Expansion Of Angiosperms Research Articles

Geographical and life-history traits associated with low and high species richness across angiosperm families.

The phenomenal expansion of angiosperms has prompted many investigations into the factors driving their diversification, but there remain significant gaps in our understanding of flowering plant species diversity. Using the crown age of families from five studies, we used a maximum likelihood approach to classify families as having poor, predicted or high species richness (SR) using strict consensus criteria. Using these categories, we looked for associations between family SR and i) the presence of an inferred familial ancestral polyploidization event, ii) 23 life history and floral traits compiled from previously published datasets and papers, and iii) sexual system (dioecy) or genetically determined self-incompatibility (SI) mating system using an updated version of our own database and iv) geographic distribution using a new database describing the global distribution of plant species/families across realms and biomes and inferred range. We find that more than a third of angiosperm families (65%) had predicted SR, a large proportion (30.2%) were species poor, while few (4.8%) had high SR. Families with poor SR were less likely to have undergone an ancestral polyploidization event, exhibited deficits in diverse traits, and were more likely to have unknown breeding systems and to be found in only one or few biomes and realms, especially the Afrotropics or Australasia. On the other hand, families with high SR were more likely to have animal mediated pollination or dispersal, are enriched for epiphytes and taxa with an annual life history, and were more likely to harbour sporophytic SI systems. Mapping the global distribution of georeferenced taxa by their family DR, we find evidence of regions dominated by taxa from lineages with high vs low SR. These results are discussed within the context of the literature describing "depauperons" and the factors contributing to low and high biodiversity in angiosperm clades.

A complete genus-level phylogeny reveals the Cretaceous biogeographic diversification of the poppy family

Angiosperms, a trigger for the Cretaceous Terrestrial Revolution (KTR), underwent a rapid expansion and occupied all the environments during the Mid-Upper Cretaceous. Yet, Cretaceous biogeographic patterns and processes underlying the distribution of angiosperm diversity in the Northern Hemisphere are still poorly known. Here, we elucidated the biogeographic diversification of the angiosperm family Papaveraceae, an ancient Northern Hemisphere clade characterized by poor dispersal ability and high level of regional endemism. Based on both plastome and multi-locus datasets, we reconstructed a robust time-calibrated phylogeny that includes all currently recognized 45 genera of this family. Within the time-calibrated phylogenetic framework, we conducted 72 biogeographic analyses by testing the sensitivity of uncertainties of area delimitation, maxarea constraints, and the parameters of the model, i.e., j (describing jump-dispersal events) and w (modifying dispersal multiplier matrices), to ancestral range estimations. We also inferred ancestral habitat and ecological niches. Phylogenetic analyses strongly support Papaveraceae as monophyletic. Pteridophylloideae is strongly supported as sister to Hypecoideae-Fumarioideae. Our results indicate that the j parameter and number of predefined areas strongly affect ancestral range estimates, generating questionable ancestral ranges, whereas maxarea constraint and w parameter have no effect and improve model fit. After accounting for these uncertainties, our results indicate that Papaveraceae differentiated in Asian wet forests during the Lower Cretaceous and subsequently occupied the Asian and western North American arid and open areas. Three dispersals from Asia to western North America via the Bering land bridge occurred in the Mid-Upper Cretaceous, largely in agreement with the KTR. Habitat shift and ecological niche divergence resulted in the subsequent disjunctions between Asia and western North America. These findings suggest that the interplay of range expansion and niche divergence-driven vicariance might have shaped Cretaceous biogeographic patterns of angiosperms with Papaveraceae-like ecological requirements and dispersal abilities in the Northern Hemisphere, hence contributing to the knowledge on the geographic expansion of angiosperms during the KTR.

Identification and Analysis of Stress-Associated Proteins (SAPs) Protein Family and Drought Tolerance of ZmSAP8 in Transgenic Arabidopsis.

Stress-associated proteins (SAPs), a class of A20/AN1 zinc finger proteins, play vital roles in plant stress response. However, investigation of SAPs in maize has been very limited. Herein, to better trace the evolutionary history of SAPs in maize and plants, 415 SAPs were identified in 33 plant species and four species of other kingdoms. Moreover, gene duplication mode exploration showed whole genome duplication contributed largely to SAP gene expansion in angiosperms. Phylogeny reconstruction was performed with all identified SAPs by the maximum likelihood (ML) method and the SAPs were divided into five clades. SAPs within the same clades showed conserved domain composition. Focusing on maize, nine ZmSAPs were identified. Further promoter cis-elements and stress-induced expression pattern analysis of ZmSAPs indicated that ZmSAP8 was a promising candidate in response to drought stress, which was the only AN1-AN1-C2H2-C2H2 type SAP in maize and belonged to clade I. Additionally, ZmSAP8 was located in the nucleus and had no transactivation activity in yeast. Overexpressing ZmSAP8 enhanced the tolerance to drought stress in Arabidopsis thaliana, with higher seed germination and longer root length. Our results should benefit the further functional characterization of ZmSAPs.

Permian lobed Zoophycos as the product of the terrestrialization process: Behavioral innovation in the Tahkandit Limestone (Yukon River, Alaska, USA)

Paleontological survey in the remote Yukon-Charley Rivers National Preserve in Alaska led to the discovery of lobed Zoophycos from the lower Tahkandit Limestone (informally named Sandstone unit), an interval characterized by grayish-green glauconitic sandstone and conglomerate of coastal origin. The studied Zoophycos consists of a lobate skirt-like spreite bounded by a marginal tube. Smaller tongue-shaped lobes branch off from larger parent lobes that share the same tongue-like shape. Sedimentological features, together with body fossils and associated trace fossils ( Planolites, Chondrites ), indicate a shoreface habitat for the Zoophycos producer. This shallow-marine environmental setting is in contrast with the deeper bathymetries in which lobed Zoophycos are recovered in post-Palaeozoic times. The producer of the lobed Zoophycos of the Yukon River is interpreted as a deposit-feeder that used sensory-driven, directed search for locating heterogeneously distributed trophic resources. The Zoophycos producer filled its burrow with Coprolus- like fecal pellets, possibly complementing deposit feeding with microbial gardening and/or food caching. Data presented here provide useful insight into the morphological evolution and bathymetric distribution of Zoophycos , suggesting two ‘Golden Ages’ for lobed Zoophycos: (1) Carboniferous–Permian and (2) Cretaceous–Neogene. This stratigraphic distribution supports the important ecological role of major terrestrialization events, that are, the Palaeozoic expansion of land plants and the Mesozoic expansion of angiosperms. The consequent increased input of nutrients to coastal areas was an important contributor to declining trends in porewater oxygen concentrations. This phenomenon favored adaptive traits to exploit nutrient-rich but oxygen-poor niches, among which the U-shaped marginal tube of lobed Zoophycos was an efficient adaptation to bring oxygenated water into low-oxygen substrates. • New ichnological information is added to a poorly understood Permian rock unit along the Yukon River in eastern Alaska. • This report illustrates the important role of terrestralization events in driving the Zoophycos morphology. • In contrast to previous work, this study shows larger spirals of Zoophycos preserved in more inshore environments. • This is one of the oldest known trace morphologies of Zoophycos with such a complex vertical structure.

Admixture of divergent genomes facilitates hybridization across species in the family Brassicaceae.

Summary Hybridization and polyploidization are pivotal to plant evolution. Genetic crosses between distantly related species are rare in nature due to reproductive barriers but how such hurdles can be overcome is largely unknown. Here we report the hybrid genome structure of xBrassicoraphanus, a synthetic allotetraploid of Brassica rapa and Raphanus sativus.We performed cytogenetic analysis and de novo genome assembly to examine chromosome behaviors and genome integrity in the hybrid. Transcriptome analysis was conducted to investigate expression of duplicated genes in conjunction with epigenome analysis to address whether genome admixture entails epigenetic reconfiguration.Allotetraploid xBrassicoraphanus retains both parental chromosomes without genome rearrangement. Meiotic synapsis formation and chromosome exchange are avoided between nonhomologous progenitor chromosomes. Reconfiguration of transcription network occurs, and less divergent cis‐elements of duplicated genes are associated with convergent expression. Genome‐wide DNA methylation asymmetry between progenitors is largely maintained but, notably, B. rapa‐originated transposable elements are transcriptionally silenced in xBrassicoraphanus through gain of DNA methylation.Our results demonstrate that hybrid genome stabilization and transcription compatibility necessitate epigenome landscape adjustment and rewiring of cis–trans interactions. Overall, this study suggests that a certain extent of genome divergence facilitates hybridization across species, which may explain the great diversification and expansion of angiosperms during evolution.

Historia biogeográfica de las briófitas de Chile

Some biogeographical features of bryophytes associated with subtropical-temperate ecosystems in Chile are examined, including diversity, concentration of species richness, endemism, biogeographical composition and disjunctions. The historical-biogeographical interpretation of these characteristics considers the major evolutionary changes of the group in three different temporal scenarios, according to geological, paleobotanical and molecular evidences. First, simple- and complex-thalloids liverworts represented in Chile are examined, belonging to the Classes Haplomitriopsida and Marchantiopsida- Marchantiidae, archaic lineages linked to the pre-Cenozoic colonization of the Pangea mega-continent. Examples of simple-thalloids liverworts of Class Jungermanniopsida-Pelliidae, hornworts of the Class Anthocerotopsida and mosses of the Classes Bryopsida and Polytrichopsida illustrate the paleo-austral distribution patterns of bryophytes from the temperate region of Chile, related to the Gondwana breakup and expansion of Angiosperms during the Cretaceous/Paleogene. A second scenario, considers the final uplift of the Andes and the development of the Arid Diagonal of South America during the Neogene, processes that have played a major role in the biogeographical structure of the Chilean and South American floras. In this context, leafy liverworts associated with the relict forests of the Chilean semi- arid coast stand out, particularly epiphyllous liverworts of the two most diverse families of the Class Jungermanniopsida-Jungermanniidae, Lejeuneaceae and Plagiochilaceae, adapted to microclimates favored by coastal fogs. The last scenario discusses the significant role of the Pleistocene glacial cycles in the expansion of the sub-Antarctic temperate-cold element throughout central-southern Chile, and its effects on the concentration of bryophyte richness associated with Chile’s temperate-rain ecosystems.

Molecular evolution of GDP-L-galactose phosphorylase, a key regulatory gene in plant ascorbate biosynthesis.

Ascorbic acid (AsA) is a widespread antioxidant in living organisms, and plays essential roles in the growth and development of animals and plants as well as in the response to abiotic stress tolerance. The GDP-L-galactose phosphorylase (GGP) is a key regulatory gene in plant AsA biosynthesis that can regulate the concentration of AsA at the transcriptional and translational levels. The function and regulation mechanisms of GGP have been well understood; however, the molecular evolutionary patterns of the gene remain unclear. In this study, a total of 149 homologous sequences of GGP were sampled from 71 plant species covering the major groups of Viridiplantae, and the phylogenetic relationships, gene duplication and molecular evolution analyses of the genes were systematically investigated. Results showed that GGP genes are present throughout the plant kingdom and five shared whole-genome duplications and several lineage-specific whole-genome duplications were found, which led to the rapid expansion of GGPs in seed plants, especially in angiosperms. The structure of GGP genes was more conserved in land plants, but varied greatly in green algae, indicating that GGP may have undergone great differentiation in the early stages of plant evolution. Most GGP proteins had a conserved motif arrangement and composition, suggesting that plant GGPs have similar catalytic functions. Molecular evolutionary analyses showed that GGP genes were predominated by purifying selection, indicating that the gene is functionally conserved due to its vital importance in AsA biosynthesis. Most of the branches under positive selection identified by the branch-site model were mainly in the chlorophytes lineage, indicating episodic diversifying selection may contribute to the evolution of GGPs, especially in the chlorophyte lineage. The conserved function of GGP and its rapid expansion in angiosperms maybe one of the reasons for the increase of AsA content in angiosperms, enabling angiosperms to adapt to changing environments.

The rise of flowering plants in the high southern latitudes of Australia

The Early Cretaceous high-paleolatitude palynofloras from the Otway and Gippsland basins of southeastern Australia contain diverse angiosperm assemblages not described previously. Clavatipollenites hughesii recovered from the early Aptian Upper Cyclosporites hughesii subzone in the Gippsland Basin represents the first record of angiosperm pollen in Australia, coeval to records recovered from the Great Artesian and North West Shelf basins of northern Australia. Tricolpate pollen including Tricolpites variabilis, Rousea georgensis and Striatopollis spp., are first recorded in the late Albian Upper Coptospora paradoxa subzone in southeastern Australia. This represents the second oldest occurrence of tricolpate pollen in Australia, the first occurring in the older middle Albian Lower Coptospora paradoxa subzone in the northern Great Artesian Basin. By the latest Albian Phimopollenites pannosus Zone angiosperms had diversified rapidly in southeastern Australia. The delayed appearance, rise in abundance and diversification of eudicot angiosperms in the high-latitude southern basins of Australia, relative to low- and mid-latitude settings, supports a latitudinally diachronous pattern of angiosperm range expansion from warmer paleoequatorial regions to relatively cooler high-latitude settings. Increasing mean annual temperatures globally in the late Albian likely facilitated the expansion of angiosperms into high-latitude settings in the Southern and Northern hemispheres.

Canopy structure in Late Cretaceous and Paleocene forests as reconstructed from carbon isotope analyses of fossil leaves

Abstract While modern forests have their origin in the diversification and expansion of angiosperms in the Late Cretaceous and early Cenozoic, it is unclear whether the rise of closed-canopy tropical rainforests preceded or followed the end-Cretaceous extinction. The “canopy effect” is a strong vertical gradient in the carbon isotope (δ13C) composition of leaves in modern closed-canopy forests that could serve as a proxy signature for canopy structure in ancient forests. To test this, we report measurements of the carbon isotope composition of nearly 200 fossil angiosperm leaves from two localities in the Paleocene Cerrejón Formation and one locality in the Maastrichtian Guaduas Formation of Colombia. Leaves from one Cerrejón fossil assemblage deposited in a small fluvial channel exhibited a 6.3‰ range in δ13C, consistent with a closed-canopy forest. Carbon isotope values from lacustrine sediments in the Cerrejón Formation had a range of 3.3‰, consistent with vegetation along a lake edge. An even-narrower range of δ13C values (2.7‰) was observed for a leaf assemblage recovered from the Cretaceous Guaduas Formation, and suggests vegetation with an open canopy structure. Carbon isotope fractionation by Late Cretaceous and early Paleogene leaves was in all cases similar to that by modern relatives, consistent with estimates of low atmospheric CO2 during this time period. This study confirms other lines of evidence suggesting that closed-canopy forests in tropical South America existed by the late Paleocene, and fails to find isotopic evidence for a closed-canopy forest in the Cretaceous.

Synopsis of the SOFL Plant-Specific Gene Family.

SUPPRESSOR OF PHYB-4#5DOMINANT (sob5-D) was previously identified as a suppressor of the phyB-4 long-hypocotyl phenotype in Arabidopsis thaliana. Overexpression of SOB5 conferred dwarf phenotypes similar to those observed in plants containing elevated levels of cytokinin (CK) nucleotides and nucleosides. Two SOB-FIVE- LIKE (SOFL) proteins, AtSOFL1 and AtSOFL2, which are more similar at the protein level to each other than they are to SOB5, conferred similar phenotypes to the sob5-D mutant when overexpressed. We used protein sequences of founding SOFL gene family members to perform database searches and identified a total of 289 SOFL homologs in genomes of 89 angiosperm species. Phylogenetic analysis results implied that the SOFL gene family emerged during the expansion of angiosperms and later evolved into four distinct clades. Among the newly identified gene family members are four previously unreported Arabidopsis SOFLs. Multiple sequence alignment of the 289 SOFL protein sequences revealed two highly conserved domains; SOFL-A and SOFL-B. We used overexpression and site-directed mutagenesis studies to demonstrate that SOFL domains are necessary for SOB5 and AtSOFL1’s overexpression phenotypes. Examination of the subcellular localization patterns of founding Arabidopsis thaliana SOFLs suggested they may be localized in the cytoplasm and/or the nucleus. Overall, we report that SOFLs are a plant-specific gene family characterized by two conserved domains that are important for function.

The case of Darwinylus marcosi (Insecta: Coleoptera: Oedemeridae): A Cretaceous shift from a gymnosperm to an angiosperm pollinator mutualism

ABSTRACTAbundant gymnosperm pollen grains associated with the oedemerid beetle Darwinylus marcosi Peris, 2016 were found in Early Cretaceous amber from Spain. This discovery provides confirmatory evidence for a pollination mutualism during the mid Mesozoic for the family Oedemeridae (Coleoptera), which today is known to pollinate only angiosperms. As a result, this new record documents a lateral host-plant transfer from an earlier gymnosperm to a later angiosperm, indicating that pollination of the latter is a derived condition within Oedemeridae. This new fossil record exemplifies one of the 4 ecological-evolutionary pollinator cohorts now known to have existed during the global shift from a gymnosperm to an angiosperm dominated global flora. Currently, all direct evidence for pollination during the 35 million-year interval of the mid Cretaceous gymnosperm-to-angiosperm transition entails recognition of gymnosperm pollen grains on insect mouthparts and other body contact surfaces, while analogous records involving angiosperms are lacking. The gathering evidence indicates that angiosperm pollination was preceded by at least 4 gymnosperm pollination modes that served as a functional and ecological prelude to the rise and expansion of angiosperms.

Changes to Cretaceous surface fire behaviour influenced the spread of the early angiosperms.

Summary Angiosperms evolved and diversified during the Cretaceous period. Early angiosperms were short‐stature weedy plants thought to have increased fire frequency and mortality in gymnosperm forest, aiding their own expansion. However, no explorations have considered whether the range of novel fuel types that diversified throughout the Cretaceous also altered fire behaviour, which should link more strongly to mortality than fire frequency alone.We measured ignitability and heat of combustion in analogue Cretaceous understorey fuels (conifer litter, ferns, weedy and shrubby angiosperms) and used these data to model palaeofire behaviour.Variations in ignition, driven by weedy angiosperms alone, were found to have been a less important feedback to changes in Cretaceous fire activity than previously estimated. Our model estimates suggest that fires in shrub and fern understories had significantly greater fireline intensities than those fuelled by conifer litter or weedy angiosperms, and whilst fern understories supported the most rapid fire spread, angiosperm shrubs delivered the largest amount of heat per unit area.The higher fireline intensities predicted by the models led to estimates of enhanced scorch of the gymnosperm canopy and a greater chance of transitioning to crown fires. Therefore, changes in fire behaviour driven by the addition of new Cretaceous fuel groups may have assisted the angiosperm expansion.

Tectonic-driven climate change and the diversification of angiosperms.

In 1879, Charles Darwin characterized the sudden and unexplained rise of angiosperms during the Cretaceous as an "abominable mystery." The diversification of this clade marked the beginning of a rapid transition among Mesozoic ecosystems and floras formerly dominated by ferns, conifers, and cycads. Although the role of environmental factors has been suggested [Coiffard C, Gómez B (2012) Geol Acta 10(2):181-188], Cretaceous global climate change has barely been considered as a contributor to angiosperm radiation, and focus was put on biotic factors to explain this transition. Here we use a fully coupled climate model driven by Mesozoic paleogeographic maps to quantify and discuss the impact of continental drift on angiosperm expansion and diversification. We show that the decrease of desertic belts between the Triassic and the Cretaceous and the subsequent onset of long-lasting humid conditions during the Late Cretaceous were driven by the breakup of Pangea and were contemporaneous with the first rise of angiosperm diversification. Positioning angiosperm-bearing fossil sites on our paleobioclimatic maps shows a strong match between the location of fossil-rich outcrops and temperate humid zones, indicating that climate change from arid to temperate dominance may have set the stage for the ecological expansion of flowering plants.

The enigma of the rise of angiosperms: can we untie the knot?

Multiple hypotheses have been put forward to explain the rise of angiosperms to ecological dominance following the Cretaceous. A unified scheme incorporating all these theories appears to be an inextricable knot of relationships, processes and plant traits. Here, we revisit these hypotheses, categorising them within frameworks based on plant carbon economy, resistance to climatic stresses, nutrient economy, biotic interactions and diversification. We maintain that the enigma remains unresolved partly because our current state of knowledge is a result of the fragmentary nature of palaeodata. This lack of palaeodata limits our ability to draw firm conclusions. Nonetheless, based on consistent results, some inferences may be drawn. Our results indicate that a complex multidriver hypothesis may be more suitable than any single-driver theory. We contend that plant carbon economy and diversification may have played an important role during the early stages of gymnosperms replacement by angiosperms in fertile tropical sites. Plant tolerance to climatic stresses, plant nutrition, biotic interactions and diversification may have played a role in later stages of angiosperm expansion within temperate and harsh environments. The angiosperm knot remains partly tied, but to unravel it entirely will only be feasible if new discoveries are made by scientific communities.

467 Taxonomic and palaeoecological implications of mountain floras from the Okhotsk-Chukotka volcanic belts in connection with problem of evolution and expansion of angiosperms

467 Taxonomic and palaeoecological implications of mountain floras from the Okhotsk-Chukotka volcanic belts in connection with problem of evolution and expansion of angiosperms

Early Cretaceous Gleicheniaceae and Matoniaceae (Gleicheniales) from Alexander Island, Antarctica

The first Cretaceous representatives of Matoniaceae and Gleicheniaceae in the Antarctic are described from the Albian flora of Alexander Island. Two new species are recognized, one is assigned to Matonia, and another to the emended gleicheniaceous form-genus Gleicheniaceaephyllum. The taxonomy of fossil genera assigned to Gleicheniaceae is problematic, and as an alternative to the invalid genus Gleichenites, we propose that Microphyllopteris be utilised as a form-genus for species that have either an unknown branching pattern or pseudo-dichotomous branching without resting buds; Microphyllopteris is reserved for ferns that cannot be ascribed conclusively to Gleicheniaceae. Alternatively, Gleicheniaceaephyllum should be used for gleicheniaceous material that has a resting bud amongst two to four primary branches. Gleicheniaceaephyllum acutum sp. nov. is preserved as sterile and fertile fronds, pinnae and pinnules. The other new species, Matonia jeffersonii sp. nov., occurs as sterile and fertile pinnae and pinnules, and is aligned to Matoniaceae based on the presence of circular-oval sori with peltate indusia. The two taxa are inferred to have either a pedate or scrambling habit. These morphologies are relatively common at Alexander Island, compared to other high latitude fossil floras, and are interpreted as an adaptation to the high disturbance, polar conditions and unusual photoperiod experienced at Alexander Island during the Early Cretaceous. The theory that angiosperms caused the demise of ferns is discussed in relation to the Matoniaceae and Gleicheniaceae, and there does not appear to be association between angiosperm expansion and the decline of these two fern families.

Insect Mouthparts: Ascertaining the Paleobiology of Insect Feeding Strategies

One of the most intensively examined and abundantly documented structures in the animal world is insect mouthparts. Major structural types of extant insect mouthparts are extensive, consisting of diverse variations in element structure within each of the five mouthpart regions—labrum, hypopharynx, mandibles, maxillae, and labium. Numerous instances of multielement fusion both within and among mouthpart regions result in feeding organs capable of ingesting in diverse ways foods that are solid, particulate, and liquid in form. Mouthpart types have a retrievable and interpretable fossil history in well-preserved insect deposits. In addition, the trace-fossil record of insect-mediated plant damage, gut contents, coprolites, and insect-relevant floral features provides complementary data documenting the evolution of feeding strategies during the past 400 million years.From a cluster analysis of insect mouthparts, I recognize 34 fundamental mouthpart classes among extant insects and their geochronological evolution by a five-phase pattern. This pattern is characterized, early in the Devonian, by coarse partitioning of food by mandibulate and piercing-and-sucking mouthpart classes, followed by a rapid rise in herbivore mouthpart types for fluid- and solid-feeding during the Late Carboniferous and Early Permian. Mouthpart innovation during the Late Triassic to Early Jurassic added mouthpart classes for fluid and aquatic particle-feeding. This ecomorphological expansion of mouthpart design was associated with the radiation of holometabolous insects, especially Diptera. The final phase of mouthpart class expansion occurred during the Late Jurassic and Early Cretaceous, with addition of surface-fluid-feeding mouthpart classes that subsequently became important during the ecological expansion of angiosperms. Conclusions about the evolution of mouthpart design are based on the mapping of phenetic mouthpart classes onto (ideally) cladistic phylogenies of lineages bearing those same mouthpart classes. The plotting of phenetic and associated ecological attributes onto baseline phylogenies is one of the most important uses of cladistic data.

Insect diversity in the fossil record.

Insects possess a surprisingly extensive fossil record. Compilation of the geochronologic ranges of insect families demonstrates that their diversity exceeds that of preserved vertebrate tetrapods through 91 percent of their evolutionary history. The great diversity of insects was achieved not by high origination rates but rather by low extinction rates comparable to the low rates of slowly evolving marine invertebrate groups. The great radiation of modern insects began 245 million years ago and was not accelerated by the expansion of angiosperms during the Cretaceous period. The basic trophic machinery of insects was in place nearly 100 million years before angiosperms appeared in the fossil record.

Zoogeography of the Coleopterous Family Chauliognathidae

ZOOGEOGRAPHIC philosophy to date has been based primarily on the distribution of vertebrate animals though it would seem that a study of invertebrate forms could also make an important contribution. Recognition of zoogeographic principles can be useful to the better understanding of other disciplines such as systematics and phylogeny. It has opened a fresh approach to my general study of Cantharoidea and has contributed to the delineation of a new family, Chauliognathidae (Miskimen, 1961), as well as suggesting other changes in cantharoid classification which will be treated in future papers. The fundamentals of invertebrate distribution are substantially the same as those affecting vertebrates, especially the poikilotherms (Darlington, 1948). The salient difference is timing. Almost all present-day insect families and many genera were well established at the close of the Cretaceous period while mammals and birds, on the distribution of which many prevailing zoogeographic conclusions are based, were only in very early stages of their evolution. Relatively few major evolutionary changes have occurred among insects since that time; the principal innovations are adaptations, usually at the generic level, correlated with new food sources made available by the Upper Cretaceous angiosperm expansion (Axelrod, 1952; Dorf, 1955). Geology and climatology of the past are particularly useful in the understanding of contemporary insect zoogeography because of a paucity of fossil evidence. Such indirect evidence, as well as inferences drawn from contemporary observations, assumes great importance when resolving puzzling geographic discontinuities and distributions. The beetle family, Chauliognathidae, is an especially good example. Chauliognathids are divided into two tribes, Chauliognathini and Ichthyurini, which are found today in rather restricted localities. The group is admirably suited to zoogeographic study because there are comparatively few species; these agree in generally limited spatial ranges, rather restricted habitats, and relative lack of association with man. Careful survey of their distribution adds materially to the appreciation of relationships between the various taxa, including species. In this paper, geology and climate will be discussed as a unit, followed by correlation with probable origins and dispersions to give a continuous picture of events leading to the contemporary distribution of Chauliognathidae.