Grimaldi and Engel 2005 Current Views on Insect Diversity
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Grimaldi and Engel 2005 Current Views on Insect Diversity Mina Krenz Dan Chou Phylogenomics resolves the timing and paern of insect evolution Misof B. et al 2014 Why Study Insect Phylogeny? • Insects represent one of the earliest animals to make their way to terrestrial and aquatic environments • Due to having a multitude of species, the phylogeny of insects are still well debated to this day • Must reconstruct timelines of insect diversification in order to understand the changes in their physiology and morphology Summary of the methods • 1478 nuclear genes from 144 extant taxa • More than 2.5 gigabases from each of the 103 species they studied • Estimating divergence events via 37 fossil records • Maximum Likelihood of mutations in DNA and amino acids (rather than maximum parsimony) Boot-Strapping • Acts as a resampling method in statistics • Selects a number of base pairs to see how sensitive results are to exclusion of some data. • Resampling data to see how robust and strong it is against randomization. • See conflict in data by providing: confidence intervals, variance, errors, etc. Paleoptera Relationship between Odonates and Ephemeroptera • The data indicates that Odonata and Ephemeroptera are nested within the Paleoptera • However, this analysis is supported by a low boot- strapping • What might this mean? o Odonates and Ephemeroptera possibly paraphyletic? • Why is it difficult to determine the relationships in Paleoptera? Relationship between Odonates and Ephemeroptera • Paleoptera only have two extant lineages, even though they derive from all the way back in the Carboniferous • Most likely Odonata and Ephemeroptera divided shortly after Paleoptera and Neoptera diverged, so huge variance between two Orders • Long time span for Ephemeroptera and Odonata to diverge through gene mutations Neoptera Polyneoptera, Holometabola/Endometabolous and Paraneoptera*** Holometabolous • Strong statistical support for the well-nested group of Holometabolous • High diversity of Hymenoptera, Diptera, and Lepidoptera in early Cretaceous Polyneoptera • Previous study supported the monophyly in groups such as Hexapoda, Insecta, Pterygota, Neoptera, Paraneoptera, and Holometabolous • However, there was weaker support for a monophyletic Polyneoptera (Kjer et al 2006) Polyneoptera • Misof et al strongly support the monophyletic group of the Polyneoptera • Boom in diversity of Blattodea, Mantodea, and Plasmodea in Permean extinction. Polyneoptera Holometabolous /Endopterygota Paraneoptera • The results suggest a diverge of Psocodea from the rest of the Paraneoptera, forming a paraphyletic group • The results show that Psocodea in fact a sister taxa to the Holometabolous • Yet this claim does not have statistical support • Why include this data if it is not backed? Age of Psocodea Taxa? • The study claims that that parasitic lice (Menopan and Pediculus) arose in around 53mya with the emergence of the avian and mammalian taxa. • However, their analysis looks at the crown clade, rather than the stem clade from the remaining • Stem shows an arrival of parasitic lice ~130 mya, at the arrival of feathered theropod dinosaurs Food for Thought • In your opinion, how to these results compare to what has been presented in class/in the book (i.e. what critiques do you have for this study)? • What other data or tests could have been used to make this study more reliable? • Study of genomics is still a fairly new field of science • Science based on certain assumptions and interpretations of the data; continuously changing and growing Gullan and Cranston, 2014 Food for Thought • What evolutionary/environmental factors may have given rise to these diverse groups? Phylogeny of the Ants: Diversificaon in the Age of Angiosperms Moreau C.S., Bell C.D., Vila R., ArchiBald B., Pierce N.E. Ants • Key roles in symBio6c interac6ons • Soil Aeraon • Nutrient cycling • Dominant in terrestrial landscape: – 11,800 species – evolu6onary history poorly resolved Main points • Extant ants arose much earlier than previously proposed: 75-125 mya • Began to diversify late Cretaceous – Early Eocene – 60-100 mya • This me period corresponds with the rise of angiosperms and many herBivorous insects Past phylogenies of Family Formicidae • Past phylogenies proposed using morphological traits and molecular data with less data • This phylogeny constructed from large-scale molecular data – 4.5 kB of sequence data – Six gene regions from 139 of 288 extant genera • Represents 19 of 20 suBfamilies Stas6cal Analysis • Maximum likelihood Bootstrap • Bayesian posterior proBabili6es • Maximum parsimony Bootstrap Major lineages in Formicidae • Three main clades: – Leptanilloid (sister taxon to all other ants) • One suBfamily: Leptanillinae – Poneroid • 5 suBfamilies • AmBlyoponinae lacked support – Formicoid • Contains remaining 13 suBfamilies Monophyly • 14 recovered as Monophyle6c with strong support – Leptanillinae (100%) • Early morphological phylogenies do not show this at Basal posi6on • Basal posi6on shows early tergosternal fusion of 3rd & 4th abdominal segments, lost secondarily • These characters are labile/homoplasious • 19 recovered as monophyle6c • Cerapachyinae paraphyly Bolton = Proposed “poneromorph” clade -AmBlyoponinae -Ectatomminae -Heteroponerinae -Paraponerinae -Ponerinae -Procer6inae These results exclude Ectatomminae and Heteroponerinae, But add Agroecomyrmecinae -Represented By Tatuidris Tatusia Bolton = Proposed “poneromorph” clade using morphological analyses -AmBlyoponinae -Ectatomminae -Heteroponerinae -Paraponerinae -Ponerinae -Procer6inae These results exclude Ectatomminae and Heteroponerinae, But add Agroecomyrmecinae Historical placement of Heteroponerinae and Ectatomminae • Heteroponerinae in formicoid clade is unexpected – Historically in poneromorph clade – Same goes for Ectatomminae (closely related to Heteroponerinae) Fossil record • Oldest reliable fossils containing Formicidae are ~100 million yrs. old from early cretaceous in French & Burnese amBers • Implies earlier history than expected of Formicidae • Results show an even earlier history… 140-168 Million years old! Much older than previous es6mate Based on fossil record Previous studies showing early history of Formicidae • Previous studies By Brady and Ward used molecular data to arrive at an es6mate of 130-140 Million years old – But…. Although these are similar dates the Moreau et al. study used: • Wider sampling • Addi6onal fossils à Leads to an even older es6mate! (140-168 mil years old) Results • Diversificaon of major Formicidae lineages ocurred: – Beginning of Early Paleoceneto Late cretaceous (60-100 Mya) • Ancestors of major suBfamilies present 75-125 mya • If they were present much earlier, why did they take so long to diversify? • Previous fossil record indicates later evolu6on Correspondence with Angiosperm radiaon • Rise in Angiosperm dominated forests was essen6al to the diversificaon of ants – Why would this happen? Discussion Given the importance of plants in determining the 6ming of evolved traits in insects, as well as human’s adverse impact on nature (eg: deforestaon), is it possiBle that insect evolu6on is Being dampened? Would insects Be Bemer off without humans or are all organisms interconnected and important for others to thrive, despite some downfalls? Lineages through 6me plot • LTT plot: Accumulaon of ant lineages around ~100 (following angiosperm radiaon) – Also seen in Coleopteran & Hemipteran diversificaon Is there something wrong with the way this Histogram was constructed? Are LTT graphs a good method for researchers to infer phylogene6c relaonships? Why or why not? Why the correla,on between Angiosperm radiaon and diversificaon of Formicidae? • Forests are more diverse – Wider array of habitats • Expansion of herBivorous insects – Provided direct food source – Indirect food source: honeydew – Shio in diet à evolu6on of social Behaviors Significance? • Evolu6onary inves6gaon of life history, ecology, Biogeography in order to: – OBserve paerns of diversificaon and distriBu6on of this dominant group of insects • This highlights need for conservaon of ant habitats to foster Biodiversity to further research poorly understood evolu6onary history The Fossil record and Macroevolu6onary history of Beetles Smith D.M. and Marcot J.D. Main points • Compiled a database of gloBal Beetle fossil data in order to study evolu6onary history • Polyphaga responsible for most taxononmic richness of Beetles – Also increase in diversificaon rate in Cretaceous like Formicidae, But not due to Angiosperm radiaon • OBserved mechanisms that inhiBited Beetle ex6nc6on rather than mechanisms promo6ng speciaon Polyphagan vs. Non-Polyphagan diversificaon • Degree of dietary variaon and specializaon within suBgroups in Polyphaga – Algae, fluid feeders, carnivores, xylophages • Non-polyphagans first to appear in fossil record – Reach peak of family richness in Triassic – Jurassic: low originaon rates and higher ex6nc6on rates than Polyphaga • Polyphagans surpass richness of non- polyphagans in Jurassic – Established early and longlived (family ex6nc6on rate of zero) Increase in diversificaon rate of Polyphagans Polyphagans diversificaon rate surpasses that of non-polyphagans (who have a higher ex6nc6on rate in this 6me period) Non-polyphagans reach family richness peak Origin of non-polyphagans Should this middle-cretacean increase in the diversificaon rate of Polyphagans be ariButed to the rise in Angiosperms during the same 6me period? Like that of the Ant paper? AmBer deposits • Instead of ariBu6ng this