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Systematic Relationships among Birds. Hypotheses for the evolution of birds Determines how your bird guide is (or will be) arranged Contentious Morphology What to measure, convergent evolution Fossils Missing links / sample bias Lack of soft parts DNA How to calibrate
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Systematic Relationships among Birds • Hypotheses for the evolution of birds • Determines how your bird guide is (or will be) arranged • Contentious • Morphology • What to measure, convergent evolution • Fossils • Missing links / sample bias • Lack of soft parts • DNA • How to calibrate • Which and how many genes to use • What species to include in analysis (outgroups especially)
Fundamental Divisions • Paleognathae • Primitive palate • Monophyly of many flightless birds • Hoatzin primitive but not basal to Neognathae • Opisthocomiformes • Galloanserae is basal and monphyletic In Rheas, Tinamous, Ostrich, Cassowaries, Emu, Kiwi, and extinct Moas and Elephant Birds the vomar bone extends back to articulate with palatines and pterygoids to separate both from the parashenoid rostrum
Tertiary Big Bang? K-T boundary (end of Cretaceous; 65 my) Volcanism, mountain building, regression of continental seas meteor in Mexico (Chicxulub crater in Yucatan) Dinosaur extinctions Theory in accordance with Fossil record (Feduccia 2003)
(Ericson et al. 2002) (Cracraft 2001)
Ancestor of Passeriformes was Gondwanan; divergence began before KT • Acanthisitta = New Zealand Wrens are most primitive passerine and sister to Suboscines and Oscines (songbirds) • They diverged from other passerines about 82my, ancestors moved across connected Antarctic • Oscine and Suboscine songbirds began to diverge about 77my (Barker et al. 2004)
Red = Australasia Green = Africa and Eurasia Blue = North and South America Grey = ambiguous ancestral / current area (Barker et al. 2004)
So, the Real Explosion was After the KT event (Barker et al. 2004)
Why Did Some Birds Survive Mass Extinction? • Body Size? • Smaller requires less energy, shorter generation times • Southern Distribution? • Early passerine ancestor on New Zealand • Beaks? • Crush hard seeds that survive impact • Ground/water dwelling • Avoid impacts of forest destruction • Field et al. 2018 Current Biology 28:1825-1831
50ka During the last several million years land connections (via Beringia) between New and Old Worlds waxed and waned with glaciation. Beringia was dry and offered land passage.
Corvids Invade Our World Tied to changes in vegetation and sea level Tertiary forests of Australia were being replaced by deserts perhaps forcing corvid ancestors (related to Birds of Paradise and Orioles) to leave Australia and head north in Oligocene and Miocene to Asia, following northward movement of tropical forests 6-8 mya in Miocene New World Jay ancestor from forests of southeast Asia, radiate in South America (first) and North America 15,000 ya – 2 mya in Pleistocene Old World Jay (Gray Jay) Nutcracker Magpie Crow Raven (from article on evolution of cats; Johnson et al. 2006; Science 311:73-77)
Ravens • 4 clades diverging in Africa 1.7-3.8my • Corvus corax ancestor diverges (closest relative is C. albus) shortly thereafter • C. corax invades New World 2my and new and old world ravens begin independent evolution • Old world raven spins off Canary Island Raven 650,000 yr • New world ravens spins off Chihuahuan Raven • C. corax reinvades New World 15,000 years ago
Complexity Revealed By Genetic Analyses • Common Raven • ~15,000 years ago old world ravens again invaded the new world via Beringia • Holarctic and California clade of ravens now found in North America, but they are becoming more similar, not diverging as they had in past. • Giving us new insights into what constitutes a “species”
Scrub-jays Florida Scrub-jay Fig. 1. FROM DELANEY et al. 2008---Distribution of Island and Western scrub-jays, with associated geographic trends in morphological characteristics. Species distributions are adapted from Curry et al. (2002); orange = Aphelocoma insularis, green = Californica group of Aphelocoma californica, light blue = Woodhouseii group of Aphelocoma californica, and dark blue = sumichrasti group of Aphelocoma californica. Currently we recognize 3 species, but there are most likely 5 and maybe 6 isolation has been of paramount importance and novel selective pressures from foods eaten (oaks versus other seeds)
CITATION: McCormack, Peterson, Delaney, Knowles, in prep. Unicolor Jay (Aphelocoma unicolor) A. un. concolor A. un. guerrerensis A. un. oaxacae A. un. griscomi A. un. unicolor
A. ul. arizonae A. ul. couchii A. ul. wollweberi A. ul. gracilis A. ul. potosina A. ul. ultramarina A. ul. colimae Mexican Jay • The 7 Aphelocoma ultramarina traditional subspecies are given in abbreviations (A. ul. Xxx) on the map, the colors refer to the 4 groups I refer to in the text now. These could be labeled as the Central group (orange), the Eastern group (yellow), the Ultramarina group (blue), and the Wollweberi group (green). The four groups are from McCormack et al. 2008. • McCormack, J. E., Peterson, A. T., Bonaccorso, E., and T. B. Smith. 2008. Speciation in the highlands of Mexico: genetic and phenotypic divergence in the Mexican jay (Aphelocoma ultramarina). Molecular Ecology 17:2505-2521.
Rapid diversification of the 3 major clades • Deep divergence in A. unicolor • Major phylogenetic structure in Mexican jays • Florida scrub-jays basal within scrub-jay complex McCormack, Peterson, Delaney, Knowles, in prep. Also could cite: Delaney et al. 2008, Auk (for scrub jays) McCormack, Peterson, Bonaccorso, and Smith 2008, Mol Ecol (for Mexican jays)
Timing of diversification in Aphelocoma McCormack, Peterson, Delaney, Knowles, in prep. • Diversification much older than previously thought • Major lineages diverging quickly in the late Miocene • Transvolcanic Mexican jay ancient lineage • Few divergence events in the major glacial period (<0.7 mya)
Unicolor Jay • Major division N and S of the Isthmus (6% div) • Pliocene • Divergences north of Isthmus potentially linked to glacial cycles • Time frame correct for when Mexico was influenced by glacial activity further north 2.8 (3.9,1.3) Honduras , Guatemala 0.5 (1.2,0.3) 3.5 (6.0,2.5) 0.9 (2.0,0.5) A. un. concolor A. un. guerrerensis A. un. oaxacae A. un. griscomi A. un. unicolor McCormack, Peterson, Delaney, Knowles, in prep.
4.6 (7.4,3.7) 1.8 (2.4,0.9) 2.8 (3.7,1.5) • Genetic variation is not continuous & no haplotype sharing among clades • Transvolcanic Mexican jays have a long and independent history • New major clade in the rugged mountains of the Central Plateau • Pleistocene glacial cycles likely not the cause of major diversification events (diversification preceeded the effect of glacial cycles in Mexico which were 650000 yr) • West group very structured • North to south (food diffs) • East/West division in sky islands (10,000yr isolation by dispersal barriers) A. ul. arizonae A. ul. couchii A. ul. wollweberi A. ul. gracilis A. ul. potosina A. ul. ultramarina A. ul. colimae McCormack, Peterson, Bonaccorso, & Smith Mol Ecol 2008
Conservation Implications • Widespread, common species may actually be comprised of many, relatively rare and specialized species • Areas on the fringes of a “species” distribution and therefore perhaps not all that important might be in the core of revised “species” and therefore of high significance to biodiversity • Mexican Highlands are one such area with potentially many endemic species that we currently fail to recognize
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