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Dactylorhiza sambucina (L.) Soo is a polycarpic perennial herb occurring in the Central European, Eastern European, and Balkan floristic provinces. At the European scale, the IUCN considers it a species of “least concern”. This paper reviews the taxonomic status, morphology, distribution, habitat requirements, mycorrhizal associations, and life cycle of D. sambucina , with special emphasis on its reproduction. We also summarize information on chromosome numbers and genetic variation. Our data from 12 years of monitoring D. sambucina in the Czech Republic show that three to four leaves have to be produced prior to flowering; plants with five and more leaves flower regularly. Juvenile plants near adult plants suggest recruitment from seeds. About 20% of our 450 monitored plants underwent dormancy (failure of mature plants to produce above-ground parts in one or more growing seasons), the maximum duration being eight years. After reappearance, these individuals were usually sterile for the next year. Mortality was highest (24%) at the seedling stage. Regarding the purple/yellow flower colour polymorphism that characterizes D. sambucina , we found no correlation between morph frequency and soil properties (pH, calcium content), population density, or altitude above sea level.

It is plausible that specialized ecological interactions constrain geographic ranges. We address this question in neotropical bees, Centris and Epicharis, that collect oils from flowers of Calceolariaceae, Iridaceae, Krameriaceae, Malpighiaceae, Plantaginaceae, or Solanaceae, with different species exploiting between one and five of these families, which either have epithelial oil glands or hair fields. We plotted the level of oil-host specialization on a clock-dated phylogeny for 22 of the 35 species of Epicharis and 72 of the 230 species of Centris (genera that are not sister genera) and calculated geographic ranges (km(2) ) for 23 bee species based on collection data from museum specimens. Of the oil-offering plants, the Malpighiaceae date to the Upper Cretaceous, whereas the other five families are progressively younger. The stem and crown groups of the two bee genera date to the Cretaceous, Eocene, and Oligocene. Shifts between oil hosts from different families are common in Centris, but absent in Epicharis, and the direction is from flowers with epithelial oil glands to flowers with oil hairs, canalized by bees' oil-collecting apparatuses, suitable for piercing epithelia or mopping oil from hair fields. With the current data, a link between host specialization and geographic range size could not be detected.

Summary Experimental data on the perception of day length and temperature in dormant temperate zone trees are surprisingly scarce. In order to investigate when and where these environmental signals are perceived, we carried out bagging experiments in which buds on branches of Fagus sylvatica, Aesculus hippocastanum and Picea abies trees were exposed to natural light increase or kept at constant 8-h days from December until June. Parallel experiments used twigs cut from the same trees, harvesting treated and control twigs seven times and then exposing them to 8- or 16-h days in a glasshouse. Under 8-h days, budburst in Fagus outdoors was delayed by 41 d and in Aesculus by 4 d; in Picea, day length had no effect. Buds on nearby branches reacted autonomously, and leaf primordia only reacted to light cues in late dormancy after accumulating warm days. Experiments applying different wavelength spectra and high-resolution spectrometry to buds indicate a phytochrome-mediated photoperiod control. By demonstrating local photoperiodic control of buds, revealing the time when these signals are perceived, and showing the interplay between photoperiod and chilling, this study contributes to improved modelling of the impact of climate warming on photosensitive species.

International audience; AbstractGreen roofs, which have become mandatory on new flat-topped buildings in many cities, increase habitat connectivity for wildlife and have contributed to a boom in urban beekeeping. The ecological benefits or risks of green roofs for wild bees (bee species other than the domesticated honey bee, Apis mellifera), however, have not been comprehensively analyzed. We therefore reviewed studies on insects caught on green roofs in Asia, Europe, or North America between 1992 and early 2017 and extracted information on wild bees. The resulting species list includes 236 Apidae identified in 35 studies, with thermophilic species probably overrepresented because roofs provide warm and dry habitats. The percentage of cavity-nesting bees on roofs is higher than that on nearby ground, while the percentage of pollen specialists is lower. Data are almost completely lacking on the reproductive success of bees on green roofs, the effect of roof age on bee diversity, and the genetic or demographic benefits of increased habitat connectivity. Our list of the bee species so far reported on green roofs will help in the selection and implementation of suitable soils, nesting aids, and plantings.

Differing from most animals, plants have an indeterminate body plan, allowing them to add new body parts throughout their lifetime. The realized modular construction of a plant is the result of endogenous processes and exogenous constraints. Plant architectural analysis provides a holistic approach to whole-plant development by disentangling endogenous processes from exogenous constraints. Such analyses have focused more on trees than herbs. We here report a rare growth form discovered in the tropical herb Centella asiatica (Apiaceae) using micro-computed X-ray tomography to understand organogenesis in the meristem. Seeds of C. asiatica germinate to give rise to an orthotropic monochasium (sympodium) with terminal inflorescences. A stolon subsequently emerges from the oldest leaf axils of the sympodium. The primary growth of stolons is indeterminate (monopodial), and they only bear scale leaves. The stolon not only produces new leafy sympodial shoots at each node but also an axillary inflorescence. To place this growth form in context, we illustrate all architectural models so far reported from herbs. Our study highlights the complexity of herb architectures and illustrates the use of a novel methodology to decipher plant branching modes.

We analysed the geographical origins and divergence times of the West Indian hummingbirds, using a large clock-dated phylogeny that included 14 of the 15 West Indian species and statistical biogeographical reconstruction. We also compiled a list of 101 West Indian plant species with hummingbird-adapted flowers (90 of them endemic) and dated the most species-rich genera or tribes, with together 41 hummingbird-dependent species, namely Cestrum (seven spp.), Charianthus (six spp.), Gesnerieae (75 species, c. 14 of them hummingbird-pollinated), Passiflora (ten species, one return to bat-pollination) and Poitea (five spp.), to relate their ages to those of the bird species. Results imply that hummingbirds colonized the West Indies at least five times, from 6.6 Mya onwards, coming from South and Central America, and that there are five pairs of sister species that originated within the region. The oldest of the dated plant groups diversified 9.1, 8.5, and 5.4 Mya, simultaneous with or slightly before the extant West Indian bird radiations. The time frame of the coevolved bird/flower mutualisms obtained here resembles that recently inferred for North America, namely 5–9 Mya. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114, 848–859.

SummaryType specimens are permanently preserved biological specimens that fix the usage of species names. This method became widespread from 1935 onwards and is now obligatory. We used DNA sequencing of types and more recent collections of wild and cultivated melons to reconstruct the evolutionary history of the genus Citrullus and the correct names for its species. We discovered that the type specimen of the name Citrullus lanatus, prepared by a Linnaean collector in South Africa in 1773, is not the species now thought of as watermelon. Instead, it is a representative of another species that is sister to C. ecirrhosus, a tendril-less South African endemic. The closest relative of the watermelon instead is a West African species. Our nuclear and plastid data furthermore reveal that there are seven species of Citrullus, not four as assumed. Our study implies that sweet watermelon originates from West, not southern Africa as previously believed, and that the South African citron melon has been independently domesticated. These findings affect and explain numerous studies on the origin of these two crops that led to contradictory results because of the erroneous merging of several distinct species.

SummaryAnt-plant symbioses involve over 110 ant species in five subfamilies that are facultative or obligate occupants of stem, leaf or root domatia formed by hundreds of ant-plant species. The phylogenetic distribution and geological ages of these associations, and the frequency of gains or losses of domatium, are largely unknown. We compiled an up-to-date list of ant domatium-bearing plants, estimated their probable true number from model-based statistical inference, generated dated phylogenies that include c. 50% of ant-plant lineages, and traced the occurrence of domatia and extrafloral nectaries on a 1181-species tree, using likelihood and Bayesian methods. We found 681 vascular plants with domatia (159 genera in 50 families) resulting from minimally 158 inferred domatium origins and 43 secondary losses over the last 19 Myr. The oldest African ant-plant symbioses are younger than those in Australasia and the Neotropics. The best statistical model suggests that the true number of myrmecophytes may approach 1140 species. The phylogenetic distribution of ant-plants shows that domatia evolved from a range of pre-adapted morphological structures and have been lost frequently, suggesting that domatia have no generalizable effect on diversification. The Miocene origin of ant-plant symbioses is consistent with inferred changes in diet and behaviour during ant evolution.

Many epiphytic plants have associated with ants to gain nutrients. Here, we report a novel type of ant–plant symbiosis in Fiji where one ant species actively and exclusively plants the seeds and fertilizes the seedlings of six species of Squamellaria (Rubiaceae). Comparison with related facultative ant plants suggests that such farming plays a key role in mutualism stability by mitigating the critical re-establishment step. Mutualisms are common in nature. In Fiji, a species of ant selects, disperses and fertilizes an epiphytic plant in an exclusive symbiosis. This represents a novel example of plant farming by ants.

Mutualisms could be evolutionarily unstable, with changes in partner abundances or in the spatial context of interactions potentially promoting their dissolution. We test this prediction using the defense mutualisms between species of the Neotropical genus Cecropia and Azteca ants. A new, multigene phylogeny with representatives of all five genera of Cecropieae (most of them from the Neotropics) and half of the 61 species of Cecropia shows the West African endemic Musanga (2 spp.) as sister to Cecropia, implying dispersal from the Neotropics to Africa, with a molecular clock suggesting that this occurred about 23 Mya. Cecropia, a genus of neotropical pioneer trees, started diversifying ca. 8 Mya. We infer a single origin of specialized symbiosis with Azteca within Cecropia, eight complete losses of this symbiosis, and a potential partner shift involving the replacement of Azteca by Neoponera luteola ants. Niche space modeling based on geo-referenced occurrences of over 9000 collections representing 58 of the 61 species of Cecropia, together with several comparative analyses, implies that mutualism loss is concentrated at high altitudes and on Caribbean islands, with the surprisingly frequent breakdowns potentially facilitated by low species-specificity of interacting Cecropia and Azteca mutualists.

Background and Aims Resin is a defence against herbivores and a floral reward in a few African and South American species whose bee pollinators collect it for nest construction. Here we describe a new role for floral resin from the Asian genus Kadsura (Schisandraceae). Kadsura tepals tightly cover a globe formed by carpels (in females) or near-fused stamens with fleshy connectives (in male flowers of most, but not all species). Methods We carried out field observations at four sites in China and used pollinator behavioural assays, chemical analyses and time-calibrated insect and plant phylogenies to investigate the specificity of the interactions and their relationship to floral structure. Key Results Nocturnal resin midges (Resseliella, Cecidomyiidae) walk around on the flowers’ sexual organs to oviposit, thereby transferring pollen and wounding tissues. The larvae then develop in resin-filled chambers. Male and female floral scents are dominated by α-pinene, while the resinous exudate is dominated by caryophyllene. As revealed by barcoding of multiple midge larvae per flower species, the mutualisms are species specific and appear to have evolved over the past 6–9 million years. Conclusions Resin feeding, not pollen or ovule feeding, by midge larvae explains the abundant Kadsura exudates, highlighting the poorly known world of nocturnal flower–fly interactions.

Mutualisms that involve symbioses among specialized partners may be more stable than mutualisms among generalists, and theoretical models predict that in many mutualisms, partners exert reciprocal stabilizing selection on traits directly involved in the interaction. A corollary is that mutualism breakdown should increase morphological rates of evolution. We here use the largest ant-plant clade (Hydnophytinae), with different levels of specialization for mutualistic ant symbionts, to study the ecological context of mutualism breakdown and the response of a key symbiosis-related trait, domatium entrance hole size, which filters symbionts by size. Our analyses support three predictions from mutualism theory. First, all 12 losses apparently only occur from a generalist symbiotic state. Second, mutualism losses occurred where symbionts are scarce, in our system at high altitudes. Third, domatium entrance hole size barely changes in specialized symbiotic species, but evolves rapidly once symbiosis with ants has broken down, with a "morphorate map" revealing that hotspots of entrance hole evolution are clustered in high-altitude areas. Our study reveals that mutualistic strategy profoundly affects the pace of morphological change in traits involved in the interaction and suggests that shifts in partners' relative abundances may frequently drive reversions of generalist mutualisms to autonomy.

This special featureresults from the symposium ‘Ants 2016: ant interactions with their biotic environments’ held in Munich in May 2016 and deals with the interactions between ants and other insects, plants, microbes and fungi, studied at micro- and macroevolutionary levels with a wide range of approaches, from field ecology to next-generation sequencing, chemical ecology and molecular genetics. In this paper, we review key aspects of these biotic interactions to provide background information for the papers of this special feature. After listing the major types of biotic interactions that ants engage in, we present a brief overview of ant/ant communication, ant/plant interactions, ant/fungus symbioses, and recent insights about ants and their endosymbionts. Using a large molecular clock-dated Formicidae phylogeny, we map the evolutionary origins of different ant clades' interactions with plants, fungi and hemiptera. Ants' biotic interactions provide ideal systems to address fundamental ecological and evolutionary questions about mutualism, coevolution, adaptation and animal communication.

The length of the vegetation period (LVP), which is the time between leaf-out and leaf senescence, affects numerous ecosystem functions, including biogeochemical cycles and interspecific interactions. The evolutionary mechanisms determining LVP, however, are poorly understood, and thus, it is unknown whether innate LVPs differ between eastern North American (ENA), European and East Asian species. Here we monitored LVP in 2014–2015 in 396 Northern Hemisphere woody species grown in a common garden. We found that ENA species, under the same conditions, have three weeks (11%) shorter vegetation periods than their European and East Asian relatives, because their leaves flushed 9 ± 4 and 13 ± 4 days later and senesced 9 ± 4 and 11 ± 4 days earlier. LVPs of species introduced from Eurasia into ENA are therefore longer than those of native species, suggesting that the spread of non-natives might alter seasonal forest productivity in ENA. LVP between naturalized invasive and non-invasive species, however, did not differ, rejecting the common assumption that longer leaf presentation generally fosters invasive success. A likely explanation for the shorter LVP of ENA species is that region’s uniquely high inter-annual temperature variation. These results highlight the footprint of regional climate history, which will affect forest response to climate change. Comparing 396 Northern Hemisphere woody plant species grown under identical conditions in a common garden, the authors show that eastern North American species have 11% shorter vegetation periods than their European and East Asian relatives.

Combining molecular cytogenetics and phylogenetic modelling of chromosome number change can shed light on the types of evolutionary changes that may explain the haploid numbers observed today. Applied to the monocot family Araceae, with chromosome numbers of 2n = 8 to 2n = 160, this type of approach has suggested that descending dysploidy has played a larger role than polyploidy in the evolution of the current chromosome numbers. To test this, we carried out molecular cytogenetic analyses in 14 species from 11 genera, using probes for telomere repeats, 5S rDNA and 45S rDNA and a plastid phylogenetic tree covering the 118 genera of the family, many with multiple species. We obtained new chromosome counts for six species, modelled chromosome number evolution using all available counts for the family and carried out fluorescence in situ hybridization with three probes (5S rDNA, 45S rDNA and Arabidopsis-like telomeres) on 14 species with 2n = 14 to 2n = 60. The ancestral state reconstruction provides support for a large role of descending dysploidy in Araceae, and interstitial telomere repeats (ITRs) were detected in Anthurium leuconerum, A. wendlingeri and Spathyphyllum tenerum, all with 2n = 30. The number of ITR signals in Anthurium (up to 12) is the highest so far reported in angiosperms, and the large repeats located in the pericentromeric regions of A. wendlingeri are of a type previously reported only from the gymnosperms Cycas and Pinus. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177, 15–26.

A major concern in molecular clock dating is how to use information from the fossil record to calibrate genetic distances from DNA sequences. Here we apply three Bayesian dating methods that differ in how calibration is achieved— "node dating" (ND) in BEAST, "total evidence" (TE) dating in MrBayes, and the "fossilized birth-death" (FBD) in FDPPDiv— to infer divergence times in the royal ferns. Osmundaceae have 16-17 species in four genera, two mainly in the Northern Hemisphere and two in South Africa and Australasia; they are the sister clade to the remaining leptosporangiate ferns. Their fossil record consists of at least 150 species in ∼17 genera. For ND, we used the five oldest fossils, whereas for TE and FBD dating, which do not require forcing fossils to nodes and thus can use more fossils, we included up to 36 rhizomes and frond compression/impression fossils, which for TE dating were scored for 33 morphological characters. We also subsampled 10%, 25%, and 50% of the 36 fossils to assess model sensitivity. FBD-derived divergence ages were generally greater than those inferred from ND; two of seven TE-derived ages agreed with FBD-obtained ages, the others were much younger or much older than ND or FBD ages. We prefer the FBD-derived ages because they best fit the Osmundales fossil record (including Triassic fossils not used in our study). Under the preferred model, the clade encompassing extant Osmundaceae (and many fossils) dates to the latest Paleozoic to Early Triassic; divergences of the extant species occurred during the Neogene. Under the assumption of constant speciation and extinction rates, the FBD approach yielded speciation and extinction rates that overlapped those obtained from just neontological data. However, FBD estimates of speciation and extinction are sensitive to violations in the assumption of continuous fossil sampling; therefore, these estimates should be treated with caution. (Bayesian inference; fossilized birth-death dating; molecular clock calibration; node dating; total evidence dating; royal ferns.)

Gene trees of holoparasitic plants usually show distinctly longer branch lengths than seen in photosynthetic closest relatives. Such substitution rate jumps have made it difficult to infer the absolute divergence times of parasites. An additional problem is that parasite clades often lack a fossil record. Using nuclear and mitochondrial DNA sequences of Apodanthaceae, a worldwide family of endoparasites living inside Fabaceae and Salicaceae, we compared several dating approaches: (i) an uncorrelated lognormal (UCLN) model calibrated with outgroup fossils, (ii) ages of host lineages as a maximal age in an UCLN model, (iii) user-assigned local clocks, and (iv) outgroup-fossil-calibrated random local clocks (RLC) with varying prior probabilities on the number of permitted rate changes (RLCu and RLCp models), a variable that has never been explored. The resulting dated phylogenies include all 10 species of the family, three in Australia, one in Iran, one in Africa, and the remainder in the Americas. All clock models infer a drastic rate jump between nonparasitic outgroups and Apodanthaceae, but since they distribute the rate heterogeneity differently, they result in much-different age estimates. Bayes factors using path and stepping-stone sampling indicated that the RLCp model fit poorly, while for matR, topologically unconstrained RLCu and UCLN models did not differ significantly and for 18S, the UCLN model was preferred. Under the equally well fitting models, the Apodanthaceae appear to be a relatively old clade, with a stem age falling between 65 and 81 my, the divergence of Apodanthes from Pilostyles between 36 and 57 my ago, and the crown age of the Australian clade 8–18 my ago. In our study system, host-age calibrations did not yield well-constrained results, but they may work better in other parasite clades. For small data sets where statistical convergence can be reached even with complex models, random local clocks should be explored as an alternative to the exclusive reliance on UCLN clocks.

The economically most important group of bees is the “corbiculates”, or pollen basket bees, some 890 species of honeybees (Apis), bumblebees (Bombus), stingless bees (Meliponini), and orchid bees (Euglossini). Molecular studies have indicated that the corbiculates are closest to the New World genera Centris, with 230 species, and Epicharis, with 35, albeit without resolving the precise relationships. Instead of concave baskets, these bees have hairy hind legs on which they transport pollen mixed with floral oil, collected with setae on the anterior and middle legs. We sampled two-thirds of all Epicharis, a third of all Centris, and representatives of the four lineages of corbiculates for four nuclear gene regions, obtaining a well-supported phylogeny that has the corbiculate bees nested inside the Centris/Epicharis clade. Fossil-calibrated molecular clocks, combined with a biogeographic reconstruction incorporating insights from the fossil record, indicate that the corbiculate clade arose in the New World and diverged from Centris 84 (72–95) mya. The ancestral state preceding corbiculae thus was a hairy hind leg, perhaps adapted for oil transport as in Epicharis and Centris bees. Its replacement by glabrous, concave baskets represents a key innovation, allowing efficient transport of plant resins and large pollen/nectar loads and freeing the corbiculate clade from dependence on oil-offering flowers. The transformation could have involved a novel function of Ubx, the gene known to change hairy into smooth pollen baskets in Apis and Bombus.

Absolute times from calibrated DNA phylogenies can be used to infer lineage diversification, the origin of new ecological niches, or the role of long distance dispersal in shaping current distribution patterns. Molecular-clock dating of non-vascular plants, however, has lagged behind flowering plant and animal dating. Here, we review dating studies that have focused on bryophytes with several goals in mind, (i) to facilitate cross-validation by comparing rates and times obtained so far; (ii) to summarize rates that have yielded plausible results and that could be used in future studies; and (iii) to calibrate a specieslevel phylogeny for Nothoceros, a model for plastid genome evolution in hornworts. Including the present work, there have been 18 molecular clock studies of liverworts, mosses, or hornworts, the majority with fossil calibrations, a few with geological calibrations or dated with previously published plastid substitution rates. Over half the studies cross-validated inferred divergence times by using alternative calibration approaches. Plastid substitution rates inferred for ‘‘bryophytes’’ are in line with those found in angiosperm studies, implying that bryophyte clock models can be calibrated either with published substitution rates or with fossils, with the two approaches testing and cross-validating each other. Our phylogeny of Nothoceros is based on 44 accessions representing all suspected species and a matrix of six markers of nuclear, plastid, and mitochondrial DNA. The results show that Nothoceros comprises 10 species, nine in the Americas and one in New Zealand (N. giganteus), with the divergence between the New Zealand species and its Chilean sister species dated to the Miocene and therefore due to long-distance dispersal. Based on the new tree, we formally transfer two species of Megaceros into Nothoceros, resulting in the new combinations N. minarum (Nees) J.C. Villarreal and N. schizophyllus (Gottsche ex Steph.) J.C. Villarreal, and we also newly synonymize eight names described in Megaceros.

Summary � Leaf out phenology affects a wide variety of ecosystem processes and ecological interactions and will take on added significance as leaf out times increasingly shift in response to warming temperatures associated with climate change. There is, however, relatively little information available on the factors affecting species differences in leaf out phenology. � An international team of researchers from eight Northern Hemisphere temperate botanical gardens recorded leaf out dates of c. 1600 woody species in 2011 and 2012. � Leaf out dates in woody species differed by as much as 3 months at a single site and exhibited strong phylogenetic and anatomical relationships. On average, angiosperms leafed out earlier than gymnosperms, deciduous species earlier than evergreen species, shrubs earlier than trees, diffuse and semi-ring porous species earlier than ring porous species, and species with smaller diameter xylem vessels earlier than species with larger diameter vessels. The order of species leaf out was generally consistent between years and among sites. � As species distribution and abundance shift due to climate change, interspecific differences in leaf out phenology may affect ecosystem processes such as carbon, water, and nutrient cycling. Our open access leaf out data provide a critical framework for monitoring and modelling such changes going forward.

A well-timed phenology is essential for plant growth and reproduction, but species-specific phenological strategies are still poorly understood. Here, we use a common garden approach to compare biannual leaf-out data for 495 woody species growing outdoors in Munich, 90% of them not native to that climate regime. For three species, data were augmented by herbarium dates for 140-year-long time series. We further meta-analysed 107 temperate-zone woody species in which leaf-out cues have been studied, half of them also monitored here. Southern climate-adapted species flushed significantly later than natives, and photoperiod- and chilling- sensitive species all flushed late. The herbarium method revealed the extent of species-specific climate tracking. Our results forecast that: (1) a northward expansion of southern species due to climate warming will increase the number of late flushers in the north, counteracting documented and expected flushing time advances; and (2) photoperiod- and chilling-sensitive woody species cannot rapidly track climate warming.

The lily family Colchicaceae consists of geophytic herbs distributed on all continents except the Neotropics. It is particularly diverse in southern Africa, where 80 of the 270 species occur. Colchicaceae exhibit a wide range of ploidy levels, from 2n = 14 to 2n = 216. To understand where and how this cytogenetic diversity arose, we generated multilocus phylogenies of the Colchicaceae and the Colchicum clade that respectively included 85 or 137 species plus relevant outgroups. To infer the kinds of events that could explain the observed numbers in the living species (dysploidy, polyploidization, or demi-duplication, i.e. fusion of gametes of different ploidy), we compared a series of likelihood models on phylograms, penalized likelihood ultrametric trees, and relaxed clock chronograms that contained the 58 or 112 species with published chromosome counts. While such models involve simplification and cannot address the processes behind chromosomal rearrangements, they can help frame questions about...

Aim Likelihood analyses of ancestral ranges require a parameterized model that consists of a time-calibrated phylogeny, an ‘adjacency matrix’ of allowed or forbidden area connections, and an ‘area–dispersal’ matrix with probabilities for discrete periods of time. The approach is implemented in the software Lagrange. Because it can incorporate information about past continental positions, the approach has been used in historical biogeographical studies of relatively old clades. Surprisingly, no study has evaluated the interactions among these input matrices. Here we use the lily family Colchicaceae and artificial data to study the relative effect of the input matrices on final estimates. Location Africa, Australia, Eurasia, North America and South America. Methods Using eight plastid, mitochondrial and nuclear DNA regions from 85 of the c. 280 species of Colchicaceae (representing all genera and the entire geographical range) and relevant outgroups, we obtained a well-resolved phylogeny dated with a molecular clock. We then assigned species to six geographical distributions and carried out 22 Lagrange runs in which we modified the adjacency and dispersal matrices, the latter with zero, two or four time periods and one, three or five dispersal probabilities. For a second data set, the areas at deep nodes in the empirical tree were modified by shuffling species distributions. Models were compared based on global log-likelihoods. Results The adjacency matrix strongly determined the outcome, while time slices and dispersal probability categories had minor effects. Ancestral areas reconstructed at most nodes were unaffected by the different input matrices. Colchicaceae are likely to have originated in Cretaceous East Gondwana, initially diversified in Australia (c. 67 Ma), reached southern Africa during the Palaeocene–Eocene, and from there extended their range to Southeast Asia (probably through Arabia) and then North America (through Beringia). Main conclusions At least in small data sets, Lagrange models should be tested with sensitivity analyses as carried out here, concentrating on constrained versus unconstrained adjacency matrices, and it should be good practice to report the set-up of both input matrices, not just the dispersal matrix, which is the less decisive of the two.

The circumscription of the five to eight species of Luffa, as well as their correct names, have long been problematic. Experts on the genus, most recently C. Heiser and C. Jeffrey, have disagreed on the number of species in the New World and the application of the name L. operculata, which in turn affected the names L. quinquefida and L. sepium. Heiser used classic biosystematic methods, including experimental crossing, to infer species boundaries, but neither researcher had today's option of using DNA sequences for this purpose. We sequenced 51 accessions of Luffa, representing the geographic range of the genus and as much as possible topotypical or type material. Phylogenies from four non-coding plastid regions and the nuclear ribosomal DNA spacer region show that eight clades of specimens have geographic-morphological coherence. Heiser's view that Luffa has three species in the New World is supported, and there are four species in tropical and subtropical Asia. Australia has an endemic species...

• Premise of the study: The Foulden Maar lake sediments in Otago, South Island, New Zealand, date to the earliest Miocene and provide an important picture of the diversity of the Australasian biota, paleoecology, and climate at a time when New Zealand had a smaller land area than today. The diverse rainforest contains many taxa now restricted to Australia, New Caledonia, or South America. The presence of Luzuriaga-like fossils in these deposits is important for understanding Alstroemeriaceae evolution and the biogeography of genera shared between New Zealand and South America.• Methods: Leaves and a flower with in situ pollen that resemble extant Luzuriaga are described and placed phylogenetically. Geographic range information and a molecular clock model for the Alstroemeriaceae were used to investigate possible biogeographic scenarios and the influence of the new fossil on inferred divergence times.• Key results: Luzuriaga peterbannisteri Conran, Bannister, Mildenh., & D.E.Lee sp. nov. represents the first macrofossil record for Alstroemeriaceae. An associated Luzuriaga-like flower with in situ fossil pollen of Liliacidites contortus Mildenh. & Bannister sp. nov. is also described. The biogeographic analysis suggests that there have been several dispersal events across the Southern Ocean for the genus, with the fossil representing a now-extinct New Zealand lineage.• Conclusions: Luzuriaga was present in Early Miocene New Zealand, indicating a long paleogeographic history for the genus, and L. peterbannisteri strengthens biogeographic connections between South America and Australasia during the Oligocene and earliest Miocene.

Most plant–pollinator interactions are mutualistic, involving rewards provided by flowers or inflorescences to pollinators. Antagonistic plant–pollinator interactions, in which flowers offer no rewards, are rare and concentrated in a few families including Araceae. In the latter, they involve trapping of pollinators, which are released loaded with pollen but unrewarded. To understand the evolution of such systems, we compiled data on the pollinators and types of interactions, and coded 21 characters, including interaction type, pollinator order, and 19 floral traits. A phylogenetic framework comes from a matrix of plastid and new nuclear DNA sequences for 135 species from 119 genera (5342 nucleotides). The ancestral pollination interaction in Araceae was reconstructed as probably rewarding albeit with low confidence because information is available for only 56 of the 120–130 genera. Bayesian stochastic trait mapping showed that spadix zonation, presence of an appendix, and flower sexuality were correlated with pollination interaction type. In the Araceae, having unisexual flowers appears to have provided the morphological precondition for the evolution of traps. Compared with the frequency of shifts between deceptive and rewarding pollination systems in orchids, our results indicate less lability in the Araceae, probably because of morphologically and sexually more specialized inflorescences.

Studies over the past ten years have shown that the crown groups of most conifer genera are only about 15-25 Ma old. The genus Picea (spruces, Pinaceae), with around 35 species, appears to be no exception. In addition, molecular studies of co-existing spruce species have demonstrated frequent introgression. Perhaps not surprisingly therefore previous phylogenetic studies of species relationships in Picea, based mostly on plastid sequences, suffered from poor statistical support. We therefore generated mitochondrial, nuclear, and further plastid DNA sequences from carefully sourced material, striking a balance between alignability with outgroups and phylogenetic signal content. Motif duplications in mitochondrial introns were treated as characters in a stochastic Dollo model; molecular clock models were calibrated with fossils; and ancestral ranges were inferred under maximum likelihood. In agreement with previous findings, Picea diverged from its sister clade 180 million years ago (Ma), and the most recent common ancestor of today's spruces dates to 28 Ma. Different from previous analyses though, we find a large Asian clade, an American clade, and a Eurasian clade. Two expansions occurred from Asia to North America and several between Asia and Europe. Chinese P. brachytyla, American P. engelmannii, and Norway spruce, P. abies, are not monophyletic, and North America has ten, not eight species. Divergence times imply that Pleistocene refugia are unlikely to be the full explanation for the relationships between the European species and their East Asian relatives. Thus, northern Norway spruce may be part of an Asian species complex that diverged from the southern Norway spruce lineage in the Upper Miocene, some 6 Ma, which can explain the deep genetic gap noted in phylogeographic studies of Norway spruce. The large effective population sizes of spruces, and incomplete lineage sorting during speciation, mean that the interspecific relationships within each of the geographic clades require further studies, especially based on genomic information and population genetic data.

Using molecular data, we recently showed that Carica papaya L. is sister to a Mexican/Guatemalan clade of two genera, Jarilla Rusby with three species and Horovitzia V.M. Badillo with one. These species are herbs or thin-stemmed trees and may be of interest for future genomics-enabled papaya breeding. Here we clarify the correct names of J. heterophylla (Cerv. ex La Llave) Rusby and J. caudata (Brandegee) Standl., which were confused in a recent systematic treatment of Jarilla (McVaugh 2001). We designate epitypes for both, provide weblinks to type specimens, a key to the species of Jarilla and Horovitzia, and notes on their habitats and distribution.

Betulaceae, with 120–150 species in six genera, are a family of Fagales that occurs mainly in the Northern Hemisphere. Previous studies of the evolution of Alnus, Betula, Carpinus, Corylus, Ostrya and Ostryopsis have relied on a relatively small number of sequence data and molecular clocks with fixed-point calibrations. We exploited GenBank to construct Betulaceae matrices of up to 900 sequence accessions and 9300 nucleotides of nuclear and plastid DNA; we also computed species consensus sequences to build 46- and 29-species matrices that strike a balance between species sampling and nucleotide sampling. Trees were rooted on Ticodendraceae and Casuarinaceae, and divergence times were inferred under relaxed and strict molecular clocks, using alternative fossil constraints. The data support the traditional two subfamilies, Betuloideae (Alnus, Betula) and Coryloideae, and show that Ostryopsis is sister to Ostrya/Carpinus. The fossil record and molecular clocks calibrated with alternating fossils indicate that the stem lineage of Betulaceae dates back to the Upper Cretaceous, the two subfamilies to the Palaeocene and the most recent common ancestors of each of the living genera to the mid- to late Miocene. A substitution rate shift in Coryloideae between 25 and 15 Mya preceded the mid-Miocene climatic optimum and may be linked to temperate niches that became available following the mid-Miocene. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 172, 465–477.

SUMMARY Comparisons of closely related species are needed to understand the fine-scale dynamics of retrotransposon evolution in flowering plants. Towards this goal, we classified the long terminal repeat (LTR) retrotransposons from six diploid and one tetraploid species of Orobanchaceae. The study species are the autotrophic, non-parasitic Lindenbergia philippensis (as an out-group) and six closely related holoparasitic species of Orobanche [O. crenata, O. cumana, O. gracilis (tetraploid) and O. pancicii] and Phelipanche (P. lavandulacea and P. ramosa). All major plant LTR retrotransposon clades could be identified, and appear to be inherited from a common ancestor. Species of Orobanche, but not Phelipanche, are enriched in Ty3/Gypsy retrotransposons due to a diversification of elements, especially chromoviruses. This is particularly striking in O. gracilis, where tetraploidization seems to have contributed to the Ty3/Gypsy enrichment and led to the emergence of seven large species-specific families of chromoviruses. The preferential insertion of chromoviruses in heterochromatin via their chromodomains might have favored their diversification and enrichment. Our phylogenetic analyses of LTR retrotransposons from Orobanchaceae also revealed that the Bianca clade of Ty1/Copia and the SMART-related elements are much more widely distributed among angiosperms than previously known.

Intuitively, interannual spring temperature variability (STV) should influence the leaf-out strategies of temperate zone woody species, with high winter chilling requirements in species from regions where spring warming varies greatly among years. We tested this hypothesis using experiments in 215 species and leaf-out monitoring in 1585 species from East Asia (EA), Europe (EU) and North America (NA). The results reveal that species from regions with high STV indeed have higher winter chilling requirements, and, when grown under the same conditions, leaf out later than related species from regions with lower STV. Since 1900, STV has been consistently higher in NA than in EU and EA, and under experimentally short winter conditions NA species required 84% more spring warming for bud break, EU ones 49% and EA ones only 1%. These previously unknown continental-scale differences in phenological strategies underscore the need for considering regional climate histories in global change models.

Background With some 700 species, the pantropical Crotalaria is among the angiosperm’s largest genera. We sampled 48% of the species from all sections (and representatives of the 15 remaining Crotalarieae genera) for nuclear and plastid DNA markers to infer changes in climate niches, flower morphology, leaf type, and chromosome numbers. Results Crotalaria is monophyletic and most closely related to African Bolusia (five species) from which it diverged 23 to 30 Ma ago. Ancestral state reconstructions reveal that leaf and flower types are conserved in large clades and that leaf type is uncorrelated to climate as assessed with phylogenetically-informed analyses that related compound vs. simple leaves to the mean values of four Bioclim parameters for 183 species with good occurrence data. Most species occur in open habitats

BMC evolutionary biology 16, 248 (2016). doi:10.1186/s12862-016-0827-5, Published by BioMed Central, London

A growing body of evidence is pointing to an important role of horizontal gene transfer (HGT) in the evolution of higher plants. However, reports of HGTs of transposable elements (TEs) in plants are still scarce and only one case is known of a class II transposon horizontally transferred between grasses. To investigate possible TE transfers in dicots, we performed transcriptome screening in the obligate root parasite Phelipanche aegyptiaca (Orobanchaceae), data-mining in the draft genome assemblies of four other Orobanchaceae, gene cloning, gene annotation in species with genomic information and a molecular phylogenetic analysis. We discovered that the broomrape genera Phelipanche and Orobanche acquired two related nuclear genes (christened BO transposase genes), a new group of the hAT superfamily of class II transposons, from Asian Sisymbrieae or a closely related tribe of Brassicaceae, by HGT. The collinearity of the flanking genes, lack of a classic border structure and low expression levels suggest that BO transposase genes cannot transpose in Brassicaceae, whereas they are highly expressed in P. aegyptiaca.

Cynomoriaceae, one of the last unplaced families of flowering plants, comprise one or two species or subspecies of root parasites that occur from the Mediterranean to the Gobi Desert. Using Illumina sequencing, we assembled the mitochondrial and plastid genomes as well as some nuclear genes of a Cynomorium specimen from Italy. Selected genes were also obtained by Sanger sequencing from individuals collected in China and Iran, resulting in matrices of 33 mitochondrial, 6 nuclear, and 14 plastid genes and rDNAs enlarged to include a representative angiosperm taxon sampling based on data available in GenBank. We also compiled a new geographic map to discern possible discontinuities in the parasites' occurrence. Cynomorium has large genomes of 13.70-13.61 (Italy) to 13.95-13.76 pg (China). Its mitochondrial genome consists of up to 49 circular subgenomes and has an overall gene content similar to that of photosynthetic angiosperms, while its plastome retains only 27 of the normally 116 genes. Nuclear, plastid and mitochondrial phylogenies place Cynomoriaceae in Saxifragales, and we found evidence for several horizontal gene transfers from different hosts, as well as intracellular gene transfers.

SummaryObligate mutualisms require filtering mechanisms to prevent their exploitation by opportunists, but ecological contexts and traits facilitating the evolution of such mechanisms are largely unknown. We investigated the evolution of filtering mechanisms in an epiphytic ant-plant symbiotic system in Fiji involving Rubiaceae and dolichoderine ants, using field experiments, metabolomics, X-ray micro-computed tomography (micro-CT) scanning and phylogenetics. We discovered a novel plant reward consisting of sugary sap concealed in post-anthetic flowers only accessible to Philidris nagasau workers that bite through the thick epidermis. In five of the six species of Rubiaceae obligately inhabited by this ant, the nectar glands functioned for 10 d after a flower's sexual function was over. Sugar metabolomics and field experiments showed that ant foraging tracks sucrose levels, which only drop at the onset of fruit development. Ontogenetic analyses of our focal species and their relatives revealed a 25-fold increase in nectary size and delayed fruit development in the ant-rewarding species, and Bayesian analyses of several traits showed the correlated evolution of sugar rewards and symbiosis specialization. Concealed floral nectar forestalls exploitation by opportunists (generalist ants) and stabilizes these obligate mutualisms. Our study pinpoints the importance of partner choice mechanisms in transitions from facultative to obligate mutualisms.

Few angiosperms have distinct Y chromosomes. Among those that do are Silene latifolia (Caryophyllaceae), Rumex acetosa (Polygonaceae) and Coccinia grandis (Cucurbitaceae), the latter having a male/female difference of 10% of the total genome (female individuals have a 0.85 pg genome, male individuals 0.94 pg), due to a Y chromosome that arose about 3 million years ago. We compared the sequence composition of male and female C. grandis plants and determined the chromosomal distribution of repetitive and organellar DNA with probes developed from 21 types of repetitive DNA, including 16 mobile elements. The size of the Y chromosome is largely due to the accumulation of certain repeats, such as members of the Ty1/copia and Ty3/gypsy superfamilies, an unclassified element and a satellite, but also plastome- and chondriome-derived sequences. An abundant tandem repeat with a unit size of 144 bp stains the centromeres of the X chromosome and the autosomes, but is absent from the Y centromere. Immunostaining with pericentromere-specific markers for anti-histone H3Ser10ph and H2AThr120ph revealed a Y-specific extension of these histone marks. That the Y centromere has a different make-up from all the remaining centromeres raises questions about its spindle attachment, and suggests that centromeric or pericentromeric chromatin might be involved in the suppression of recombination.

Based on morphological and molecular data, calcareous dinoflagellates (Thoracosphaeraceae, Peridiniales) are a monophyletic group comprising the three major clades Ensiculifera/Pentapharsodinium, Thoracosphaera/Pfiesteria, and Scrippsiella sensu lato. We used stratigraphically well-documented first occurrences of particular archeopyle types to constrain relaxed Bayesian molecular clocks applied to nuclear rRNA sequences of 18 representatives of the three main clades. By comparing divergence estimates obtained in differently calibrated clocks with first stratigraphic occurrences of taxa not themselves used as constraints, we identified plausible divergence times for several subclades of calcareous dinoflagellates. The initial diversification of extant calcareous dinoflagellates probably took place in the Late Jurassic, with the three main clades all established by the Cretaceous. The two mesoepicystal operculum types observed in calcareous dinoflagellates probably evolved independently from simple apical archeopyles. Based on our taxon sample, the K/T boundary had relatively little effect on the diversity of the group, with several lineages dating to before 65 mya (million years ago). The first stratigraphic occurrences of key taxa, such as Thoracosphaera and Calciodinellum (not themselves used as constraints), are in agreement with the molecular time estimates. Conflicts that involve "Calciodinellum"levantinum, Leonella, Pentapharsodinium, Pernambugia, and the Scrippsiella trochoidea species complex may be due to inaccurate assignment of fossils because of high morphological homoplasy and insufficient knowledge of the extant diversity of calcareous dinoflagellates.

PREMISE OF THE STUDY: Caricaceae include six genera and 34 species, among them papaya, a model species in plant sex chromosome research. The family was held to have a conserved karyotype with 2n = 18 chromosomes, an assumption based on few counts. We examined the karyotypes and genome size of species from all genera to test for possible cytogenetic variation. METHODS: We used fluorescent in situ hybridization using standard telomere, 5S, and 45S rDNA probes. New and published data were combined with a phylogeny, molecular clock dating, and C values (available for similar to 50% of the species) to reconstruct genome evolution. KEY RESULTS: The African genus Cylicomorpha, which is sister to the remaining Caricaceae (all neotropical), has 2n = 18, as do the species in two other genera. A Mexican clade of five species that includes papaya, however, has 2n = 18 (papaya), 2n = 16 (Horovitzia cnidoscoloides), and 2 n = 14 (Jarilla caudata and J. heterophylla;third Jarilla not counted), with the phylogeny indicating that the dysploidy events occurred similar to 16.6 and similar to 5.5 million years ago and that Jarilla underwent genome size doubling (similar to 450 to 830-920 Mbp/haploid genome). Pericentromeric interstitial telomere repeats occur in both Jarilla adjacent to 5S rDNA sites, and the variability of 5S rDNA sites across all genera is high. CONCLUSIONS: On the basis of outgroup comparison, 2n = 18 is the ancestral number, and repeated chromosomal fusions with simultaneous genome size increase as a result of repetitive elements accumulating near centromeres characterize the papaya clade. These results have implications for ongoing genome assemblies in Caricaceae.

Delphinieae (Ranunculaceae) comprise ∼650 species of temperate herbs in Eurasia, North America, and Africa. Their zygomorphic flowers have been the object of numerous studies in morphology, ecology, and developmental genetics, and new phylogenetic insights make it timely to synthesize knowledge about their evolution. Key features of Delphinieae flowers are unusual nectaries consisting of paired organs of the inner perianth whorl that are completely enclosed by a single dorsal organ of the outer whorl. We investigated the floral development of five annual, unicarpellate species of Delphinium, focusing on perianth organization. The results show that the nectar-storing organ in these species results from the postgenital fusion of two primordia of the internal perianth whorl. Eleven floral traits traced on a phylogeny of Delphinieae reveal only two homoplasies in the perianth, namely, the nightcap shape of the dorsal organ of the external perianth whorl and the reduction of the internal perianth whorl to two ...

Ribulose-1,5-Biphosphate-carboxylase-oxygenase (RuBisCO) has a crucial role in carbon fixation but a slow catalytic rate, a problem overcome in some plant lineages by physiological and anatomical traits that elevate carbon concentrations around the enzyme. Such carbon-concentrating mechanisms are hypothesized to have evolved during periods of low atmospheric CO 2 . Hornworts, the sister to vascular plants, have a carbon-concentrating mechanism that relies on pyrenoids, proteinaceous bodies mostly consisting of RuBisCO. We generated a phylogeny based on mitochondrial and plastid sequences for 36% of the approximately 200 hornwort species to infer the history of gains and losses of pyrenoids in this clade; we also used fossils and multiple dating approaches to generate a chronogram for the hornworts. The results imply five to six origins and an equal number of subsequent losses of pyrenoids in hornworts, with the oldest pyrenoid gained ca . 100 Mya, and most others at 2 ], and the maintenance of pyrenoids during episodes of high [CO 2 ] all argue against the previously proposed relationship between pyrenoid origin and low [CO 2 ]. The selective advantages, and costs, of hornwort pyrenoids thus must relate to additional factors besides atmospheric CO 2 .

Papaya (Carica papaya) is a crop of great economic importance, and the species was among the first plants to have its genome sequenced. However, there has never been a complete species-level phylogeny for the Caricaceae, and the crop's closest relatives are therefore unknown. We investigated the evolution of the Caricaceae based on sequences from all species and genera, the monospecific Carica, African Cylicomorpha with two species, South American Jacaratia and Vasconcellea with together c. 28 species, and Mexican/Guatemalan Jarilla and Horovitzia with four species. Most Caricaceae are trees or shrubs; the species of Jarilla, however, are herbaceous. We generated a matrix of 4711 nuclear and plastid DNA characters and used maximum likelihood (ML) and Bayesian analysis to infer species relationships, rooting trees on the Moringaceae. Divergence times were estimated under relaxed and strict molecular clocks, using different subsets of the data. Ancestral area reconstruction relied on a ML approach. The deepest split in the Caricaceae occurred during the Late Eocene, when the ancestor of the Neotropical clade arrived from Africa. In South America, major diversification events coincide with the Miocene northern Andean uplift and the initial phase of the tectonic collision between South America and Panama resulting in the Panamanian land bridge. Carica papaya is sister to Jarilla/Horovitzia, and all three diverged from South American Caricaceae in the Oligocene, 27 (22-33) Ma ago, coincident with the early stages of the formation of the Panamanian Isthmus. The discovery that C. papaya is closest to a clade of herbaceous or thin-stemmed species has implications for plant breeders who have so far tried to cross papaya only with woody highland papayas (Vasconcellea).

 Premise of the study: Understanding the fl exibility of monocot genomes requires a phylogenetic framework, which so far is available for few of the ca. 2800 genera. Here we use a molecular tree for the South American genus Alstroemeria to place karyological information, including fl uorescent in situ hybridization (FISH) signals, in an explicit evolutionary context.  Methods: From a phylogeny based on plastid, nuclear, and mitochondrial sequences for most species of Alstroemeria , we selected early-branching (Chilean) and derived (Brazilian) species for which we obtained 18S-25S and 5S rDNA FISH signals; we also analyzed chromosome numbers, 1C-values, and telomere FISH signals (in two species).  Key results: Chromosome counts for Alstroemeria cf. rupestris and A. pulchella confi rm 2 n = 16 as typical of the genus, which now has chromosomes counted for 29 of its 78 species. The rDNA sites are polymorphic both among and within species, and interstitial telomeric sites in Alstroemeria cf. rupestris suggest chromosome fusion.  Conclusions: In spite of a constant chromosome number, closely related species of Alstroemeria differ drastically in their rDNA, indicating rapid increase, decrease, or translocations of these genes. Previously proposed Brazilian and Chilean karyotype groups are not natural, and the n = 8 chromosomes in Alstroemeria compared to n = 9 in its sister genus Bomarea may result from a Robertsonian fusion.

We used next-generation sequencing to characterize the genomes of nine species of Orobanchaceae of known phylogenetic relationships, different life forms, and including a polyploid species. The study species are the autotrophic, nonparasitic Lindenbergia philippensis, the hemiparasitic Schwalbea americana, and seven nonphotosynthetic parasitic species of Orobanche (Orobanche crenata, Orobanche cumana, Orobanche gracilis (tetraploid), and Orobanche pancicii) and Phelipanche (Phelipanche lavandulacea, Phelipanche purpurea, and Phelipanche ramosa). Ty3/Gypsy elements comprise 1.93%–28.34% of the nine genomes and Ty1/Copia elements comprise 8.09%–22.83%. When compared with L. philippensis and S. americana, the nonphotosynthetic species contain higher proportions of repetitive DNA sequences, perhaps reflecting relaxed selection on genome size in parasitic organisms. Among the parasitic species, those in the genus Orobanche have smaller genomes but higher proportions of repetitive DNA than those in Phelipanche, mostly due to a diversification of repeats and an accumulation of Ty3/ Gypsy elements. Genome downsizing in the tetraploid O. gracilis probably led to sequence loss across most repeat types.

The family Araceae (3790 species, 117 genera) has one of the oldest fossil records among angiosperms. Ecologically, members of this family range from free-floating aquatics (Pistia and Lemna) to tropical epiphytes. Here, we infer some of the macroevolutionary processes that have led to the worldwide range of this family and test how the inclusion of fossil (formerly occupied) geographical ranges affects biogeographical reconstructions. Using a complete genus-level phylogeny from plastid sequences and outgroups representing the 13 other Alismatales families, we estimate divergence times by applying different clock models and reconstruct range shifts under different models of past continental connectivity, with or without the incorporation of fossil locations. Araceae began to diversify in the Early Cretaceous (when the breakup of Pangea was in its final stages), and all eight subfamilies existed before the K/T boundary. Early lineages persist in Laurasia, with several relatively recent entries into Africa, South America, South-East Asia and Australia. Water-associated habitats appear to be ancestral in the family, and DNA substitution rates are especially high in free-floating Araceae. Past distributions inferred when fossils are included differ in nontrivial ways from those without fossils. Our complete genus-level time-scale for the Araceae may prove to be useful for ecological and physiological studies.

Hallucinogenic or toxic species of Brunfelsia (Solanaceae: Petunieae) are important in native cultures throughout South America, and the genus also contains several horticulturally important species. An earlier morphological revision of the c. 50 species recognized three main groups, one consisting of the 23 Antillean species, another of southern South American and Andean species, and a third of species from the Amazon Basin and Guiana Shield. Based on plastid and nuclear DNA sequences from up to 65 accessions representing 80% of the species, we generated a phylogeny and a calibrated chronogram for Brunfelsia to infer clade expansion and shifts in pollinators and fruit types. Brunfelsia flowers offer nectar, and attract lepidoptera, hummingbirds, or bees; the fruits are dry or fleshy. Our results imply that Brunfelsia is 16-21 Myr old and entered the Antilles from South America early during its history, with subsequent expansion along the island arc. The ancestor of the Antillean clade was hawk-moth-pollinated and had fleshy capsules, perhaps facilitating dispersal by birds. The only shift to hummingbird pollination occurred on Cuba, which also harbors the largest single radiation, with 11 species (10 included in our study) that apparently arose over the past 4 Myr. Jamaica, Hispaniola, and Puerto Rico each sustained smaller radiations. The data also reveal at least one new species.