repositories. We surveyed 4113 alpha-taxonomic studies in representative journals for 2002, 2010, and 2018, and found an increasing yet comparatively limited use of molecular data in species diagnosis and description. In 2018, of the 2661 papers published in specialized taxonomic journals, molecular data were widely used in mycology (94%), regularly in vertebrates (53%), but rarely in botany (15%) and entomology (10%). Images play an important role in taxonomic research on all taxa, with photographs used in >80% and drawings in 58% of the surveyed papers. The use of omics (high-throughput) approaches or 3D documentation is still rare. Improved archiving strategies for metabarcoding consensus reads, genome and transcriptome assemblies, and chemical and metabolomic data could help to mobilize the wealth of high-throughput data for alpha-taxonomy. Because long-term—ideally perpetual—data storage is of particular importance for taxonomy, energy footprint reduction via less storage-demanding formats is a priority if their information content suffices for the purpose of taxonomic studies. Whereas taxonomic assignments are quasifacts for most biological disciplines, they remain hypotheses pertaining to evolutionary relatedness of individuals for alpha-taxonomy. For this reason, an improved reuse of taxonomic data, including machine-learning-based species identification and delimitation pipelines, requires a cyberspecimen approach—linking data via unique specimen identifiers, and thereby making them findable, accessible, interoperable, and reusable for taxonomic research. This poses both qualitative challenges to adapt the existing infrastructure of data centers to a specimen-centered concept and quantitative challenges to host and connect an estimated \documentclass[12pt]{minimal} \usepackage{amsmath} \usepackage{wasysym} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{amsbsy} \usepackage{upgreek} \usepackage{mathrsfs} \setlength{\oddsidemargin}{-69pt} \begin{document} }{}$ \le $\end{document}2 million images produced per year by alpha-taxonomic studies, plus many millions of images from digitization campaigns. Of the 30,000–40,000 taxonomists globally, many are thought to be nonprofessionals, and capturing the data for online storage and reuse therefore requires low-complexity submission workflows and cost-free repository use. Expert taxonomists are the main stakeholders able to identify and formalize the needs of the discipline; their expertise is needed to implement the envisioned virtual collections of cyberspecimens. [Big data; cyberspecimen; new species; omics; repositories; specimen identifier; taxonomy; taxonomic data.]

Hundreds of land plant lineages have independently evolved separate sexes in either gametophytes (dioicy) or sporophytes (dioecy), but 43% of all dioecious angiosperms are found in just 34 entirely dioecious clades, suggesting that their mode of sex determination evolved a long time ago. Here, we review recent insights on the molecular mechanisms that underlie the evolutionary change from individuals that each produce male and female gametes to individuals specializing in the production of just one type of gamete. The canonical model of sex chromosome evolution in plants predicts that two sex-determining genes will become linked in a sex-determining region (SDR), followed by expanding recombination suppression, chromosome differentiation and, ultimately, degeneration. Experimental work, however, is showing that single genes function as master regulators in model systems, such as the liverwort Marchantia and the angiosperms Diospyros and Populus. In Populus, this type of regulatory function has been demonstrated by genome editing. In other systems, including Actinidia, Asparagus and Vitis, two coinherited factors appear to independently regulate female and male function, yet sex chromosome differentiation has remained low. We discuss the best-understood systems and evolutionary pathways to dioecy, and present a meta-analysis of the sizes and ages of SDRs. We propose that limited sexual conflict explains why most SDRs are small and sex chromosomes remain homomorphic. It appears that models of increasing recombination suppression with age do not apply because selection favours mechanisms in which sex determination depends on minimal differences, keeping it surgically precise.

Some ~15.000 angiosperms are dioecious, but mechanisms of sex determination in plants remain poorly understood. In particular, how Y chromosomes evolve and degenerate, and whether dosage compensation evolves as a response, are matters of debate. Here we focus on Coccinia grandis, a dioecious cucurbit with the highest level of X/Y heteromorphy recorded so far. We identified sex-linked genes using RNA sequences from a cross and a model-based method termed SEX-DETector. Parents and F1 individuals were genotyped and the transmission patterns of SNPs then analysed. In the >1300 sex-linked genes studied, X-Y divergence was 0.13 - 0.17, and substantial Y degeneration is implied by an average Y/X expression ratio of 0.63 and an inferred gene loss on the Y of ~40%. We also found reduced Y gene expression being compensated by elevated expression of corresponding genes on the X and an excess of sex-biased genes on the sex chromosomes. Molecular evolution of sex-linked genes in C. grandis is thus comparable to that in Silene latifolia, another dioecious plant with a strongly heteromorphic XY system, and cucurbits are the fourth plant family in which dosage compensation is described, suggesting it might be common in plants.

Ongoing global warming is causing phenological shifts that affect photosynthesis and growth rates in temperate woody species. However, the effects of seasonally uneven climate warming-as is occurring in much of Europe, where the winter/spring months are warming twice as fast than the summer/autumn months-on autumn growth cessation (completion of overwintering buds) and leaf senescence, and possible carry-over effects between phenophases, remain under-investigated. We conducted experiments in which we exposed saplings of canopy and understory species to 4 °C warming in winter/spring, summer/autumn, or all year to disentangle how the timing of bud break, bud set completion, and leaf senescence is affected by seasonally uneven warming. All-year warming led to significantly delayed leaf senescence, but advanced bud set completion; summer/autumn warming only delayed leaf senescence; and winter/spring warming advanced both bud set and senescence. The non-parallel effects of warming on bud completion and leaf senescence show that leaf senescence alone is an inadequate proxy for autumn growth cessation in trees and counterintuitively suggest that continued uneven seasonal warming will advance cessation of primary growth in autumn, even when leaf senescence is delayed. Phenological responses to warming treatments (earlier spring onset, later autumn senescence) were more than twice as high in understory species than in canopy species, which can partly be explained by the absence of carry-over effects among phenophases in the former group. This underscores the need to consider differences among plant functional types when forecasting the future behaviour of ecosystems.

Orchids are globally distributed, a feature often attributed to their tiny dustlike seeds. They were ancestrally terrestrial but in the Eocene expanded into tree canopies, with some lineages later returning to the ground, providing an evolutionarily replicated system. Because seeds are released closer to the ground in terrestrial species than in epiphytic ones, seed traits in terrestrials may have been under selective pressure to increase seed dispersal efficiency. In this study, we test the expectations that seed airspace-a trait known to increase seed flotation time in the air-is (i) larger in terrestrial lineages and (ii) has increased following secondary returns to a terrestrial habit. We quantified and scored 20 seed traits in 121 species and carried out phylogenetically informed analyses. Results strongly support both expectations, suggesting that aerodynamic traits even in dust seeds are under selection to increase dispersal ability, following shifts in average release heights correlated with changes in habit.

Phenological mismatch results when interacting species change the timing of regularly repeated phases in their life cycles at different rates. We review whether this continuously ongoing phenomenon, also known as trophic asynchrony, is becoming more common under ongoing rapid climate change. In antagonistic trophic interactions, any mismatch will have negative impacts for only one of the species, whereas in mutualistic interactions, both partners are expected to suffer. Trophic mismatch is therefore expected to last for evolutionarily short periods, perhaps only a few seasons, adding to the difficulty of attributing it to climate change, which requires long-term data. So far, the prediction that diverging phenologies linked to climate change will cause mismatch is most clearly met in antagonistic interactions at high latitudes in the Artic. There is limited evidence of phenological mismatch in mutualistic interactions, possibly because of strong selection on mutualists to have co-adapted phenological strategies. The study of individual plasticity, population variation, and the genetic bases for phenological strategies is in its infancy. Recent work on woody plants revealed the large imprint of historic climate change on temperature, chilling, and day-length thresholds used by different species to synchronize their phenophases, which in the Northern Hemisphere has led to biogeographic phenological regions in which long-lived plants have adapted to particular interannual and intermillennial amplitudes of climate change.

Reconstruction of palaeobiomes, ancient communities that exhibit a physiognomic and functional structure controlled by their environment, depends on proxies from different disciplines. Based on terrestrial mammal fossils, the late Miocene vegetation from China to the eastern Mediterranean and East Africa has been reconstructed as a single cohesive biome with increasingly arid conditions, with modern African savannahs the surviving remnant. Here, we test this reconstruction using plant fossils spanning 14-4 million years ago from sites in Greece, Bulgaria, Turkey, the Tian Shan Mountains and Baode County in China, and East Africa. The western Eurasian sites had a continuous forest cover of deciduous or evergreen angiosperms and gymnosperms, with 15% of 1,602 fossil occurrences representing conifers, which were present at all but one of the sites. Raup-Crick analyses reveal high floristic similarity between coeval eastern Mediterranean and Chinese sites, and low similarity between Eurasian and African sites. The disagreement between plant-based reconstructions, which imply that late Miocene western Eurasia was covered by evergreen needleleaf forests and mixed forests, and mammal-based reconstructions, which imply a savannah biome, throws into doubt the approach of inferring Miocene precipitation and open savannah habitats solely from mammalian dental traits. Organismal communities are constantly changing in their species composition, and neither animal nor plant traits by themselves are sufficient to infer entire ancient biomes. The plant fossil record, however, unambiguously rejects the existence of a cohesive savannah biome from eastern Asia to northeast Africa.