[dinosaur] Evolution of crocodilian nesting + keratins in skin appendages of terrestrial vertebrates (free pdfs) .
Ben Creisler <[email protected]> Thu, 12 Dec 2019 23:23:44 -0800
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--000000000000b9eef8059990be60 Content-Type: text/plain; charset="UTF-8" Content-Transfer-Encoding: Quoted-printable Ben [email protected]=0A=0ARecent non-dino papers that may be = of interest, with free pdfs:=0A=0A=0AFree pdf:=0A=0AChristopher M. Murray, = Brian I. Crother & J. Sean Doody (2019)=0AThe evolution of crocodilian nest= ing ecology and behavior.=0AEcology and Evolution (advance online publicati= on)=0Adoi: https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__doi.org_1= 0.1002_ece3.5859&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI= &r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVk= QmguaNo8WBJAQuxfnaOIg&s=3DeV38NkO28QLosw0f5FCita7ceRcyOxHyjjtU22AcMY8&e=3D= =20=0Ahttps://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__onlinelibrary.= wiley.com_doi_10.1002_ece3.5859&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHh= pN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjm= GQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3DZ4Y5RFLn3chOmfkvmTEMLw2DhSH2cOd-8o= lBwTcHsyc&e=3D=20=0Ahttps://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__= onlinelibrary.wiley.com_doi_10.1002_ece3.5859&d=3DDwIFaQ&c=3DclK7kQUTWtAVEO= VIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9= SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3DZ4Y5RFLn3chOmfkvmTEM= Lw2DhSH2cOd-8olBwTcHsyc&e=3D=20=0A=0AFree pdf:=0Ahttps://urldefense.proofpo= int.com/v2/url?u=3Dhttps-3A__onlinelibrary.wiley.com_doi_pdf_10.1002_ece3.5= 859&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IF= aUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQ= uxfnaOIg&s=3DHqxpakvJRM5JjLr88pV_v96UrnQoo34uC82-MhbnTBs&e=3D=20=0A=0ACroco= dilians comprise an ancient and successful lineage of archosaurs that=0Arep= eatedly raises questions on how they survived a mass extinction and=0Aremai= ned relatively unchanged for ~100 million years. Was their success due=0Ato= the change=E2=80=90resistant retention of a specific set of traits over ti= me=0A(phylogenetic conservatism) or due to flexible, generalist capabilitie= s=0A(e.g., catholic diets, phenotypic plasticity in behavior), or some=0Aco= mbination of these? We examined the evolution of reproductive ecology and= =0Abehavior of crocodilians within a phylogenetic perspective, using 14 tra= its=0Afor all 24 species to determine whether these traits were phylogeneti= cally=0Aconstrained versus (ecologically) convergent. Our analysis revealed= that=0Athe ancestral crocodilian was a mound nester that exhibited both ne= st=0Aattendance and defense. Nesting mode exhibited 4=E2=80=935 transformat= ions from=0Amound to hole nesting, a convergence of which habitat may have = been a=0Adriving factor. Hole nesters were more likely to nest communally, = but this=0Aassociation may be biased by scale. Although there were exceptio= ns, mound=0Anesters typically nested during the wet season and hole nesters= during the=0Adry season; this trait was relatively conserved, however. Abo= ut two=E2=80=90thirds=0Aof species timed their nesting with the wet season,= while the other third=0Atimed their hatching with the onset of the wet sea= son. Nest attendance and=0Adefense were nearly ubiquitous and thus exhibite= d phylogenetic=0Aconservatism, but attendance lodging was diverse among spe= cies, showing=0Amultiple reversals between water and burrows. Collectively,= our analysis=0Areveals that reproductive trait evolution in crocodilians r= eflects=0Aphylogenetic constraint (nest attendance, nest defense), ecologic= al=0Aconvergence (seasonal timing of nesting, nest attendance lodging), or = both=0A(mode of nesting). Some traits (e.g., communal nesting and mode of n= esting)=0Awere autocorrelated. Our analysis provides a framework for addres= sing=0Ahypotheses raised for why there has been trait convergence in reprod= uctive=0Aecology and behavior in crocodilians and why some traits remained= =0Aphylogenetically conserved.=0A=0A=0A=3D=3D=3D=0A=0AFree pdf:=0A=0AFloria= n Ehrlich, Julia Lachner, Marcela Hermann, Erwin Tschachler & Leopold=0AEck= hart (2019)=0AConvergent evolution of cysteine-rich keratins in hard skin a= ppendages of=0Aterrestrial vertebrates.=0AMolecular Biology and Evolution, = msz279,=0Adoi: https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__doi.o= rg_10.1093_molbev_msz279&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7= CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZY= KXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3DzQw-Yii3H5klENiXQaX20S7c11CCQikdHMRaTX__R= Bc&e=3D=20=0Ahttps://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__academi= c.oup.com_mbe_advance-2Darticle_doi_10.1093_molbev_msz279_5652086&d=3DDwIFa= Q&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZ= gecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3D= HunNReCsmYMAoWf7UYYwoeecC-lQ6xqLhQiWHAtp7Z0&e=3D=20=0A=0A=0ATerrestrial ver= tebrates have evolved hard skin appendages, such as scales,=0Aclaws, feathe= rs and hair that play crucial roles in defense, predation,=0Alocomotion, an= d thermal insulation. The mechanical properties of these skin=0Aappendages = are largely determined by cornified epithelial components.=0ASo-called "hai= r keratins", cysteine-rich intermediate filament proteins=0Athat undergo co= valent cross-linking via disulfide bonds, are the crucial=0Astructural prot= eins of hair and claws in mammals and hair keratin orthologs=0Aare also pre= sent in lizard claws, indicating an evolutionary origin in a=0Ahairless com= mon ancestor of amniotes. Here, we show that reptiles and birds=0Ahave also= other cysteine-rich keratins which lack cysteine-rich orthologs=0Ain mamma= ls. In addition to hard acidic (type I) sauropsid-specific (HAS)=0Akeratins= , we identified hard basic (type II) sauropsid-specific (HBS)=0Akeratins wh= ich are conserved in lepidosaurs, turtles, crocodilians and=0Abirds. Immuno= histochemical analysis with a newly made antibody revealed=0Aexpression of = chicken HBS1 keratin in the cornifying epithelial cells of=0Afeathers. Mole= cular phylogenetics suggested that the high cysteine contents=0Aof HAS and = HBS keratins evolved independently from the cysteine-rich=0Asequences of ha= ir keratin orthologs, thus representing products of=0Aconvergent evolution.= In conclusion, we propose an evolutionary model in=0Awhich HAS and HBS ker= atins evolved as structural proteins in epithelial=0Acornification of repti= les and at least one HBS keratin was co-opted as a=0Acomponent of feathers = after the evolutionary divergence of birds from=0Areptiles. Thus, cytoskele= tal proteins of hair and feathers are products of=0Aconvergent evolution an= d evolutionary co-option to similar biomechanical=0Afunctions in clade-spec= ific hard skin appendages.=0A= --000000000000b9eef8059990be60 Content-Type: text/html; charset="UTF-8" Content-Transfer-Encoding: Quoted-printable <div dir=3D"ltr"><div><br></div>Ben Creisler<div><a href=3D"mailto:bcreisle= [email protected]">[email protected]</a></div><div><br></div><div>Recent non-di= no papers that may be of interest, with free pdfs:<br><div><div><br></div><= div><br></div><div>Free pdf:</div><div><br>Christopher M. Murray, Brian I. = Crother & J. Sean Doody (2019)<br>The evolution of crocodilian nesting = ecology and behavior.<br>Ecology and Evolution (advance online publication)= <br>doi: <a href=3D"https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__= doi.org_10.1002_ece3.5859&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOU= HhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m= =3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3DeV38NkO28QLosw0f5FCi= ta7ceRcyOxHyjjtU22AcMY8&e=3D">https://doi.org/10.1002/ece3.5859</a><br>= <a href=3D"https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__onlinelib= rary.wiley.com_doi_10.1002_ece3.5859&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOV= Igvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJg= FW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3DZ4Y5RFLn3= chOmfkvmTEMLw2DhSH2cOd-8olBwTcHsyc&e=3D">https://onlinelibrary.wiley.co= m/doi/10.1002/ece3.5859</a><br><a href=3D"https://urldefense.proofpoint.com= /v2/url?u=3Dhttps-3A__onlinelibrary.wiley.com_doi_10.1002_ece3.5859&d= =3DDwMFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO= 4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaN= o8WBJAQuxfnaOIg&s=3DZ4Y5RFLn3chOmfkvmTEMLw2DhSH2cOd-8olBwTcHsyc&e= =3D">https://onlinelibrary.wiley.com/doi/10.1002/ece3.5859</a><br><br>Free = pdf:<br><a href=3D"https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__o= nlinelibrary.wiley.com_doi_pdf_10.1002_ece3.5859&d=3DDwMFaQ&c=3DclK= 7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRC= KHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s= =3DHqxpakvJRM5JjLr88pV_v96UrnQoo34uC82-MhbnTBs&e=3D">https://onlinelibr= ary.wiley.com/doi/pdf/10.1002/ece3.5859</a><br><br>Crocodilians comprise an= ancient and successful lineage of archosaurs that repeatedly raises questi= ons on how they survived a mass extinction and remained relatively unchange= d for ~100 million years. Was their success due to the change=E2=80=90resis= tant retention of a specific set of traits over time (phylogenetic conserva= tism) or due to flexible, generalist capabilities (e.g., catholic diets, ph= enotypic plasticity in behavior), or some combination of these? We examined= the evolution of reproductive ecology and behavior of crocodilians within = a phylogenetic perspective, using 14 traits for all 24 species to determine= whether these traits were phylogenetically constrained versus (ecologicall= y) convergent. Our analysis revealed that the ancestral crocodilian was a m= ound nester that exhibited both nest attendance and defense. Nesting mode e= xhibited 4=E2=80=935 transformations from mound to hole nesting, a converge= nce of which habitat may have been a driving factor. Hole nesters were more= likely to nest communally, but this association may be biased by scale. Al= though there were exceptions, mound nesters typically nested during the wet= season and hole nesters during the dry season; this trait was relatively c= onserved, however. About two=E2=80=90thirds of species timed their nesting = with the wet season, while the other third timed their hatching with the on= set of the wet season. Nest attendance and defense were nearly ubiquitous a= nd thus exhibited phylogenetic conservatism, but attendance lodging was div= erse among species, showing multiple reversals between water and burrows. C= ollectively, our analysis reveals that reproductive trait evolution in croc= odilians reflects phylogenetic constraint (nest attendance, nest defense), = ecological convergence (seasonal timing of nesting, nest attendance lodging= ), or both (mode of nesting). Some traits (e.g., communal nesting and mode = of nesting) were autocorrelated. Our analysis provides a framework for addr= essing hypotheses raised for why there has been trait convergence in reprod= uctive ecology and behavior in crocodilians and why some traits remained ph= ylogenetically conserved.<br><br><br>=3D=3D=3D</div><div><br></div><div>Fre= e pdf:<br><br>Florian Ehrlich, Julia Lachner, Marcela Hermann, Erwin Tschac= hler & Leopold Eckhart (2019)<br>Convergent evolution of cysteine-rich = keratins in hard skin appendages of terrestrial vertebrates.<br>Molecular B= iology and Evolution, msz279, <br>doi: <a href=3D"https://urldefense.proofp= oint.com/v2/url?u=3Dhttps-3A__doi.org_10.1093_molbev_msz279&d=3DDwMFaQ&= amp;c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_= Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxf= naOIg&s=3DzQw-Yii3H5klENiXQaX20S7c11CCQikdHMRaTX__RBc&e=3D">https:/= /doi.org/10.1093/molbev/msz279</a><br><a href=3D"https://urldefense.proofpo= int.com/v2/url?u=3Dhttps-3A__academic.oup.com_mbe_advance-2Darticle_doi_10.= 1093_molbev_msz279_5652086&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BO= UHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&= m=3DYhZkUjmGQX0_ZYKXAFBEVkQmguaNo8WBJAQuxfnaOIg&s=3DHunNReCsmYMAoWf7UYY= woeecC-lQ6xqLhQiWHAtp7Z0&e=3D">https://academic.oup.com/mbe/advance-art= icle/doi/10.1093/molbev/msz279/5652086</a></div><div><br><br>Terrestrial ve= rtebrates have evolved hard skin appendages, such as scales, claws, feather= s and hair that play crucial roles in defense, predation, locomotion, and t= hermal insulation. The mechanical properties of these skin appendages are l= argely determined by cornified epithelial components. So-called "hair = keratins", cysteine-rich intermediate filament proteins that undergo c= ovalent cross-linking via disulfide bonds, are the crucial structural prote= ins of hair and claws in mammals and hair keratin orthologs are also presen= t in lizard claws, indicating an evolutionary origin in a hairless common a= ncestor of amniotes. Here, we show that reptiles and birds have also other = cysteine-rich keratins which lack cysteine-rich orthologs in mammals. In ad= dition to hard acidic (type I) sauropsid-specific (HAS) keratins, we identi= fied hard basic (type II) sauropsid-specific (HBS) keratins which are conse= rved in lepidosaurs, turtles, crocodilians and birds. Immunohistochemical a= nalysis with a newly made antibody revealed expression of chicken HBS1 kera= tin in the cornifying epithelial cells of feathers. Molecular phylogenetics= suggested that the high cysteine contents of HAS and HBS keratins evolved = independently from the cysteine-rich sequences of hair keratin orthologs, t= hus representing products of convergent evolution. In conclusion, we propos= e an evolutionary model in which HAS and HBS keratins evolved as structural= proteins in epithelial cornification of reptiles and at least one HBS kera= tin was co-opted as a component of feathers after the evolutionary divergen= ce of birds from reptiles. Thus, cytoskeletal proteins of hair and feathers= are products of convergent evolution and evolutionary co-option to similar= biomechanical functions in clade-specific hard skin appendages.<br><br><br= ></div><div><br></div></div></div></div>=0A= --000000000000b9eef8059990be60--