Data, metrics, and methods for arthropod and fungal herbivory at the dawn of angiosperm diversification: The Rose Creek plant assemblage of Nebraska, U.S.A.

The data presented in this article are related to the research article titled "Arthropod and fungal herbivory at the dawn of angiosperm diversification: The Rose Creek plant assemblage of Nebraska, U.S.A." (Xiao et al., 2021). These data correspond to an examination of arthropod and fungal herbivory on 2084 plant specimens from the Early Cretaceous (late Albian) Rose Creek locality of southeastern Nebraska, USA. Ten datasets have been assembled to describe and contextualize the diversity and intensity of herbivory at Rose Creek, as documented in Appendices of the online supplementary material. Appendices S4 and S5 provide a list and the frequency distributions by major clade and species/morphotype of all plant taxa examined. Appendix S6 outlines general procedures for documenting herbivory on plants and how the data was acquired. Appendix S9a and S9b provide rarefaction analyses for plant taxa to demonstrate sampling sufficiency, which is paralleled by rarefaction analyses of Appendix S9c and S9d that indicate sampling of damage types are robust. The comprehensive dataset of Appendix S12 lists plant taxa horizontally by major clade/group and species/morphotype versus vertically listed feeding classes, functional feeding groups (FFGs) and damage types (DTs). The basic metrics of DTs, feeding event occurrences, DT host-plant specialization, and number of matrix cells are displayed, with data subtotals and totals. This data matrix serves as the central source of data for the study, and records the six metrics of DT richness, DT frequency, DT host-plant specialization, percent of area herbivorized, and feeding event occurrences. Three of these metrics are used for establishing component community structure of the three most herbivorized taxa (Figs 8-10), and the relationships among plant hosts and FFGs in the non-metric multidimensional scaling analysis (Fig. 11) (Xiao et al., 2021). Appendix S15 is a list DTs, with their assigned host-plant specialization of 1 for generalized, 2 for intermediate specificity, and 3 for specialized. Appendix S16 is a table that provides plant surface areas (cm2) and their percentages that have been removed due to herbivory. Appendix S18 provides descriptions and ancillary data for 14 new DTs described from Rose Creek. A listing of the herbivory index (herbivorized surface area divided by total surface area) of plant assemblages and individual plant species in Appendix S19 provides comparisons among Rose Creek, other fossil, and modern plant assemblages. Lastly, Appendix S23 lists from the literature of arthropod species forming the well-documented herbivore component communities of five modern plant species to the three most herbivorized taxa at Rose Creek shown in Fig. 12. Some of the metrics used to quantitatively measure the diversity and intensity of herbivory are recent, such as feeding event occurrences, whereas others such as herbivorized surface area and host-plant specialization values have had a longer use in plant-arthropod studies.

Early Cretaceous mealybug herbivory on a laurel highlights the deep-time history of angiosperm-scale insect associations.

Insect fluid-feeding on fossil vascular plants is an inconspicuous and underappreciated mode of herbivory that can provide novel data on the evolution of deep-time ecological associations and indicate the host-plant preferences of ancient insect herbivores. Previous fossil studies have documented piercing-and-sucking herbivory but often are unable to identify culprit insect taxa. One line of evidence are punctures and scale-insect impression marks made by piercing-and-sucking insects that occasionally provide clues to the systematic identities and relationships of particular insect herbivores. We report here the earliest occurrences of piercing and sucking on early angiosperms as evidenced by scale insect covers, impression marks, punctures and body fossils - notably a mealybug - from the Lower Cretaceous Rose Creek Flora of the Dakota Formation (c. 103 Ma), in southeastern Nebraska, USA. The mealybug, two other scale insect taxa, and several distinctive damage types on laurel leaves and seed-plant stems at Rose Creek document a diverse guild of piercing-and-sucking insects on early angiosperms. The discovery of an Early Cretaceous female mealybug indicates an early herbivorous association with a laurel host. These data provide direct evidence for co-associations and possible coevolution of scale insects and their plant hosts during early angiosperm diversification.

Occurrence of Phoma Sacc. in the phyllosphere of Neogene Siwalik forest of Arunachal sub-Himalaya and its palaeoecological implications.

The present study reports in situ occurrence of two new epiphyllous fungal species of Phomites (comparable to modern genus Phoma Sacc.) on angiospermic leaf remains recovered from the Siwalik sediments (middle Miocene to early Pleistocene) of Arunachal Pradesh, eastern Himalaya. We describe two new species i.e. Phomites siwalicus Vishnu, Khan et Bera S, sp. nov. and Phomites neogenicus Vishnu, Khan et Bera S, sp. nov. on the basis of structural details of pycnidia. The pycnidium is a globose or slightly lens-shaped, ostiolate with a collar layer consisting of thick walled cells, sunken in leaf cuticle, with one-celled conidiospores and short-ampulliform conidiogenous cells. Host leaves resemble to those of extant Dipterocarpus C. F. Gaertn., Shorea Roxb. ex C. F. Gaertn. (Dipterocarpaceae), Dysoxylum Blume (Meliaceae), and Poaceae Barnhart. In situ occurrence of two Phomites morphotypes on the said leaf remains suggests a possible host-parasite interaction in the moist evergreen forest of Arunachal sub-Himalaya during Mio-Pleistocene period. The occurrence of Phomites in appreciable numbers indicates a humid climate favored by high rate of precipitation during Siwalik sedimentation, which is also consistent with our previously published climatic data obtained from the study of the macroscopic plant remains.

Paleocene Ipomoea (Convolvulaceae) from India with implications for an East Gondwana origin of Convolvulaceae.

The morning glory family, Convolvulaceae, is globally important in medicine and food crops. The family has worldwide distribution in a variety of habitats; however, its fossil record is very poorly documented. The current fossil record suggests an origin in North America, which is in contrast to molecular data that indicate an East Gondwana origin. We report Ipomoea leaves from the late Paleocene (Thanetian; 58.7-55.8 million years ago) of India, which was a part of East Gondwana during this time. This is the earliest fossil record for both the family Convolvulaceae and the order Solanales. This suggests that the sister families Convolvulaceae and Solanaceae diverged before the Eocene in Gondwana-derived continents. The evidence presented here supports the conclusion from molecular phylogenetic analysis of an East Gondwana origin of Convolvulaceae.

The evolutionary convergence of mid-Mesozoic lacewings and Cenozoic butterflies.

Mid-Mesozoic kalligrammatid lacewings (Neuroptera) entered the fossil record 165 million years ago (Ma) and disappeared 45 Ma later. Extant papilionoid butterflies (Lepidoptera) probably originated 80-70 Ma, long after kalligrammatids became extinct. Although poor preservation of kalligrammatid fossils previously prevented their detailed morphological and ecological characterization, we examine new, well-preserved, kalligrammatid fossils from Middle Jurassic and Early Cretaceous sites in northeastern China to unravel a surprising array of similar morphological and ecological features in these two, unrelated clades. We used polarized light and epifluorescence photography, SEM imaging, energy dispersive spectrometry and time-of-flight secondary ion mass spectrometry to examine kalligrammatid fossils and their environment. We mapped the evolution of specific traits onto a kalligrammatid phylogeny and discovered that these extinct lacewings convergently evolved wing eyespots that possibly contained melanin, and wing scales, elongate tubular proboscides, similar feeding styles, and seed-plant associations, similar to butterflies. Long-proboscid kalligrammatid lacewings lived in ecosystems with gymnosperm-insect relationships and likely accessed bennettitalean pollination drops and pollen. This system later was replaced by mid-Cretaceous angiosperms and their insect pollinators.

Montsechia, an ancient aquatic angiosperm.

The early diversification of angiosperms in diverse ecological niches is poorly understood. Some have proposed an origin in a darkened forest habitat and others an open aquatic or near aquatic habitat. The research presented here centers on Montsechia vidalii, first recovered from lithographic limestone deposits in the Pyrenees of Spain more than 100 y ago. This fossil material has been poorly understood and misinterpreted in the past. Now, based upon the study of more than 1,000 carefully prepared specimens, a detailed analysis of Montsechia is presented. The morphology and anatomy of the plant, including aspects of its reproduction, suggest that Montsechia is sister to Ceratophyllum (whenever cladistic analyses are made with or without a backbone). Montsechia was an aquatic angiosperm living and reproducing below the surface of the water, similar to Ceratophyllum. Montsechia is Barremian in age, raising questions about the very early divergence of the Ceratophyllum clade compared with its position as sister to eudicots in many cladistic analyses. Lower Cretaceous aquatic angiosperms, such as Archaefructus and Montsechia, open the possibility that aquatic plants were locally common at a very early stage of angiosperm evolution and that aquatic habitats may have played a major role in the diversification of some early angiosperm lineages.

Fertile structures with in situ spores of a dipterid fern from the Triassic in southern China.

Clathropteris was a typical dipterid fern with well documented fossil record and was widely dispersed during the Mesozoic; however, our knowledge of fertile structures including in situ spores for this genus is still very limited. Here we report well-preserved compression specimens of Clathropteris obovata Oishi from the Late Triassic of Guangyuan, Sichuan Province, China. The specimens show round to oval and exindusiate sori, vertical to oblique annuli in sporangia, and in situ trilete spores with verrucate and baculate sculptures, which are comparable to dispersed spore genera of Converrucosisporites and Conbaculatisporites. Comparisons of relevant fossil taxa suggest that specimens of C. obovata from Triassic of China provide for the first time in Asia the detailed fertile structures with in situ spore characters of dipterid fossil Clathropteris. Unlike living Dipteris, Mesozoic fossils of Dipteridaceae show a high diversity and a range of complex morphology of in situ spores, thus are significant for the evolutionary links between Dipteridaceae and other related fern clade, including Gleicheniaceae and Matoniaceae of the Gleicheniales.

Coryphoid palm leaf fossils from the Maastrichtian-Danian of Central India with remarks on phytogeography of the Coryphoideae (Arecaceae).

PREMISE OF RESEARCH: A large number of fossil coryphoid palm wood and fruits have been reported from the Deccan Intertrappean beds of India. We document the oldest well-preserved and very rare costapalmate palm leaves and inflorescence like structures from the same horizon. METHODOLOGY: A number of specimens were collected from Maastrichtian-Danian sediments of the Deccan Intertrappean beds, Ghughua, near Umaria, Dindori District, Madhya Pradesh, India. The specimens are compared with modern and fossil taxa of the family Arecaceae. PIVOTAL RESULTS: Sabalites dindoriensis sp. nov. is described based on fossil leaf specimens including basal to apical parts. These are the oldest coryphoid fossil palm leaves from India as well as, at the time of deposition, from the Gondwana- derived continents. CONCLUSIONS: The fossil record of coryphoid palm leaves presented here and reported from the Eurasian localities suggests that this is the oldest record of coryphoid palm leaves from India and also from the Gondwana- derived continents suggesting that the coryphoid palms were well established and wide spread on both northern and southern hemispheres by the Maastrichtian-Danian. The coryphoid palms probably dispersed into India from Europe via Africa during the latest Cretaceous long before the Indian Plate collided with the Eurasian Plate.

Fruit structure in Magnoliaceae s.l. and Archaeanthus and their relationships.

PREMISE OF THE STUDY: The family Magnoliaceae s.l. is a basal angiosperm family with two subfamilies-Magnolioideae and Liriodendroideae, which differ by the types and structure of their fruits and seeds. The late Albian genus Archaeanthus shares many features of its reproductive organs with Magnoliaceae s.l., but its pericarp anatomy was never studied in detail. A broad-scale carpological investigation of Archaeanthus and Magnoliaceae s.l. was undertaken to reveal the nature of the similarities in fruit structure and to reconstruct Archaeanthus pericarp anatomy. These data are important to determine the early stages of fruit morphogenesis and thus to clarify relationships of Archaeanthus to the taxa of Magnoliaceae s.l. METHODS: The pericarp anatomy was studied with light microscopy, SEM, and polarizing microscopy. KEY RESULTS: The dehiscent, polyspermous follicles shed from the receptacle of Archaeanthus share similarities with dehiscent follicles of Magnoliaceae s.s. and shedding nutlets of Liriodendron. The seeds of Archaeanthus are dorsiventrally flattened, ovoid, and encircled with a single circular wing. The pericarps of all the taxa studied are differentiated into exocarp (epidermis), multilayered mesocarp, and endocarp (fiber-like sclereids). The mesocarp consists of parenchyma with scattered secretory cells and sclereid clusters (Magnoliaceae s.s., Archaeanthus) or composed by sclerenchyma (Liriodendron). CONCLUSIONS: The specializations of dehiscent multifollicles of unknown Cretaceous ancestors for different modes of seed and fruitlet dispersal formed the basis for the differentiation of two evolutionary lines with their divergence occurring more than 100 million years ago: Magnoliaceae s.s. and the Archaeanthus-Liriodendroidea-Liriodendron line (Liriodendraceae s.l.) within the order Magnoliales.

Rise to dominance of angiosperm pioneers in European Cretaceous environments.

The majority of environments are dominated by flowering plants today, but it is uncertain how this dominance originated. This increase in angiosperm diversity happened during the Cretaceous period (ca. 145-65 Ma) and led to replacement and often extinction of gymnosperms and ferns. We propose a scenario for the rise to dominance of the angiosperms from the Barremian (ca. 130 Ma) to the Campanian (ca. 84 Ma) based on the European megafossil plant record. These megafossil data demonstrate that angiosperms migrated into new environments in three phases: (i) Barremian (ca. 130-125 Ma) freshwater lake-related wetlands; (ii) Aptian-Albian (ca. 125-100 Ma) understory floodplains (excluding levees and back swamps); and (iii) Cenomanian-Campanian (ca. 100-84 Ma) natural levees, back swamps, and coastal swamps. This scenario allows for the measured evolution of angiosperms in time and space synthesizing changes in the physical environment with concomitant changes in the biological environment. This view of angiosperm radiation in three phases reconciles previous scenarios based on the North American record. The Cretaceous plant record that can be observed in Europe is exceptional in many ways. (i) Angiosperms are well preserved from the Barremian to the Maastrichtian (ca. 65 Ma). (ii) Deposits are well constrained and dated stratigraphically. (iii) They encompass a full range of environments. (iv) European paleobotany provides many detailed studies of Cretaceous floras for analysis. These factors make a robust dataset for the study of angiosperm evolution from the Barremian to the Campanian that can be traced through various ecosystems and related to other plant groups occupying the same niches.

Palynological composition of a Lower Cretaceous South American tropical sequence: climatic implications and diversity comparisons with other latitudes.

PREMISE OF THE STUDY: Reconstruction of floristic patterns during the early diversification of angiosperms is impeded by the scarce fossil record, especially in tropical latitudes. Here we collected quantitative palynological data from a stratigraphic sequence in tropical South America to provide floristic and climatic insights into such tropical environments during the Early Cretaceous. METHODS: We reconstructed the floristic composition of an Aptian-Albian tropical sequence from central Colombia using quantitative palynology (rarefied species richness and abundance) and used it to infer its predominant climatic conditions. Additionally, we compared our results with available quantitative data from three other sequences encompassing 70 floristic assemblages to determine latitudinal diversity patterns. KEY RESULTS: Abundance of humidity indicators was higher than that of aridity indicators (61% vs. 10%). Additionally, we found an angiosperm latitudinal diversity gradient (LDG) for the Aptian, but not for the Albian, and an inverted LDG of the overall diversity for the Albian. Angiosperm species turnover during the Albian, however, was higher in humid tropics. CONCLUSIONS: There were humid climates in northwestern South America during the Aptian-Albian interval contrary to the widespread aridity expected for the tropical belt. The Albian inverted overall LDG is produced by a faster increase in per-sample angiosperm and pteridophyte diversity in temperate latitudes. However, humid tropical sequences had higher rates of floristic turnover suggesting a higher degree of morphological variation than in temperate regions.

A eudicot from the Early Cretaceous of China.

The current molecular systematics of angiosperms recognizes the basal angiosperms and five major angiosperm lineages: the Chloranthaceae, the magnoliids, the monocots, Ceratophyllum and the eudicots, which consist of the basal eudicots and the core eudicots. The eudicots form the majority of the angiosperms in the world today. The flowering plants are of exceptional evolutionary interest because of their diversity of over 250,000 species and their abundance as the dominant vegetation in most terrestrial ecosystems, but little is known of their very early history. In this report we document an early presence of eudicots during the Early Cretaceous Period. Diagnostic characters of the eudicot fossil Leefructus gen. nov. include simple and deeply trilobate leaves clustered at the nodes in threes or fours, basal palinactinodromous primary venation, pinnate secondary venation, and a long axillary reproductive axis terminating in a flattened receptacle bearing five long, narrow pseudo-syncarpous carpels. These morphological characters suggest that its affinities are with the Ranunculaceae, a basal eudicot family. The fossil co-occurs with Archaefructus sinensis and Hyrcantha decussata whereas Archaefructus liaoningensis comes from more ancient sediments. Multiple radiometric dates of the Lower Cretaceous Yixian Formation place the bed yielding this fossil at 122.6-125.8 million years old. The earliest fossil records of eudicots are 127 to 125 million years old, on the basis of pollen. Thus, Leefructus gen. nov. suggests that the basal eudicots were already present and diverse by the latest Barremian and earliest Aptian.

Global CO2 rise leads to reduced maximum stomatal conductance in Florida vegetation.

A principle response of C3 plants to increasing concentrations of atmospheric CO(2) (CO(2)) is to reduce transpirational water loss by decreasing stomatal conductance (g(s)) and simultaneously increase assimilation rates. Via this adaptation, vegetation has the ability to alter hydrology and climate. Therefore, it is important to determine the adaptation of vegetation to the expected anthropogenic rise in CO(2). Short-term stomatal opening-closing responses of vegetation to increasing CO(2) are described by free-air carbon enrichments growth experiments, and evolutionary adaptations are known from the geological record. However, to date the effects of decadal to centennial CO(2) perturbations on stomatal conductance are still largely unknown. Here we reconstruct a 34% (±12%) reduction in maximum stomatal conductance (g(smax)) per 100 ppm CO(2) increase as a result of the adaptation in stomatal density (D) and pore size at maximal stomatal opening (a(max)) of nine common species from Florida over the past 150 y. The species-specific g(smax) values are determined by different evolutionary development, whereby the angiosperms sampled generally have numerous small stomata and high g(smax), and the conifers and fern have few large stomata and lower g(smax). Although angiosperms and conifers use different D and a(max) adaptation strategies, our data show a coherent response in g(smax) to CO(2) rise of the past century. Understanding these adaptations of C3 plants to rising CO(2) after decadal to centennial environmental changes is essential for quantification of plant physiological forcing at timescales relevant for global warming, and they are likely to continue until the limits of their phenotypic plasticity are reached.

Climate forcing due to optimization of maximal leaf conductance in subtropical vegetation under rising CO2.

Plant physiological adaptation to the global rise in atmospheric CO(2) concentration (CO(2)) is identified as a crucial climatic forcing. To optimize functioning under rising CO(2), plants reduce the diffusive stomatal conductance of their leaves (g(s)) dynamically by closing stomata and structurally by growing leaves with altered stomatal densities and pore sizes. The structural adaptations reduce maximal stomatal conductance (g(smax)) and constrain the dynamic responses of g(s). Here, we develop and validate models that simulate structural stomatal adaptations based on diffusion of CO(2) and water vapor through stomata, photosynthesis, and optimization of carbon gain under the constraint of a plant physiological cost of water loss. We propose that the ongoing optimization of g(smax) is eventually limited by species-specific limits to phenotypic plasticity. Our model reproduces observed structural stomatal adaptations and predicts that adaptation will continue beyond double CO(2). Owing to their distinct stomatal dimensions, angiosperms reach their phenotypic response limits on average at 740 ppm and conifers on average at 1,250 ppm CO(2). Further, our simulations predict that doubling today's CO(2) will decrease the annual transpiration flux of subtropical vegetation in Florida by ≈60 W·m(-2). We conclude that plant adaptation to rising CO(2) is altering the freshwater cycle and climate and will continue to do so throughout this century.

Fruits and foliage of Pueraria (Leguminosae, Papilionoideae) from the Neogene of Eurasia and their biogeographic implications.

PREMISE OF THE STUDY: Pueraria (Leguminosae, Papilionoideae) is native in East Asia, South Asia, Southeast Asia, and Oceania and is well known as a rampant invasive weed in the southeastern United States (P. montana; better known as kudzu), but relatively little is known about its early evolution and biogeographic origin. • METHODS: On the basis of comparative analyses of the fruit and leaflet architecture of closely related extant and fossil taxa, we studied the fossil history and biogeography of Pueraria. • KEY RESULTS: Fossil Pueraria is recognized on the basis of distinctive fruit and foliage from the Mio-Pliocene of middle latitudes in China, Japan, Abkhazia, and Croatia. Recognition of P. miothunbergiana from the Mio-Pliocene of China and Japan is reinforced by a trifoliolate leaf as well as isolated lateral and terminal leaflets. Pueraria shanwangensis sp. nov. represents the first recognition of fossil Pueraria fruits. This fruit species co-occurs with P. miothunbergiana in the Middle Miocene Shanwang flora and possibly represents the same population. Pueraria maxima (Unger) comb. nov., previously named as Dolichites maximus or Desmodium maximum, is recognized on the basis of leaflets from the Miocene of Croatia and Abkhazia. Other prior fossil reports of Pueraria and Dolichites are reevaluated. • CONCLUSIONS: Pueraria had begun to diversify by at least the Middle Miocene and had spread into the Mio-Pliocene subtropical and temperate floras of the Balkan Peninsula, the Caucasus, and eastern Asia, which suggests the present diversity of this genus in tropical Asia and Oceania might have originated from the mid-latitudes of Eurasia.

Tertiary leaf fossils of Mangifera (Anacardiaceae) from Li Basin, Thailand as examples of the utility of leaf marginal venation characters.

The mango (Mangifera) is the most important commercial fruit in Asia and is popular worldwide. Because of uncertainty in its origin and biogeography, the study of Mangifera could be of value in elucidating its current genetic diversity and geographical distribution. We report here two new species of compressed leaves of Mangifera (Anacardiaceae) from upper Oligocene-lower Miocene lacustrine sediments in the Li Basin, northern Thailand. Leaf form and venation characters of fossil and extant leaves are presented with detailed marginal venation patterns that were found to be important in assigning the fossil leaves to Mangifera paleoindica and M. paleocaloneura. These two fossil species of Mangifera and the present occurrence of basal species of Mangifera in Thailand suggest that northern Thailand could be an area with a long history of evolution and diversification of Mangifera, particularly M. indica.

An Early Cretaceous fruit with affinities to Ceratophyllaceae.

A new genus and species with affinities to Ceratophyllaceae from the Lower Cretaceous of Kansas, USA is reported. The fruits of Donlesia dakotensis gen. et sp. nov. are smooth achenes with two winged lateral spines, two winged facial spines, one stylar spine, and a long peduncle with a distinctive groove. The facial spines are arranged perpendicular to the lateral spines. The seed contains two cotyledons and a central plumule. Dichotomous leaves associated with Donlesia dakotensis fruits are whorled, petiolate, and lack denticles. The fruit morphology of D. dakotensis differs from extant and fossil members of Ceratophyllaceae in its tetra-radial symmetry, the presence of two winged facial spines, a long peduncle, and the lack of other appendages on the fruit surface. It seems to be more closely related to section Ceratophyllum of extant Ceratophyllum. This relationship suggests that section Ceratophyllum of extant Ceratophyllum may represent the ancestral members of the ancient Ceratophyllum lineage. The presence of this species in the Dakota Formation further confirms that there were diverse aquatic plants living in freshwater lakes along the eastern margin of the Western Interior Seaway during the Early Cretaceous.

A climatic and taxonomic comparison between leaf litter and standing vegetation from a Florida swamp woodland.

One method to determine past climate has been the use of leaf morphological characteristics of fossil leaves quantified using modern climate and canopy leaf characteristics. Fossil assemblages are composed of abscised leaves, and climate may be more accurately determined by using leaves from leaf litter instead of the canopy. To better understand whether taphonomic processes make a difference in this relationship, a north-central Florida woodland was sampled to determine the morphologically based climate estimates from these leaves. Leaves from woody, dicotyledonous plants were collected and identified, then compared using presence/absence data and analyzed using several linear regression equations and the CLAMP data set. Although the majority of standing vegetation was reflected in leaf litter, some inconsistencies were observed, which may reflect plant community structure or sampling technique. Mean annual temperature (MAT) and growing season precipitation (GSP) were estimated from leaf litter morphological characters and living leaves. Overall, values for MAT estimated from litter and living leaves were cooler than actual MATs, although several accurate and high estimates were obtained depending on the predictive method used. Estimated GSP values were higher than actual GSPs. Statistically, no difference was observed between MAT and GSP estimates derived from leaf litter vs. estimates derived from living leaves, with one exception.

Early steps of angiosperm pollinator coevolution.

The hypothesis that early flowering plants were insect-pollinated could be tested by an examination of the pollination biology of basal angiosperms and the pollination modes of fossil angiosperms. We provide data to show that early fossil angiosperms were insect-pollinated. Eighty-six percent of 29 extant basal angiosperm families have species that are zoophilous (of which 34% are specialized) and 17% of the families have species that are wind-pollinated, whereas basal eudicot families and basal monocot families more commonly have wind and specialized pollination modes (up to 78%). Character reconstruction based on recent molecular trees of angiosperms suggests that the most parsimonious result is that zoophily is the ancestral state. Combining pollen ornamentation, size, and aperture characteristics and the abundance of single-species pollen clumps of Cenomanian angiosperm-dispersed pollen species from the Dakota Formation demonstrates a dominance of zoophilous pollination (76% versus 24% wind pollination). The zoophilous pollen species have adaptations for pollination by generalist insects (39%), specialized pollen-collecting insects (27%), and other specialized pollinators (10%). These data quantify the presences of more specialized pollination modes during the mid-Cretaceous angiosperm diversification.

The impact of Miocene atmospheric carbon dioxide fluctuations on climate and the evolution of terrestrial ecosystems.

The Miocene is characterized by a series of key climatic events that led to the founding of the late Cenozoic icehouse mode and the dawn of modern biota. The processes that caused these developments, and particularly the role of atmospheric CO2 as a forcing factor, are poorly understood. Here we present a CO2 record based on stomatal frequency data from multiple tree species. Our data show striking CO2 fluctuations of approximately 600-300 parts per million by volume (ppmv). Periods of low CO2 are contemporaneous with major glaciations, whereas elevated CO2 of 500 ppmv coincides with the climatic optimum in the Miocene. Our data point to a long-term coupling between atmospheric CO2 and climate. Major changes in Miocene terrestrial ecosystems, such as the expansion of grasslands and radiations among terrestrial herbivores such as horses, can be linked to these marked fluctuations in CO2.