Literature Review: Arthropod Evolution

by Carl Strang

If you’re a bug nerd you’ll enjoy the following notes on research from 2013. Especially significant were studies of butterflies and moths, and an eye-opening paper on periodical cicadas. This concludes my literature review until next winter.

Butterflies and moths had their origin in the Triassic Period according to recent studies, though the first ones were more like caddis flies than like this red-spotted purple.

Butterflies and moths had their origin in the Triassic Period according to recent studies, though the first ones were more like caddis flies than like this red-spotted purple.

Zhang, W, et al. 2013. New fossil Lepidoptera (Insecta: Amphiesmenoptera) from the Middle Jurassic Jiulongshan Formation of northeastern China. PLoS ONE 8(11): e79500. doi:10.1371/journal.pone.0079500  They found 15 species of early moths representing at least 3 families in Chinese deposits, and details of wing venation led to the conclusion that the Lepidoptera (moths and butterflies) diverged from the Trichoptera (caddis flies) by the early Jurassic Period.

Wahlberg, N, CW Wheat, C Peña 2013. Timing and patterns in the taxonomic diversification of Lepidoptera (butterflies and moths). PLoS ONE 8(11): e80875. doi:10.1371/journal.pone.0080875  They estimated timings of major episodes of speciation in the major groups of butterflies and moths. Their results point to a Triassic origin of Lepidoptera, around 215 million years ago. The timing of diversification episodes at least in some cases corresponds to times when plants were diversifying, and also after the end-Cretaceous mass extinction. Coevolution of lepidoptera with their larval food plants appears to be an important theme. They give origin ages for major Lepidoptera groups (in millions of years ago): Gracillarioidea 120, Yponomeutoidea 117, Glechioidea 106 (these first three are small moths, many of them leaf miners), Papilionoidea 104 (butterflies), Pyraloidea (including many local pyralid moths) 93, Bombycoidea (including sphinx moths) 84, Geometroidea (including inchworm moths) 83, Noctuoidea (the enormous owlet moth group) 82, Tortricoidea (including leaf-folding caterpillars) 68. All these groups are represented by local species.

The Chicago region’s 17-year periodical cicadas: Magicicada septendecim, left, and M. cassini.

The Chicago region’s 17-year periodical cicadas: Magicicada septendecim, left, and M. cassini.

Sota, Teiji, Satoshi Yamamoto, John R. Cooley, Kathy B.R. Hill, Chris Simon, and Jin Yoshimura. 2013. Independent divergence of 13- and 17-y life cycles among three periodical cicada lineages. Proc. Nat. Acad. Sci. 110:6919-6924. They sequenced a number of genes from nuclear and mitochondrial DNA from all known species and broods, and estimated divergence times based on general research that has been done on insect mitochondria. There are three species groups (referred to as Decim, Cassini, and Decula), each of which contains northern 17-year species and southern 13-year species. In any location, the species in the different groups emerge at the same time. The results clearly separated the three groups, and tied together the species within each group (e.g., 13-year Decim are more closely related to 17-year Decim than to 13-year Cassini). Furthermore, each species group is divided into eastern, central and western genetic clusters (this pattern has been documented in other organisms as well; for the most part, Illinois cicadas are in western clusters, Indiana ones in central clusters). Each cluster contains both 13- and 17-year species, “suggesting that life cycle divergence occurred independently in the three regions.” Analyses estimated that the western Cassini divergence of 13-year and 17-year species took place 23,000 years ago, 10,000 years for Decim. Population sizes for both Decim and Cassini groups appear to have been small during the last glacial period, but expanded greatly starting 10,000 years ago. The sequence appears to have been allopatric speciation of the 3 ancestral species, with the species later becoming sympatric and independently splitting into 13- and 17-year cicadas. “Surprisingly, however, the divergence of 13- and 17-y cicadas was asynchronous among the species groups and occurred repeatedly even within a species group.” The implication is “that the three Magicicada groups shared multiple refugia during the last glacial maximum.” The 13-/17-year splits occurred after the last glacial maximum, within the last 23,000 years, “suggesting that the life cycle divergence in Magicicada is closely associated with global climatic fluctuations and shorter growing seasons in the north versus the south.” However, the species groups themselves separated in the Pliocene, and their shared long lives suggest that this did not originate because of glacial climate influences. This shifting between 13- and 17-year life cycles suggests a common genetic basis among the species, and indicates a somewhat plastic nature of this trait. The coordination among species at a given location seems best explained by the selective advantage of low numbers of an invading species into the range of another, surviving best when sheltered by the established species’ numbers.

Zhao, Z, et al. 2013. The mitochondrial genome of Elodia flavipalpis Aldrich (Diptera: Tachinidae) and the evolutionary timescale of tachinid flies. PLoS ONE 8(4): e61814. doi:10.1371/journal.pone.0061814  Their genomic study traced the evolutionary relationships of the parasitic fly family Tachinidae, and molecular clock analysis calibrated to the fossil record points to the middle Eocene as the time of the family’s origin.

Brewer, MS, and JE Bond. 2013. Ordinal-level phylogenomics of the arthropod class Diplopoda (millipedes) based on an analysis of 221 nuclear protein-coding loci generated using next-generation sequence analyses. PLoS ONE 8(11): e79935. doi:10.1371/journal.pone.0079935  They place the ancestral millipedes at 510mya (million years ago), with major groupings established by 200mya.

Lucky, A, MD Trautwein, BS Guénard, MD Weiser, RR Dunn. 2013. Tracing the rise of ants – out of the ground. PLoS ONE 8(12): e84012. doi:10.1371/journal.pone.0084012     A phylogenetic analysis points to soil rather than leaf litter as the nesting habitat for the earliest ant species.

Literature Review: Mammal Evolution

by Carl Strang

Today’s collection of notes from the 2013 scientific literature focuses on mammals and their evolution. As the notes reveal, some of these topics are controversial among researchers.

This migrating bat chose a famous resting place during its journey: the Aldo Leopold shack in Wisconsin. Bats are the subject of two of the following studies.

This migrating bat chose a famous resting place during its journey: the Aldo Leopold shack in Wisconsin. Bats are the subject of two of the following studies.

Chang-Fu Zhou, Shaoyuan Wu, Thomas Martin, Zhe-Xi Luo. 2013. A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations. Nature 500 (7461): 163 DOI: 10.1038/nature12429  They described a newly discovered Jurassic proto-mammal, Megaconus mammaliaformis, and found evidence that traits such as hair and fur originated well before the rise of the first true mammals. The squirrel-sized Megaconus had a heel spur, similar to poisonous spurs found on modern egg-laying mammals, such as male platypuses. It had mammalian dental features, and legs and feet that point to a gait similar to that of modern armadillos. At the same time it had a reptilian middle ear, ankle bones and vertebral column.

O’Leary, Maureen, et al. 2013. The placental mammal ancestor and the post-K-Pg radiation of placentals. Science 339:662-667. Using fossil materials and an extensive character analysis, they conclude that the ancestral placental mammal from which all major surviving groups evolved lived just after the beginning of the Paleocene. This conflicts with molecular clock data that place the appearance of many groups including bats, rodents, and even-toed ungulates back in the Cretaceous. They combine the characters of the early fossils to produce a hypothetical common ancestor, an insectivorous animal resembling a shrew with a long tail.

Zhang, Guojie, et al. 2013. Comparative analysis of bat genomes provides insight into the evolution of flight and immunity. Science 339: 456-460. They did whole-genome comparisons of nuclear DNA of a Myotis and a flying fox. Significant sequences were found which may relate to the development of flight ability, and the immune systems also are different from those of other mammals. When compared to the genomes of other mammals, bats fall out most closely related to perissodactyls, then carnivores, with those groups splitting apart at an estimated time in the Cretaceous.

Ni, Xijun, et al. 2013. The oldest known primate skeleton and early haplorhine evolution. Nature 498 (7452): 60 DOI: 10.1038/nature12200  They describe a 55mya (early Eocene) Chinese fossil that is in the tarsier line but has features showing it to be close to the branch point leading to the tarsiers in one direction, anthropoids (primates including monkeys, apes and humans) on the other. It is tiny, the animal around 1 ounce in weight. Asia appears to be the likely center of early primate evolution.

Cahill JA, Green RE, Fulton TL, Stiller M, Jay F, et al. 2013. Genomic evidence for island population conversion resolves conflicting theories of polar bear evolution. PLoS Genet, 9(3): e1003345; DOI: 10.1371/journal.pgen.1003345  This most recent examination of polar bear and brown bear genetics concluded that, on the whole, polar bears have been separate from brown bears for about half the time that brown bears have been separate from black bears. The connections previously noted between the two species in southeast Alaska, and possibly in Ireland, appear to be the result of small polar bear populations being isolated during ice ages, and being swamped then by an influx of male brown bears. The polar bear is a more ancient species than that.

Zigouris J, Schaefer JA, Fortin C, Kyle CJ. 2013. Phylogeography and post-glacial recolonization in wolverines (Gulo gulo) from across their circumpolar distribution. PLoS ONE 8(12): e83837. doi:10.1371/journal.pone.0083837  Their analysis of mitochondrial and nuclear genes points to a single population of wolverines surviving the glacial maximum in a refugium somewhere in the Old World, then expanding into North America across the Bering Sea land bridge as the glaciers retreated. Subsequently, several North American populations differentiated. The fossil record likewise has them only in Eurasia prior to the late Pleistocene.

Andrew M. Minnis, Daniel L. Lindner. 2013. Phylogenetic evaluation of Geomyces and allies reveals no close relatives of Pseudogymnoascus destructans, comb. nov., in bat hibernacula of eastern North America. Fungal Biology, DOI: 10.1016/j.funbio.2013.07.001  As described in a ScienceDaily article. The closest relatives of the fungus causing white nose syndrome are species that live in European caves. This supports the idea that the fungus is an invasive species here, but one with which European bats coevolved and so have some immunity.

Literature Review: Mesozoic Miscellany

by Carl Strang

The Mesozoic Era continues to fascinate researchers as well as 8-year-olds. This is the time when the dinosaurs rose to their dominance of the terrestrial fauna, which they maintained through many millions of years, though evolution continued to shape and diversify them. The mammals and birds appeared, the latter as an offshoot of the dinosaurs, the former out of a different group. The flowering plants likewise made their entrance onto the stage of life. Today I share my notes from a grab-bag of studies published last year.

This is the sort of image that excites wonder about the Mesozoic. Traveling exhibit at the Brookfield Zoo.

Timothy B. Rowe, Thomas E. Macrini and Zhe-Xi Luo. 2011. Fossil evidence on origin of the mammalian brain. Science 332: 955-957. Brain reconstructions from scans of two early Jurassic pre-mammal skulls from China, Morganucodon and Hadrocodium, added to other information, suggest that the mammal brain size increase that is so important in distinguishing that group happened in 2 stages involving portions of the brain associated with olfaction, with later elaboration to accommodate increased tactile receptors connected to the fur, and then increased motor areas to coordinate responses to such increased sensory input.

P. A. Selden, C. Shih, D. Ren. A golden orb-weaver spider (Araneae: Nephilidae: Nephila) from the Middle Jurassic of China. Biology Letters, 2011; DOI: 10.1098/rsbl.2011.0228     (Described in ScienceDaily article). The most remarkable point here (apart from the spider’s size: 5 inches across its legs) is the fact that this Jurassic fossil is from a genus of orb-weavers still alive today. They are mainly tropical, with one species, Nephila clavipes, reaching the Gulf Coast of the United States.

Ge Sun, David L. Dilcher, Hongshan Wang, Zhiduan Chen. A eudicot from the Early Cretaceous of China. Nature, 2011; 471 (7340): 625 DOI: 10.1038/nature09811     (As described in ScienceDaily article). This Liaoning fossil is of the top of a plant with leaves and flowers similar to buttercups. The authors place it in the Ranunculaceae (buttercup family). It is the earliest fossil, other than pollen, of a true dicot, at 125 million years ago. It is sufficiently advanced to point to an earlier origin of the family, and of eudicots generally, which might have appeared as early as the Late Jurassic.

Snively E, Theodor JM (2011) Common Functional Correlates of Head-Strike Behavior in the Pachycephalosaur Stegoceras validum (Ornithischia, Dinosauria) and Combative Artiodactyls. PLoS ONE 6(6): e21422. doi:10.1371/journal.pone.0021422     Their comparative study resurrects the possibility that head-butting was a feature of pachycephalosaur (bone-headed dinosaur) social behavior.

Lindsay E. Zanno and Peter J. Makovicky. Herbivorous ecomorphology and specialization patterns in theropod dinosaur evolution. Proceedings of the National Academy of Sciences, 2010; DOI: 10.1073/pnas.1011924108     Theropods include all the predatory dinosaurs. These researchers looked at the coelurosaurian theropods, a diverse group whose diets were known to be varied but uncertain for many species. Using clues such as fossilized gut contents, coprolites (fossilized excrement), tooth morphology, and presence or absence of grinding stones in the gut area, they found 21 skeletal features that are closely associated with an herbivorous diet. Having established that base, they reviewed the coelurosaurs and found that extreme carnivores (e.g., Velociraptor) are a minority in the group. Extreme carnivory may in fact have been derived from an earlier omnivorous or even herbivorous ancestry. One feature, the toothless beak (endpoint of an evolutionary process of progressive tooth loss, and evolving in more than one group independently), is associated with herbivory and is of special interest because the coelurosaurs include the birds. The toothless beak is associated with the gastric mill. In some cases (Therizinosaurians) there is no sign of a gastric mill, and a battery of grinding teeth is retained. Three groups of coelurosaurs, the tyrannosauroids, the Compsognathidae and Dromaeosauridae, are carnivorous, but most others in the group are found to be herbivorous. Of 90 species, 24 fall out as carnivorous and 44 as herbivorous (the remaining ones omnivorous or undetermined).

Fowler DW, Freedman EA, Scannella JB, Kambic RE (2011) The Predatory Ecology of Deinonychus and the Origin of Flapping in Birds. PLoS ONE 6(12): e28964. doi:10.1371/journal.pone.0028964     They re-examined the functional morphology of the enlarged hind foot claw of Deinonychus and other “raptor” dinosaurs, and propose an alternative function: that instead of using it as a slashing tool in attacks on prey larger than themselves, the predators used it as a clasping tool to help immobilize prey of similar or smaller body size for killing and dismemberment with the teeth. The authors further suggest that the wing-like forearms could be flapped to increase the predator’s stability during this process, and that this might be relevant to the evolution of flight in birds.

Prehistoric Life 12

by Carl Strang

This year’s winter series is a review of the prehistoric life and geologic history of northeast Illinois. Each chapter will summarize current understanding, gleaned from the literature, of what was going on with life on Earth in a particular span of time, what we know about the local landscape, and what we can say about local life. I include some references, particularly to papers published in the journal Science which commonly is available at public libraries. Contact me if you need sources for other items. The Earth is so old that every imaginable environment was here at some point, from ocean depths to mountaintops, from equatorial tropics to tundra, and from wetlands to desert.

Yes, it’s a dinosaur, and yes, we’ve reached the Jurassic Period, but this one won’t appear until later.

Jurassic Period (208-144 million years ago)

The Jurassic Period was named in 1795 for layers of rocks in the Jura Mountains between France and Switzerland.

Life on Earth. Reef-forming corals went extinct at the end of the Triassic Period. New ones evolved 8-10 million years later, but by the late Jurassic, bivalves called rudists had begun to displace corals as the primary reef formers, and retained that status through the rest of the Mesozoic Era (Science 312:857).

Gymnosperms including conifers and cycads were the dominant woody plants. No flowering plants appeared, yet. However, there is evidence that insect-mediated pollination by a suite of scorpionflies was taking place in non-angiosperm seed plants (Science 326: 840).

Ants were a major new insect group. A side-necked turtle is known from the late Jurassic of Europe. Phytosaurs went extinct in the early Jurassic, but the similar crocodilians diversified.

Sauropods were the principal herbivores of the Jurassic Period.

Dinosaurs became the dominant terrestrial animals, and ranged even into deserts. Prosauropods left the most abundant vertebrate fossils of the early Jurassic. Their presumed offshoot group the sauropods, the familiar enormous four-legged dinosaurs with long necks and long tails, appeared in the early Jurassic, and by the late Jurassic reached their peak diversity, having become the dominant herbivores. They had several advantages that allowed them to have such large body sizes and numerical abundance. These included a large gut capacity enabling a long residence time for contents, which made chewing or grinding of plant food unnecessary and allowed digestion of large masses of food. Without a heavy chewing apparatus the head could be relatively small and the neck long, allowing a higher reach (in fact, sauropod skulls were so fragile and light that they seldom are found; complete skulls are known for only 8 of 120 sauropod species). The efficient, avian style air exchange system complete with bone-lightening air sacs was an important large size enabler. Other advantages were a high metabolic rate, and the production of large numbers of small young rather than a few large ones (Science 322:200-201). Sauropod genera from one famous Jurassic fossil source, the Morrison formation, include Camarasaurus, Apatosaurus (“Brontosaurus;” O.C. Marsh, an overzealous paleontologist, found and named Apatosaurus, then 2 years later found a larger sauropod he named Brontosaurus. The latter fossil turned out to be yet another Apatosaurus, which remains the correct name because of priority), Diplodocus (the size of five elephants), Ultrasaurus (approached blue whale size), Supersaurus, and Brachiosaurus. The last also is known from Portugal, and north and east Africa.

Predatory carnosaurs were few in mid-Jurassic times, but had become a large and varied group by the late Jurassic. The Morrison formation includes Ceratosaurus and the larger Allosaurus. Tooth marks from the latter are known on Apatosaurus bones, but scavenging is a possibility. Ornitholestes, also from the Morrison formation, had long grasping front legs. Compsognathus is known from the forested eastern seashore of our then continent in the late Jurassic of Germany.  Early ornithopods (2-legged ornithischian herbivores) appeared in Asia in the early Jurassic, and there was a major dispersal into North America in the mid-Jurassic. The stegosaurs appeared in the middle Jurassic in Europe and Asia, also spread to North America, and peaked during the late Jurassic Period with only a few surviving into the Cretaceous. Ankylosaurs appeared in late Jurassic Europe, China, and the western U.S. and Canada.

Stegosaurs also were characteristic of the late Jurassic. This was another of the sculptures at the Brookfield Zoo’s temporary display in 2009.

An ecological study of late Jurassic fossil assemblages found that arid regions were dominated by large-bodied, high-reaching herbivores and their likewise large predators. Wet, densely vegetated forest regions had more abundant dinosaurs of small to medium size. Intermediate environments had a diversity of large and small forms.

Of course there were the big predators, too. Here a theropod fossil is mounted so as to be attacking a stegosaur in a temporary 2005 exhibit at the Field Museum.

Ichthyosaurs flourished early, but declined during the Jurassic. Plesiosaurs were represented throughout, and beyond, the Jurassic. Pterosaurs (winged reptiles) were diverse in Europe through the Jurassic, with long-tailed ones dominant for most of the period. They declined in the late Jurassic, and were replaced by short-tailed ones that appeared then. These continued into the Cretaceous. Only a few Jurassic pterosaur fossils are known from North America, but conditions for preserving them here were poor. Pterosaurs now are known to have been fur-covered (though some suspect this “fur” was closer to feathers than to hairs), and probably were warm-blooded.

Ichthyosaurs swam the Jurassic seas. This sculpture is at a significant fossil site for the group in Nevada.

Birds (Archaeopteryx) appeared in the late Jurassic, and the concensus now is that they were an offshoot of the two-legged, predatory theropod dinosaurs. In fact, Archaeopteryx and some feathered members of the dromaeosaur family of theropods were very similar to one another (Paleobiology 32:417). The range of fossils indicates that the dromaeosaurs, their close relatives of the troodontid family, and birds all started out small (Science 317:1378). One small troodontid may have had flight capability with 4 wings (the forelimbs, plus winglike clusters of quill feathers on the legs; Nature 461: 640). The larger, later members of those two dinosaur groups such as Velociraptor were derived from small ancestors. But Velociraptor retained quill feathers on its forelimbs (Science 317:1721). Early in 2009 an ornithischian dinosaur was found with fuzz that well may have been proto-feathers (Nature 458:333). The sauropods and theropods both had air sac systems which connect them to the birds as well. Mammals diversified, but still had produced none of the groups we see today.

Local landscape. There are no Jurassic deposits in Illinois. Surrounding geology informs us that this area remained low land. The closest Jurassic bedrock is in small areas of the Oklahoma panhandle, and the Black Hills of western South Dakota. Pangaea began to break up in the early Jurassic, with southern Gondwana and northern Laurasia separating by the middle Jurassic. A tilting of the continent permitted a sea to flood the center of North America, reaching from the current Gulf of Mexico to the Arctic, and our nearest west coast was somewhere in central to western Iowa. By the late Jurassic, eastern North America was separate from western North America and all other continents except western Europe. Our area continued to drift north, though the worldwide climate remained warm.

Local life. As in the Triassic, we don’t know what kind of vegetation was here, though again this area was dry land. Elsewhere at our then latitude, conifers, ginkgos and ferns were dominant plants. We were a little farther north of the equator thanks to continental drift, but the climate was still warm. Astrodon, a brachiosaurid, has been found in Maryland and Texas, and so is a good candidate for our local dinosaur fauna. The large predator Allosaurus has been found both in the western U.S. and in Portugal, and so may well have been here as well. The prosauropods Anchisaurus and Ammosaurus have been found in the middle Jurassic of Massachusetts, along with the theropod Podokesaurus. Camptosaurus is an early ornithopod that has turned up on all sides of this part of North America. Our area then was connected with Europe, so pterosaurs and early birds also are a possibility. Late Jurassic dinosaur assemblages typically include “three or more sauropods, two or more theropods, one or two stegosaurs, and one or two ornithopods.”

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