[dinosaur] Hadrosaur skin with 3D preservation (free pdf)
Ben Creisler <[email protected]> Tue, 31 Dec 2019 08:09:43 -0800
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--000000000000d5df90059b0230c3 Content-Type: text/plain; charset="UTF-8" Content-Transfer-Encoding: Quoted-printable Ben [email protected]=0A=0AA new paper with free pdf:=0A=0AMat= teo Fabbri, Jasmina Wiemann, Fabio Manucci & Derek E. G. Briggs (2019)=0A= Three=E2=80=90dimensional soft tissue preservation revealed in the skin of = a=0Anon=E2=80=90avian dinosaur.=0APalaeontology (advance online publication= )=0Adoi: https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__doi.org_10= .1111_pala.12470&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI= &r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DvCozh0pMCUdSu291HAXhXw= OQMZdqGGu0z5ujJVAvvqE&s=3DGtbA3tInIsADQOKiwL71wY0_2wfl9qA5GIdCuBUe8WU&e=3D= =20=0Ahttps://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__onlinelibrary.= wiley.com_doi_full_10.1111_pala.12470&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU= 5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3Dv= Cozh0pMCUdSu291HAXhXwOQMZdqGGu0z5ujJVAvvqE&s=3Dj7Dcq-P9M_iR340hUIVQC0rMuyKy= Ln_ALSWp80DdA7Q&e=3D=20=0A=0AFree pdf:=0Ahttps://urldefense.proofpoint.com/= v2/url?u=3Dhttps-3A__onlinelibrary.wiley.com_doi_pdf_10.1111_pala.12470&d= =3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGo= f_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DvCozh0pMCUdSu291HAXhXwOQMZdqGGu0z5ujJVAv= vqE&s=3DEdM6TiN1VCK9oRJeNonNi45tiEY-h0dP7GuREoVCiZI&e=3D=20=0A=0AData archi= ving statement:=0AData for this study are available in the Dryad Digital Re= pository:=0Ahttps://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__doi.org_= 10.5061_dryad.3bk3j9kf1&d=3DDwIFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7C= Sfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DvCozh0pMCUdSu29= 1HAXhXwOQMZdqGGu0z5ujJVAvvqE&s=3DmLUEpblrrOtauOLRKUHEuXdR3CHIoY61tvMoLb205B= U&e=3D=20=0A=0A=0AThe most commonly preserved soft tissues associated with = ornithischian=0Adinosaurs are skin remains. The apparent resistance of hadr= osaur skin to=0Adecay, and its abundance in the fossil record relative to t= hat of other=0Atetrapods, has been attributed to factors such as thickness = and=0Acomposition. Here we report additional intrinsic factors within hadro= saur=0Askin: 3D=E2=80=90preserved eumelanin=E2=80=90bearing bodies, dermal = cells and blood vessel=0Afragments in an organic matrix composed of protein= fossilization products.=0AThe skin is much thinner than that of living mam= mals of similar size. It is=0Alikely that the preservation of hadrosaur ski= n is related to the=0Aarrangement of the layers composing it.=0A=0AThe skin= is the largest organ in the body of any vertebrate and carries out=0Aimpor= tant functions such as homeostasis, sensory reception and visual=0Asignalin= g (Landmann 1986). The skin is composed of three layers: the=0Aepidermis (o= uter), dermis and the subcutis (innermost). The epidermis is=0Acomposed of = stratified layers of keratinized cells (the stratum corneum,=0Astratum inte= rmedium and stratum basale) whereas the dermis is characterized=0Aby a stra= tum spongiosum of loose collagen fibres and a lower stratum=0Acompactum of = tightly packed, orthogonal arrays of collagen fibres (Landmann=0A1986; Coop= er & Greenberg 1992). Colouration of the skin is generally=0Adetermined by = melanophores and chromatophores and the structural layering=0Aof the integu= ment (McNamara et al. 2016; Shawkey & D'Alba 2017).=0AChromatophores and me= lanophores are usually stored at the interface between=0Athe epidermis and = dermis in vertebrates (Prum & Torres 2003, 2004; Chang et=0Aal. 2009; McNam= ara et al. 2016; Shawkey & D'Alba 2017). The innermost layer=0Aof the skin,= the subcutaneous layer, separates the integument from the rest=0Aof the bo= dy. Integument is probably the most abundant type of soft tissue=0Ain the v= ertebrate fossil record (Schweitzer 2011; Davis 2014) and it is=0Aproviding= increasingly important evidence of reptile evolution and=0Aecological dive= rsity in the past (e.g. Martill 1995; Briggs et al. 1997;=0AMartill et al. = 2000; Bell 2012; Lindgren et al. 2013, 2014, 2018; Davis=0A2014; McNamara e= t al. 2016, 2018; Yang et al. 2019).=0A=0AFossilized skin is preserved in t= hree main ways: as impressions,=0Acompressions or permineralized, three=E2= =80=90dimensional skin (Schweitzer 2011).=0AImpressions are occasionally fo= und in association with bones or footprints=0A(e.g. Lockley et al. 2004; Pa= ik et al. 2017). Compressions are preserved as=0Aa carbon=E2=80=90rich laye= r surrounding bones (e.g. Briggs et al. 1997; Qiang et=0Aal. 1998; Wang et = al. 2019). Permineralized skin is the rarest of the three=0Amodes of preser= vation (Schweitzer 2011; Bell 2012; Davis 2014) and usually=0Aoccurs in ass= ociation with three dimensionally preserved skeletons.=0APermineralized ski= n was previously thought to be a high=E2=80=90fidelity=0Amineralized replic= ation of the original organic tissue (Schweitzer 2011).=0AHowever, recent i= nvestigations of a few exceptionally preserved examples=0Ahave challenged t= his view by recovering original organic molecules.=0ADinosaurian integument= preserved as a result of early mineralization is=0Aknown in Ornithischia, = Sauropoda and Theropoda (Briggs et al. 1997; Martill=0Aet al. 2000; Bell 20= 12; Davis 2014; McNamara et al. 2018). The preservation=0Aof integument is = most prevalent in hadrosaurs (Bell 2012; Davis 2014).=0AHowever, the taphon= omic processes that favour the fossilization of skin and=0Aassociated molec= ular preservation are still poorly understood (Lyson &=0ALongrich 2011; Bel= l 2012; Davis 2014). Previous investigations of dinosaur=0Askin were perfor= med on the basal ceratopsian Psittacosaurus (Lingham=E2=80=90Soliar=0A2008)= , a nodosaur (Brown et al. 2017), and coelurosaurs (Naval=C3=B3n et al.=0A2= 015; Bell et al. 2017; McNamara et al. 2018). Traces of collagenous=0Amater= ial were found in a =E2=80=98mummified=E2=80=99 hadrosaurid from North Dako= ta, USA=0A(Manning et al. 2009). Skin colouration patterns have been recons= tructed in=0Aornithischian dinosaurs (Lingham=E2=80=90Soliar & Plodowski 20= 10; Vinther et al.=0A2016; Brown et al. 2017) but not in a hadrosaurid.=0A= =0AYPMPU 016969 is a partial hadrosaur skeleton preserving skin in the flan= k=0Aregion of the body (Fabbri et al. 2019, fig. S1). We investigated this= =0Aspecimen with the following aims: (1) to understand its taphonomy; (2) t= o=0Atest for soft tissue preservation and the presence of melanin; and (3) = to=0Aconsider previous explanations for the prevalence of hadrosaur skin=0A= preservation in the fossil record.=0A=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=0A= --000000000000d5df90059b0230c3 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>A new paper w= ith free pdf:<br><div><br></div><div>Matteo Fabbri, =C2=A0Jasmina Wiemann, = =C2=A0Fabio Manucci & Derek E. G. Briggs (2019)<br>Three=E2=80=90dimens= ional soft tissue preservation revealed in the skin of a non=E2=80=90avian = dinosaur.<br>Palaeontology (advance online publication)<br>doi: =C2=A0<a hr= ef=3D"https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__doi.org_10.111= 1_pala.12470&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p7CSfn= c_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DvCozh0pMCU= dSu291HAXhXwOQMZdqGGu0z5ujJVAvvqE&s=3DGtbA3tInIsADQOKiwL71wY0_2wfl9qA5G= IdCuBUe8WU&e=3D">https://doi.org/10.1111/pala.12470</a><br><a href=3D"h= ttps://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__onlinelibrary.wiley.c= om_doi_full_10.1111_pala.12470&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOVIgvi0N= U5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&= amp;m=3DvCozh0pMCUdSu291HAXhXwOQMZdqGGu0z5ujJVAvvqE&s=3Dj7Dcq-P9M_iR340= hUIVQC0rMuyKyLn_ALSWp80DdA7Q&e=3D">https://onlinelibrary.wiley.com/doi/= full/10.1111/pala.12470</a></div><div><br></div><div>Free pdf:<br><a href= =3D"https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__onlinelibrary.wi= ley.com_doi_pdf_10.1111_pala.12470&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOVIg= vi0NU5BOUHhpN0H8p7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW= 9SI&m=3DvCozh0pMCUdSu291HAXhXwOQMZdqGGu0z5ujJVAvvqE&s=3DEdM6TiN1VCK= 9oRJeNonNi45tiEY-h0dP7GuREoVCiZI&e=3D">https://onlinelibrary.wiley.com/= doi/pdf/10.1111/pala.12470</a><br></div><div><br>Data archiving statement:<= br>Data for this study are available in the Dryad Digital Repository: <a hr= ef=3D"https://urldefense.proofpoint.com/v2/url?u=3Dhttps-3A__doi.org_10.506= 1_dryad.3bk3j9kf1&d=3DDwMFaQ&c=3DclK7kQUTWtAVEOVIgvi0NU5BOUHhpN0H8p= 7CSfnc_gI&r=3DRy_mO4IFaUmGof_Yl9MyZgecRCKHn5g4z1CYJgFW9SI&m=3DvCozh= 0pMCUdSu291HAXhXwOQMZdqGGu0z5ujJVAvvqE&s=3DmLUEpblrrOtauOLRKUHEuXdR3CHI= oY61tvMoLb205BU&e=3D">https://doi.org/10.5061/dryad.3bk3j9kf1</a><br><b= r><br>The most commonly preserved soft tissues associated with ornithischia= n dinosaurs are skin remains. The apparent resistance of hadrosaur skin to = decay, and its abundance in the fossil record relative to that of other tet= rapods, has been attributed to factors such as thickness and composition. H= ere we report additional intrinsic factors within hadrosaur skin: 3D=E2=80= =90preserved eumelanin=E2=80=90bearing bodies, dermal cells and blood vesse= l fragments in an organic matrix composed of protein fossilization products= . The skin is much thinner than that of living mammals of similar size. It = is likely that the preservation of hadrosaur skin is related to the arrange= ment of the layers composing it.<br><br>The skin is the largest organ in th= e body of any vertebrate and carries out important functions such as homeos= tasis, sensory reception and visual signaling (Landmann 1986). The skin is = composed of three layers: the epidermis (outer), dermis and the subcutis (i= nnermost). The epidermis is composed of stratified layers of keratinized ce= lls (the stratum corneum, stratum intermedium and stratum basale) whereas t= he dermis is characterized by a stratum spongiosum of loose collagen fibres= and a lower stratum compactum of tightly packed, orthogonal arrays of coll= agen fibres (Landmann 1986; Cooper & Greenberg 1992). Colouration of th= e skin is generally determined by melanophores and chromatophores and the s= tructural layering of the integument (McNamara et al. 2016; Shawkey & D= 'Alba 2017). Chromatophores and melanophores are usually stored at the = interface between the epidermis and dermis in vertebrates (Prum & Torre= s 2003, 2004; Chang et al. 2009; McNamara et al. 2016; Shawkey & D'= Alba 2017). The innermost layer of the skin, the subcutaneous layer, separa= tes the integument from the rest of the body. Integument is probably the mo= st abundant type of soft tissue in the vertebrate fossil record (Schweitzer= 2011; Davis 2014) and it is providing increasingly important evidence of r= eptile evolution and ecological diversity in the past (e.g. Martill 1995; B= riggs et al. 1997; Martill et al. 2000; Bell 2012; Lindgren et al. 2013, 20= 14, 2018; Davis 2014; McNamara et al. 2016, 2018; Yang et al. 2019).<br><br= >Fossilized skin is preserved in three main ways: as impressions, compressi= ons or permineralized, three=E2=80=90dimensional skin (Schweitzer 2011). Im= pressions are occasionally found in association with bones or footprints (e= .g. Lockley et al. 2004; Paik et al. 2017). Compressions are preserved as a= carbon=E2=80=90rich layer surrounding bones (e.g. Briggs et al. 1997; Qian= g et al. 1998; Wang et al. 2019). Permineralized skin is the rarest of the = three modes of preservation (Schweitzer 2011; Bell 2012; Davis 2014) and us= ually occurs in association with three dimensionally preserved skeletons. P= ermineralized skin was previously thought to be a high=E2=80=90fidelity min= eralized replication of the original organic tissue (Schweitzer 2011). Howe= ver, recent investigations of a few exceptionally preserved examples have c= hallenged this view by recovering original organic molecules. Dinosaurian i= ntegument preserved as a result of early mineralization is known in Ornithi= schia, Sauropoda and Theropoda (Briggs et al. 1997; Martill et al. 2000; Be= ll 2012; Davis 2014; McNamara et al. 2018). The preservation of integument = is most prevalent in hadrosaurs (Bell 2012; Davis 2014). However, the tapho= nomic processes that favour the fossilization of skin and associated molecu= lar preservation are still poorly understood (Lyson & Longrich 2011; Be= ll 2012; Davis 2014). Previous investigations of dinosaur skin were perform= ed on the basal ceratopsian Psittacosaurus (Lingham=E2=80=90Soliar 2008), a= nodosaur (Brown et al. 2017), and coelurosaurs (Naval=C3=B3n et al. 2015; = Bell et al. 2017; McNamara et al. 2018). Traces of collagenous material wer= e found in a =E2=80=98mummified=E2=80=99 hadrosaurid from North Dakota, USA= (Manning et al. 2009). Skin colouration patterns have been reconstructed i= n ornithischian dinosaurs (Lingham=E2=80=90Soliar & Plodowski 2010; Vin= ther et al. 2016; Brown et al. 2017) but not in a hadrosaurid.<br><br>YPMPU= 016969 is a partial hadrosaur skeleton preserving skin in the flank region= of the body (Fabbri et al. 2019, fig. S1). We investigated this specimen w= ith the following aims: (1) to understand its taphonomy; (2) to test for so= ft tissue preservation and the presence of melanin; and (3) to consider pre= vious explanations for the prevalence of hadrosaur skin preservation in the= fossil record.<br>=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D=3D<br></div></div></div>= =0A= --000000000000d5df90059b0230c3--