Handbook of Avian Anatomy: Nomina Anatomica Avium

The total-evidence approach to divergence time dating uses molecular and morphological data from extant and fossil species to infer phylogenetic relationships, species divergence times, and macroevolutionary parameters in a single coherent framework. Current model-based implementations of this approach lack an appropriate model for the tree describing the diversification and fossilization process and can produce estimates that lead to erroneous conclusions. We address this shortcoming by providing a total-evidence method implemented in a Bayesian framework. This approach uses a mechanistic tree prior to describe the underlying diversification process that generated the tree of extant and fossil taxa. Previous attempts to apply the total-evidence approach have used tree priors that do not account for the possibility that fossil samples may be direct ancestors of other samples, that is, ancestors of fossil or extant species or of clades. The fossilized birth–death (FBD) process explicitly models the diversification, fossilization, and sampling processes and naturally allows for sampled ancestors. This model was recently applied to estimate divergence times based on molecular data and fossil occurrence dates. We incorporate the FBD model and a model of morphological trait evolution into a Bayesian total-evidence approach to dating species phylogenies. We apply this method to extant and fossil penguins and show that the modern penguins radiated much more recently than has been previously estimated, with the basal divergence in the crown clade occurring at ∼12.7 ∼12.7 Ma and most splits leading to extant species occurring in the last 2 myr. Our results demonstrate that including stem-fossil diversity can greatly improve the estimates of the divergence times of crown taxa. The method is available in BEAST2 (version 2.4) software www.beast2.org with packages SA (version at least 1.1.4) and morph-models (version at least 1.0.4) installed.

Bayesian Total-Evidence Dating Reveals the Recent Crown Radiation of Penguins

Testing models of macroevolution, and especially the sufficiency of microevolutionary processes, requires good collaboration between molecular biologists and paleontologists. We report such a test for events around the Late Cretaceous by describing the earliest penguin fossils, analyzing complete mitochondrial genomes from an albatross, a petrel, and a loon, and describe the gradual decline of pterosaurs at the same time modern birds radiate. The penguin fossils comprise four naturally associated skeletons from the New Zealand Waipara Greensand, a Paleocene (early Tertiary) formation just above a well-known Cretaceous/Tertiary boundary site. The fossils, in a new genus (Waimanu), provide a lower estimate of 61-62 Ma for the divergence between penguins and other birds and thus establish a reliable calibration point for avian evolution. Combining fossil calibration points, DNA sequences, maximum likelihood, and Bayesian analysis, the penguin calibrations imply a radiation of modern (crown group) birds in the Late Cretaceous. This includes a conservative estimate that modern sea and shorebird lineages diverged at least by the Late Cretaceous about 74 +/- 3 Ma (Campanian). It is clear that modern birds from at least the latest Cretaceous lived at the same time as archaic birds including Hesperornis, Ichthyornis, and the diverse Enantiornithiformes. Pterosaurs, which also coexisted with early crown birds, show notable changes through the Late Cretaceous. There was a decrease in taxonomic diversity, and small- to medium-sized species disappeared well before the end of the Cretaceous. A simple reading of the fossil record might suggest competitive interactions with birds, but much more needs to be understood about pterosaur life histories. Additional fossils and molecular data are still required to help understand the role of biotic interactions in the evolution of Late Cretaceous birds and thus to test that the mechanisms of microevolution are sufficient to explain macroevolution.

Early Penguin Fossils, Plus Mitochondrial Genomes, Calibrate Avian Evolution

The Miocene epoch (23.03–5.33 Ma) was a time interval of global warmth, relative to today. Continental configurations and mountain topography transitioned towards modern conditions, and many flora and fauna evolved into the same taxa that exist today. Miocene climate was dynamic: long periods of early and late glaciation bracketed a ∼2 Myr greenhouse interval – the Miocene Climatic Optimum (MCO). Floras, faunas, ice sheets, precipitation, pCO2, and ocean and atmospheric circulation mostly (but not ubiquitously) covaried with these large changes in climate. With higher temperatures and moderately higher pCO2 (∼400–600 ppm), the MCO has been suggested as a particularly appropriate analogue for future climate scenarios, and for assessing the predictive accuracy of numerical climate models – the same models that are used to simulate future climate. Yet, Miocene conditions have proved difficult to reconcile with models. This implies either missing positive feedbacks in the models, a lack of knowledge of past climate forcings, or the need for re‐interpretation of proxies, which might mitigate the model‐data discrepancy. Our understanding of Miocene climatic, biogeochemical, and oceanic changes on broad spatial and temporal scales is still developing. New records documenting the physical, chemical, and biotic aspects of the Earth system are emerging, and together provide a more comprehensive understanding of this important time interval. Here we review the state‐of‐the‐art in Miocene climate, ocean circulation, biogeochemical cycling, ice sheet dynamics, and biotic adaptation research as inferred through proxy observations and modelling studies.

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The Miocene: The Future of the Past

Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy: I. – Methods and characters [X1228] (matrix)

Eocene penguin remains from Seymour Island (Antarctica) are so far the old− est−known record of extinct Sphenisciformes. Rich Argentine and Polish collections of pen− guin bones from the La Meseta Formation are taxonomically revised on tarsometatarsal morphology. Two genera and four species are erected: Mesetaornis polaris gen. et sp. n., Marambiornis exilis gen. et sp. n., Delphinornis arctowskii sp. n. and D. gracilis sp. n. Moreover, the diagnoses of already described species: Anthropornis nordenskjoeldi, A. grandis, Palaeeudyptes klekowskii, P. gunnari, Archaeospheniscus wimani and Delphi− nornis larseni are revised as well. Gradual cooling of climate, changes of environment and trophic relationships, that lasted several millions years, were most probably responsible for the intense speciation and taxonomic diversification of the Middle-Late Eocene La Meseta penguins.

Taxonomic revision of Eocene Antarctic penguins based on tarsometatarsal morphology

Skeletal remains of penguins from the Eocene La Meseta Formation (Seymour Is− land, Antarctica) constitute the only extensive fossil record of Antarctic Sphenisciformes. No articulated skeletons are known, and almost all fossils occur as single isolated elements. Most of the named species are based on tarsometatarsi (for which the taxonomy was revised in 2002). Here, 694 bones (from the Polish collection) other than tarsometatarsi are reviewed, and allocated to species. They confirm previous conclusions and suggest that ten species grouped in six genera are a minimal reliable estimate of the Eocene Antarctic penguin diver− sity. The species are: Anthropornis grandis, A. nordenskjoeldi, Archaeospheniscus wimani, Delphinornis arctowskii, D. gracilis, D. larseni, Marambiornis exilis, Mesetaornis polaris, Palaeeudyptes gunnari and P. klekowskii. Moreover, diagnoses of four genera (Anthropornis, Archaeospheniscus, Delphinornis and Palaeeudyptes) and two species (P. gunnari and P. klekowskii) are supplemented with additional, non−tarsometatarsal features. Four species of the smallest penguins from the La Meseta Formation (D. arctowskii, D. gracilis, M. exilis and M. polaris) seem to be the youngest taxa within the studied assemblage - their remains come exclusively from the uppermost unit of the formation. All ten recognized species may have co−existed in the Antarctic Peninsula region during the Late Eocene epoch. Key wor ds: Antarctica, La Meseta Formation (Eocene), paleontology (penguins), taxon− omy.

Eocene penguins of Seymour Island, Antarctica: Taxonomy

Penguins (Aves: Sphenisciformes) hold much interest for many people, includ− ing (but not limited to) scientists. According to results of molecular studies, penguin his− tory began in the Cretaceous, but the oldest bones assigned to these birds are Paleocene in age. The first fossil representative of Sphenisciformes formally described was Palae− eudyptes antarcticus, and this event took place 150 years ago. Since that time, several dozens of species have been erected, though not all of them have stood a test of time. The 21st century entered new dynamics into the paleontology of penguins, and (importantly) it concerned both the new material, and new theories. This paper summarizes what we currently know about extinct penguins and indirectly suggests the most promising areas for further research. Key wor ds: Southern Hemisphere, Aves, Sphenisciformes, evolution, fossil record.

Penguin past: The current state of knowledge

Penguin bones from the Eocene La Meseta Formation (Seymour Island, Antarctic Peninsula) constitute the only extensive fossil record of Antarctic Sphenisciformes. Here, we synonymize some of the recognized genera (Anthropornis with Orthopteryx, Delphinornis with Ichtyopteryx) and species (Anthropornis nordenskjoeldi with Orthopteryx gigas, Delphinornis gracilis with Ichtyopteryx gracilis). Moreover, we suggest that Antarctic species of Anthropornis and Palaeeudyptes, so-called giant penguins, may in fact comprise only one species each instead of two, based on evidence of well-marked sexual dimorphism. We also present new estimates of body mass based on femora testifying to the impressive scope of interspecific body-size variation in Eocene Antarctic penguins.

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Aspects of diversity in early Antarctic penguins

Penguin bones from the La Meseta Formation (Seymour Island, Antarctic Penin− sula) are the only record of Eocene Antarctic Sphenisciformes. Being an abundant compo− nent of the youngest unit of the formation (Telm7), they are not so common in earlier strata. Here, I present the oldest penguin remains from the La Meseta Formation (Telm1-Telm2), often bearing close resemblance to their counterparts from younger units. Addressing the recent findings in fossil penguin systematics, I suggest there is too weak a basis for erecting new Eocene Antarctic taxa based on non−tarsometatarsal elements of penguin skeletons, and considering Oligocene species part of the studied assemblage. Finally, I conclude if the common ancestor of extant Sphenisciformes lived in the Eocene Antarctic (as suggested re− cently), penguins referred to Delphinornis seem to be prime candidates to that position. Key wor ds: Antarctica, La Meseta Formation (Eocene), paleontology (penguins), taxon− omy, evolution.

Piotr Jadwiszczak

Papers

Taxonomic revision of Eocene Antarctic penguins based on tarsometatarsal morphology

Eocene penguins of Seymour Island, Antarctica: Taxonomy

Penguin past: The current state of knowledge

Aspects of diversity in early Antarctic penguins

Eocene penguins of Seymour Island, Antarctica: The earliest record, taxonomic problems and some evolutionary considerations