Radiation Of Angiosperms Research Articles

Geography, environment and organismal traits in the diversification of a major tropical herbaceous angiosperm radiation.

The generation of plant diversity involves complex interactions between geography, environment and organismal traits. Many macroevolutionary processes and emergent patterns have been identified in different plant groups through the study of spatial data, but rarely in the context of a large radiation of tropical herbaceous angiosperms. A powerful system for testing interrelated biogeographical hypotheses is provided by the terrestrial bromeliads, a Neotropical group of extensive ecological diversity and importance. In this investigation, distributional data for 564 species of terrestrial bromeliads were used to estimate variation in the position and width of species-level hydrological habitat occupancy and test six core hypotheses linking geography, environment and organismal traits. Taxonomic groups and functional types differed in hydrological habitat occupancy, modulated by convergent and divergent trait evolution, and with contrasting interactions with precipitation abundance and seasonality. Plant traits in the Bromeliaceae are intimately associated with bioclimatic differentiation, which is in turn strongly associated with variation in geographical range size and species richness. These results emphasize the ecological relevance of structural-functional innovation in a major plant radiation.

Widespread Whole Genome Duplications Contribute to Genome Complexity and Species Diversity in Angiosperms

Gene duplications provide evolutionary potentials for generating novel functions, while polyploidization or whole genome duplication (WGD) doubles the chromosomes initially and results in hundreds to thousands of retained duplicates. WGDs are strongly supported by evidence commonly found in many species-rich lineages of eukaryotes, and thus are considered as a major driving force in species diversification. We performed comparative genomic and phylogenomic analyses of 59 public genomes/transcriptomes and 46 newly sequenced transcriptomes covering major lineages of angiosperms to detect large-scale gene duplication events by surveying tens of thousands of gene family trees. These analyses confirmed most of the previously reported WGDs and provided strong evidence for novel ones in many lineages. The detected WGDs supported a model of exponential gene loss during evolution with an estimated half-life of approximately 21.6 million years, and were correlated with both the emergence of lineages with high degrees of diversification and periods of global climate changes. The new datasets and analyses detected many novel WGDs widely spread during angiosperm evolution, uncovered preferential retention of gene functions in essential cellular metabolisms, and provided clues for the roles of WGD in promoting angiosperm radiation and enhancing their adaptation to environmental changes.

Diversity of palmately lobed leaves in the early–middle Albian of eastern Russia

Diversity of palmately lobed leaves of angiosperms of the early–middle Albian floras of the Kolyma River Basin, the Omsukchan Coal Basin, Khabarovsk and Primorye Regions was studied. Leaf fossils, previously compared with those of genera Aralia, Sassafras and Lindera, now reassigned to the fossil genus Araliaephyllum. Four new combinations and one new species are published: A. kolymense (Kryshtofovich) Golovneva, comb. nov., A. luciferum (Kryshtofovich) Golovneva, comb. nov., A. ussuriense (Krassilov) Golovneva, comb. nov., A. ievlevii (Samylina) Golovneva, comb. nov., and A. popovii Golovneva, sp. nov. The type material has been restudied in detail, and lectotypes have been selected to all newly typified species. These species share many lauralean morphological and venation features. They represent the most likely early members of this group. This relationship is based on detailed study of the leaf architecture and comparison with other fossils with studied epidermal features. This study expands our knowledge of radiation and biogeography patterns of early angiosperms in northeastern Asia.

Adaptive variation in vein placement underpins diversity in a major Neotropical plant radiation

Vein placement has been hypothesised to control leaf hydraulic properties, but the ecophysiological significance of variation in vein placement in the angiosperms has remained poorly understood. The highly diverse Neotropical Bromeliaceae offers an excellent system for exploring understudied relationships between leaf vein placement, physiological functions, and species ecology. To test key hypotheses regarding the links between vein placement, functional type divergences, and ecological diversity in the Bromeliaceae, I characterised the ratio of interveinal distance (IVD) to vein-epidermis distance (VED) in 376 species, representing all major functional types and 10% of the species diversity in the family, as well as bioclimatic properties and key leaf traits for subsets of species. There were significant differences in vein placement parameters in species of contrasting functional type, habitat association, and bioclimatic distribution. In many C3 tank-epiphytes, a greater ratio between interveinal distance and the depth of veins within the mesophyll reflects optimisation for resource foraging in shady, humid habitats. In succulent terrestrials, overinvestment in veins probably facilitates rapid recharge of water storage tissue, as well as restricting water loss. These results highlight how divergences in vein placement relate to distinctive ecophysiological strategies between and within bromeliad functional types, and provide timely insights into how structural–functional innovation has impacted the evolution of ecological diversity in a major radiation of tropical herbaceous angiosperms.

Genomic evidence reveals a radiation of placental mammals uninterrupted by the KPg boundary

The timing of the diversification of placental mammals relative to the Cretaceous-Paleogene (KPg) boundary mass extinction remains highly controversial. In particular, there have been seemingly irreconcilable differences in the dating of the early placental radiation not only between fossil-based and molecular datasets but also among molecular datasets. To help resolve this discrepancy, we performed genome-scale analyses using 4,388 loci from 90 taxa, including representatives of all extant placental orders and transcriptome data from flying lemurs (Dermoptera) and pangolins (Pholidota). Depending on the gene partitioning scheme, molecular clock model, and genic deviation from molecular clock assumptions, extensive sensitivity analyses recovered widely varying diversification scenarios for placental mammals from a given gene set, ranging from a deep Cretaceous origin and diversification to a scenario spanning the KPg boundary, suggesting that the use of suboptimal molecular clock markers and methodologies is a major cause of controversies regarding placental diversification timing. We demonstrate that reconciliation between molecular and paleontological estimates of placental divergence times can be achieved using the appropriate clock model and gene partitioning scheme while accounting for the degree to which individual genes violate molecular clock assumptions. A birth-death-shift analysis suggests that placental mammals underwent a continuous radiation across the KPg boundary without apparent interruption by the mass extinction, paralleling a genus-level radiation of multituberculates and ecomorphological diversification of both multituberculates and therians. These findings suggest that the KPg catastrophe evidently played a limited role in placental diversification, which, instead, was likely a delayed response to the slightly earlier radiation of angiosperms.

Less is more: Independent loss-of-function OCIMENE SYNTHASE alleles parallel pollination syndrome diversification in monkeyflowers (Mimulus).

Pollinator-mediated selection on flower phenotypes (e.g., shape, color, scent) is key to understanding the adaptive radiation of angiosperms, many of which have evolved specialized relationships with a particular guild of animal pollinators (e.g., birds, bats, moths, bees). E-β-Ocimene, a monoterpene produced by OCIMENE SYNTHASE (OS) in Mimulus lewisii, is a floral scent important in attracting the species' bumblebee pollinators. The taxa closely related to M. lewisii have evolved several different pollination syndromes, including hummingbird pollination and self pollination (autogamy). We are interested in how floral scent variation contributed to species diversification in this clade. We analyzed variation in E-β-ocimene emission within this Mimulus clade and explored its molecular basis through a combination of DNA sequencing, reverse transcriptase PCR, and enzyme functional analysis in vitro. We found that none of the taxa, other than M. lewisii, emitted E-β-ocimene from flowers. But the molecular basis underlying loss of E-β-ocimene emission is unique in each taxon, including deletion, missense, or frameshift mutations in the OS gene, and potential posttranscriptional downregulation. The molecular evidence suggests that parallel loss-of-function in OS is the best explanation for the observed pattern of E-β-ocimene emission, likely as the result of natural selection.

Evolutionary History of Subtilases in Land Plants and Their Involvement in Symbiotic Interactions.

Subtilases, a family of proteases involved in a variety of developmental processes in land plants, are also involved in both mutualistic symbiosis and host-pathogen interactions in different angiosperm lineages. We examined the evolutionary history of subtilase genes across land plants through a phylogenetic analysis integrating amino acid sequence data from full genomes, transcriptomes, and characterized subtilases of 341 species of diverse green algae and land plants along with subtilases from 12 species of other eukaryotes, archaea, and bacteria. Our analysis reconstructs the subtilase gene phylogeny and identifies 11 new gene lineages, six of which have no previously characterized members. Two large, previously unnamed, subtilase gene lineages that diverged before the origin of angiosperms accounted for the majority of subtilases shown to be associated with symbiotic interactions. These lineages expanded through both whole-genome and tandem duplication, with differential neofunctionalization and subfunctionalization creating paralogs associated with different symbioses, including nodulation with nitrogen-fixing bacteria, arbuscular mycorrhizae, and pathogenesis in different plant clades. This study demonstrates for the first time that a key gene family involved in plant-microbe interactions proliferated in size and functional diversity before the explosive radiation of angiosperms.

Insights into the Evolution of Hydroxyproline-Rich Glycoproteins from 1000 Plant Transcriptomes.

The carbohydrate-rich cell walls of land plants and algae have been the focus of much interest given the value of cell wall-based products to our current and future economies. Hydroxyproline-rich glycoproteins (HRGPs), a major group of wall glycoproteins, play important roles in plant growth and development, yet little is known about how they have evolved in parallel with the polysaccharide components of walls. We investigate the origins and evolution of the HRGP superfamily, which is commonly divided into three major multigene families: the arabinogalactan proteins (AGPs), extensins (EXTs), and proline-rich proteins. Using motif and amino acid bias, a newly developed bioinformatics pipeline, we identified HRGPs in sequences from the 1000 Plants transcriptome project (www.onekp.com). Our analyses provide new insights into the evolution of HRGPs across major evolutionary milestones, including the transition to land and the early radiation of angiosperms. Significantly, data mining reveals the origin of glycosylphosphatidylinositol (GPI)-anchored AGPs in green algae and a 3- to 4-fold increase in GPI-AGPs in liverworts and mosses. The first detection of cross-linking (CL)-EXTs is observed in bryophytes, which suggests that CL-EXTs arose though the juxtaposition of preexisting SPn EXT glycomotifs with refined Y-based motifs. We also detected the loss of CL-EXT in a few lineages, including the grass family (Poaceae), that have a cell wall composition distinct from other monocots and eudicots. A key challenge in HRGP research is tracking individual HRGPs throughout evolution. Using the 1000 Plants output, we were able to find putative orthologs of Arabidopsis pollen-specific GPI-AGPs in basal eudicots.

False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance

During the mid-Cretaceous, angiosperms diversified from several nondiverse lineages to their current global domination [1], replacing earlier gymnosperm lineages [2]. Several hypotheses explain this extensive radiation [3], one of which involves proliferation of insect pollinator associations in the transition from gymnosperm to angiosperm dominance. However, most evidence supports gymnosperm-insect pollinator associations, buttressed by direct evidence of pollen on insect bodies, currently established for four groups: Thysanoptera (thrips), Neuroptera (lacewings), Diptera (flies), and now Coleoptera (beetles). Each group represents a distinctive pollination mode linked to a unique mouthpart type and feeding guild [4-9]. Extensive indirect evidence, based on specialized head and mouthpart morphology, is present forone of these pollinator types, the long-proboscid pollination mode [10], representing minimally ten family-level lineages of Neuroptera, Mecoptera (scorpionflies), and Diptera [8, 10, 11]. A recurring feature uniting these pollinator modes is host associations withginkgoalean, cycad, conifer, and bennettitalean gymnosperms. Pollinator lineages bearing these pollination modes were categorized into four evolutionary cohorts during the 35-million-year-long angiosperm radiation, each defined by its host-plant associations(gymnosperm or angiosperm) and evolutionary pattern (extinction, continuation, or origination) during this interval [12]. Here, we provide the first direct evidence for one cohort, exemplified by the beetle Darwinylus marcosi, family Oedemeridae (false blister beetles), that had an earlier gymnosperm (most likely cycad) host association, later transitioning onto angiosperms [13]. This association constitutes one of four patterns explaining the plateau of family-level plant lineages generally and pollinating insects specifically during the mid-Cretaceous angiosperm radiation [12].

The vascular plants: open system of growth.

What is fascinating in plants (true also in sessile animals such as corals and hydroids) is definitely their open and indeterminate growth, as a result of meristematic activity. Plants as well as animals are characterized by a multicellular organization, with which they share a common set of genes inherited from a common eukaryotic ancestor; nevertheless, circa 1.5 billion years of evolutionary history made the two kingdoms very different in their own developmental biology. Flowering plants, also known as angiosperms, arose during the Cretaceous Period (145-65 million years ago), and up to date, they count around 235,000 species, representing the largest and most diverse group within the plant kingdom. One of the foundations of their success relies on the plant-pollinator relationship, essentially unique to angiosperms that pushed large speciation in both plants and insects and on the presence of the carpel, the structure devoted to seed enclosure. A seed represents the main organ preserving the genetic information of a plant; during embryogenesis, the primary axis of development is established by two groups of pluripotent cells: the shoot apical meristem (SAM), responsible for gene rating all aboveground organs, and the root apical meristem (RAM), responsible for producing all underground organs. During postembryonic shoot development, axillary meristem (AM) initiation and outgrowth are responsible for producing all secondary axes of growth including inflorescence branches or flowers. The production of AMs is tightly linked to the production of leaves and their separation from SAM. As leaf primordia are formed on the flanks of the SAM, a region between the apex and the developing organ is established and referred to as boundary zone. Interaction between hormones and the gene network in the boundary zone is fundamental for AM initiation. AMs only develop at the adaxial base of the leaf; thus, AM initiation is also strictly associated with leaf polarity. AMs function as new SAMs: form axillary buds with a few leaves and then the buds can either stay dormant or develop into shoot branches to define a plant architecture, which in turn affects assimilate production and reproductive efficiency. Therefore, the radiation of angiosperms was accompanied by a huge diversification in growth forms that determine an enormous morphological plasticity helping plants to environmental changes. In this review, we focused on the developmental processes of AM initiation and outgrowth. In particular, we summarized the primary growth of SAM, the key role of positional signals for AM initiation, and the dissection of molecular players involved in AM initiation and outgrowth. Finally, the interaction between phytohormone signals and gene regulatory network controlling AM development was discussed.

The phylogenetic position of eriophyoid mites (superfamily Eriophyoidea) in Acariformes inferred from the sequences of mitochondrial genomes and nuclear small subunit (18S) rRNA gene

Eriophyoid mites (superfamily Eriophyoidea) comprise >4400 species worldwide. Despite over a century of study, the phylogenetic position of these mites within Acariformes is still poorly resolved. Currently, Eriophyoidea is placed in the order Trombidiformes. We inferred the high-level phylogeny of Acari with the mitochondrial (mt) genome sequences of 110 species including four eriophyoid species, and the nuclear small subunit (18S) rRNA gene sequences of 226 species including 25 eriophyoid species. Maximum likelihood (ML), Bayesian inference (BI) and Maximum parsimony (MP) methods were used to analyze the sequence data. Divergence times were estimated for major lineages of Acari using Bayesian approaches. Our analyses consistently recovered the monophyly of Eriophyoidea but rejected the monophyly of Trombidiformes. The eriophyoid mites were grouped with the sarcoptiform mites, or were the sister group of sarcoptiform mites+non-eriophyoid trombidiform mites, depending on data partition strategies. Eriophyoid mites diverged from other mites in the Devonian (384Mya, 95% HPD, 352–410Mya). The origin of eriophyoid mites was dated to the Permian (262Mya, 95% HPD 230–307Mya), mostly prior to the radiation of gymnosperms (Triassic-Jurassic) and angiosperms (early Cretaceous). We propose that the placement of Eriophyoidea in the order Trombidiformes under the current classification system should be reviewed.

First record of microsporophyll genus Caytonanthus Thomas from Early Cretaceous beds of South Rewa Gondwana Basin, India: Its evolutionary and palaeogeographical significance

AbstractPteridosperms (seed ferns) are an extinct group of large and diverse plants. They were dominant elements during Palaeozoic and were also an important component in the Mesozoic vegetation but declined globally in the Cretaceous time with the radiation of angiosperms. The order Caytoniales of Mesozoic pteridosperm is an extinct group of seed ferns in the vegetation. They thrived for about 100 million years from the Late Triassic to the Late Cretaceous. Although they never attained dominant status, their rare occurrence in the vegetation is quite important and fascinating. For this extinct group of seed fern, Thomas () erected a new order Caytoniales which includes three organ genera Caytonia (megasporophyll), Caytonanthus (microsporophyll) and Sagenopteris (leaves). This paper is aimed to document first record of microsporophyll genus Caytonanthus, obtained from the Lower Cretaceous beds of the South Rewa Gondwana Basin of the Indian Peninsula. Its morpho‐characters, palaeogeographic distribution, along with its plausible southern origin through ancestral group like Glossopteridales, are surmised here.

Sex and the shifting biodiversity dynamics of marine animals in deep time

The fossil record of marine animals suggests that diversity-dependent processes exerted strong control on biodiversification: after the Ordovician Radiation, genus richness did not trend for hundreds of millions of years. However, diversity subsequently rose dramatically in the Cretaceous and Cenozoic (145 million years ago-present), indicating that limits on diversification can be overcome by ecological or evolutionary change. Here, we show that the Cretaceous-Cenozoic radiation was driven by increased diversification in animals that transfer sperm between adults during fertilization, whereas animals that broadcast sperm into the water column have not changed significantly in richness since the Late Ordovician (∼450 million years ago). We argue that the former group radiated in part because directed sperm transfer permits smaller population sizes and additional modes of prezygotic isolation, as has been argued previously for the coincident radiation of angiosperms. Directed sperm transfer tends to co-occur with many ecological traits, such as a predatory lifestyle. Ecological specialization likely operated synergistically with mode of fertilization in driving the diversification that began during the Mesozoic marine revolution. Plausibly, the ultimate driver of diversification was an increase in food availability, but its effects on the fauna were regulated by fundamental reproductive and ecological traits.

Phylogeny, classification and divergence times of pygmy leaf‐mining moths (Lepidoptera: Nepticulidae): the earliest lepidopteran radiation on Angiosperms?

AbstractNepticulidae represent one of the early diverging Lepidoptera lineages, and the family currently comprises over 850 described species. The larvae of the vast majority of the species are leaf miners on Angiosperms and highly monophagous, which has led to persistent ideas on coevolution with their plant hosts. We present here a molecular phylogeny based on eight gene fragments from 355 species, representing 20 out of 22 extant Nepticulidae genera. Using two fossil calibration points, we performed molecular dating to place the origin of the family in the Early Cretaceous, before the main Angiosperm diversification. Based on our results we propose a new classification, abandoning all ranks between family and genus, as well as subgenera to allow for a stable classification. The position of Enteucha Meyrick within Nepticulidae remains somewhat ambiguous, and the species‐rich cosmopolitan genus Stigmella Schrank, with nearly half of all described Nepticulidae, requires further study. Ectoedemia Busck, Zimmermannia Hering, Acalyptris Meyrick, Etainia Beirne, Parafomoria Borkowski, Muhabbetana Koçak & Kemal and Fomoria Beirne appear to have diversified in a relatively short evolutionary period, leading to short branches in the molecular phylogeny and unclear suprageneric relations. Otherwise support values throughout the phylogeny are mostly high and the species groups, genera and higher clades are discussed in respect of their supporting morphological and life‐history characters. Wing venation characters are confirmed to be mostly reliable and relevant for Nepticulidae classification, but some other previously used characters require reinterpretation. The species groups of most genera are recovered, but only partly so in the large genus Stigmella. The molecular dating results are compared with existing knowledge on the timing of the Angiosperm radiation and reveal that the diversification of Nepticulidae could largely have been contemporaneous with their hosts, although some of the genera restricted to a single plant family appear to have begun to diversify before their hosts.

Vegetation dynamics, angiosperm radiation and climatic changes in the Lusitanian Basin (Portugal) during Albian times

The mid-Cretaceous is characterised by significant evolutionary changes in terrestrial ecosystems. During this period, the typical vegetation consisting of ferns, conifers and other gymnosperms was replaced by an angiosperm-dominated flora. Long ranging and continuous records from the continental realm are mostly lacking due to the prevalence of sedimentary gaps. Here a comprehensive palynological and clay mineralogical dataset covering early Albian to early Cenomanian strata from the shallow marine São Julião section (Lusitanian Basin, Portugal) is analysed to reconstruct the vegetation composition as well as climate and weathering conditions in the continental realm. The palynological assemblage is dominated by non-saccate Classopollis and Inaperturopollenites gymnosperm pollen with low abundances of spores and angiosperm pollen. The clay mineralogical data is mainly composed of kaolinite and mica and shows two conspicuous intervals of high kaolinite abundances in the late Albian. Based on the ecological preferences of the associated parent plants of the spore-pollen assemblage, a gradual relative temperature increase is inferred in the hinterland of the Lusitanian Basin during the early Albian to early Cenomanian interval. The clay mineralogy and palynological record show that the climate in the continental hinterland of the Lusitanian Basin was semi-arid to sub-humid and punctuated by two distinct humid phases during the late Albian. Multivariate statistical analyses and a comparison with the nearby Guincho section demonstrate the significance and reproducibility of the spore-pollen record. Furthermore, the palynological assemblage implies that angiosperms and ferns occupied similar palaeo-ecological habitats, namely as understorey in Cheirolepidaceae, Araucariaceae and Cupressaceae conifer-dominated forests.

Cecilanthus polymerus, a novel multiparted flower from the mid-Cretaceous Rocky Point locality, Maryland

A flower from the early Cenomanian of northeastern Maryland, Cecilanthus polymerus gen. et sp. nov., is described using SEM and synchrotron X-ray microtomography. The flower has >20 strap-shaped tepals, ca. 50 spatulate stamens with embedded adaxial pollen sacs and possibly H-valvate dehiscence, and ca. 100 more-or-less plicate carpels. Floral phyllotaxis is whorled, with >10 parts per whorl, but slightly irregular. Pollen and ovules are not preserved, but locule shape suggests that the carpels are uniovulate. Similar characters occur in extant Magnoliales and the basal ANITA grade, but never in combination. In an attempt to resolve these ambiguities, we performed morphological phylogenetic analyses, with the arrangement of extant taxa constrained to trees based primarily on molecular data. With anther dehiscence and ovule number treated as unknown, Cecilanthus has several most-parsimonious positions: nested in Nymphaeales, nested in Magnoliales, sister to Laurales, and sister to Chloranthaceae. However, scoring anther dehiscence as H-valvate and ovule number as = 1, increases support for a position in Magnoliales. Cecilanthus demonstrates that the early radiation of angiosperms produced floral morphotypes unlike those in any surviving taxa, and it illustrates the value of characters such as pollen morphology and seed anatomy for phylogenetic placement of floral mesofossils.

The Tree of Life: China project

The Tree of Life: China project

DISCORDANT MORPHOLOGICAL DISPARITY AND TAXONOMIC DIVERSITY DURING THE CRETACEOUS ANGIOSPERM RADIATION : NORTH AMERICAN POLLEN RECORD

The Cretaceous angiosperm radiation offers an opportunity to examine patterns of morphological evolution in terrestrial plants and to compare them with patterns previously observed during radiations of marine animals. Focusing on evolution among angiosperm pollen types, I used average pairwise dissimilarity and total variance to describe changing morphological varieties (disparity) through the Cretaceous and Paleocene in North America. Angiosperm species diversity shows an approximately tenfold increase through this interval, but this taxonomic diversification is not matched by a comparable change in morphological disparity. Partitioning of morphological disparity among major pollen groups shows that most of the variance is contributed by eudicots from the Albian onwards. Constant disparity across the Cretaceous/Tertiary boundary despite decreased taxonomic diversity suggests that the Cretaceous/Tertiary extinction was not selective with respect to the pollen morphological characters analyzed here. The two measures of disparity show similar patterns. The overall pattern is robust to changes in character weighting, indicating that no one set of characters or weighting scheme is driving the pattern. Analyses of older data indicate that the initial burst of disparity in the Aptian could be due in part to analytical time-averaging. The observed incongruence between taxonomic diversity and morphological disparity suggests that morphological evolution in pollen was characterized by larger jumps early and smaller jumps later on, and is similar to that found in several groups of marine invertebrates.

Phylogenetic and Genomic Analyses Resolve the Origin of Important Plant Genes Derived from Transposable Elements.

Once perceived as merely selfish, transposable elements (TEs) are now recognized as potent agents of adaptation. One way TEs contribute to evolution is through TE exaptation, a process whereby TEs, which persist by replicating in the genome, transform into novel host genes, which persist by conferring phenotypic benefits. Known exapted TEs (ETEs) contribute diverse and vital functions, and may facilitate punctuated equilibrium, yet little is known about this process. To better understand TE exaptation, we designed an approach to resolve the phylogenetic context and timing of exaptation events and subsequent patterns of ETE diversification. Starting with known ETEs, we search in diverse genomes for basal ETEs and closely related TEs, carefully curate the numerous candidate sequences, and infer detailed phylogenies. To distinguish TEs from ETEs, we also weigh several key genomic characteristics including repetitiveness, terminal repeats, pseudogenic features, and conserved domains. Applying this approach to the well-characterized plant ETEs MUG and FHY3, we show that each group is paraphyletic and we argue that this pattern demonstrates that each originated in not one but multiple exaptation events. These exaptations and subsequent ETE diversification occurred throughout angiosperm evolution including the crown group expansion, the angiosperm radiation, and the primitive evolution of angiosperms. In addition, we detect evidence of several putative novel ETE families. Our findings support the hypothesis that TE exaptation generates novel genes more frequently than is currently thought, often coinciding with key periods of evolution.

INSECT FOLIVORY IN AN ANGIOSPERM-DOMINATED FLORA FROM THE MID-CRETACEOUS OF UTAH, U.S.A.

The Soap Wash flora contains 152 angiosperm leaf fossils from the Albian–Cenomanian boundary of central Utah. The flora exhibits many well-preserved examples of insect folivory for which the extent and type of damage on each leaf was recorded. Fewer than half of the specimens in the collection exhibit insect damage and most of the leaf damage affects less than five percent of the leaf area. Damage was distributed unevenly across the 18 leaf morphotypes described in the flora. Nineteen damage types were recognized. Most of the damage was of a generalized type, dominated by indiscriminate removal of leaf tissues (whole leaf feeding). We observed no correlation between leaf mass per area (MA) and the degree of damage by herbivores, which contrasts with Cenozoic and modern patterns. The low rate, modest area, and low diversity of damage type is similar to patterns reported through the Late Cretaceous and is consistent with a plateau in insect folivore diversity that persisted through the end of the Mesozoic. The preponderance of generalized damage types dominated by whole leaf feeding may reflect (1) the early stage in angiosperm radiation represented by this flora, and (2) the associated lag in folivore diversification and specialization.