M. JORGE GUIMARÃES ORIGINAL DISCOVERY EDITION
The Rediscovery of Archaeopteryx or How the most famous fossil in the world changes the tree of life
VOLUME I Non-rigid eggs in the bird lineage The Rediscovery of Archaeopteryx
Volume I Non-rigid eggs in the bird lineage !"#$%&'()*+*,-./*0$*12*3"%'4*56&78%94: ;<<*%&'():*%4:4%=4>2*
?(&:* 0""@* 78$* A")* 04* %4#%">6B4>C* &A* D("<4* "%* &A* #8%)C* &AB<6>&A'* &<<6:)%8)&"A:C* &A* 8A$* E"%7* F04$"A>* )(4* B"#$&A'* #4%7&))4>*0$*G4B)&"A:*.-H*8A>*.-/*"E *)(4*IG*!"#$%&'()*J8D* 8A>*4KB4#)*0$*%4=&4D4%:*E"%*)(4*#60<&B*#%4::LC*D&)("6)*D%&))4A* #4%7&::&"A*0$*)(4*#60<&:(4%:2
?(&:* 0""@* 78$* 04* #6%B(8:4>* &A* M68A)&)$* E"%* 4>6B8)&"A8
?(&:* 0""@* &:* 6A>4%* B"#$%&'()* #%")4B)&"A2* R)* D8:* %4'&:)4%4>* 8)* RA:#4S9"O54%8<* >8:* ;)&=&>8>4:* !6<)6%8&:C* N"%)6'8
N60<&:(4>*0$*;JUV?*J&E4*GB&4AB4:*!"7#6)&A'C*N"%)6'8<2 W4:&'A4>* 0$* N4>%"* Ga* 8A>* !
?"*B"A)8B)*)(4*86)("%*#<48:4*:4A>*8A*4O78&<*)"P* e"%'42'6&78%84:Q8<4%)O"A<&A42B"7 CONTENTS
!"!#$%&#'( 8
'&)"*+%$,('-."!/0"(&1 9
2*"3%$" ::
4'',+1500%*6 :;
#(&*'!5$&'*6+ ('&"+ '(+ &)"+ 0'1&+ 3%0'51+ 3'11#.+ #(+ &)"+ :8 -'*.!+%(!+#&1+*"!#1$'7"*6
#(&*'!5$&#'(+&'+7'.50"+# <8
*"15.&1 <9 .2 5<"06<8%*%8>&">4A:4*:)%6B)6%4:*<&4*=4A)%8<*)"*)(4*^4%<&A*:#4B&74A*8A>*A48%*&):*(48> c. ,2 ;A*4''O4AB<":4>*E4)6:*D&)(*8*)"")(4>*048@*<&4:*=4A)%8<*>"*)(4*^4%<&A*:#4B&74A /. d2 U=&>4AB4*"E *4'':*&A*)(4*#4<=&B*%4'&"A*"E*)(4*^4%<&A*:#4B&74A ._H _2 ;A*4''O4AB<":4>*E4)6:*&A*)(4*E%"A)*:6%E8B4*"E*)(4*^4%<&A*:#4B&74Af:*78&A*:<80 .cc c2 X%&'&A8<&)$*"E *)(4*:<80*#%"#4%*"E *)(4*^4%<&A*:#4B&74Af:*B"6A)4%:<80 .gH g2 !8%0"A8B4"6:* 4'':* 8A>* :6%E8B4* 4K#":4>* E4)6:4:* <"B8)4>* 8A)4%&"%<$* )"* )(4* ^4%<&A* ,-. :#4B&74A H2 !"#$%&'()")*+,#)-"&).$#"/0.1"(%,##)-"102-3045#6"'"728)%"(3'/"'2-"'"#(3)%,.0("%028 ,gd /2 9.1)%"72-028#"02".1)"'%)'"'2.)%0,%".,".1)":)%302"#+)(04)2 ,b. b2 ;A*&:"<8)4>*4''*&A*)(4*^4%<&A*:#4B&74Af:*B"6A)4%:<80 d-c .-2 ?(%44*=&:&0<4*E4)6:4:*8%"6A>*)(4*^4%<&A*:#4B&74Af:*0">$ ddH ..2 ?(4*A4:)&A'*^4%<&A*:#4B&74A dcd .,2 ;*E4)6:OB"A)8&A&A'*4''*8<:"*<&4:*=4A)%8<*)"*)(4*?4$<4%:*:#4B&74A* dgd .d2 X)(4%*4''O<&@4*:)%6B)6%4:*&A*)(4*?4$<4%:*:#4B&74Af:*E"::&< _-. ._2 U=&>4AB4*"E *4'':*&A*)(4*18K04%'*:#4B&74Af:*E"::&< _,. .c2 ?(4*7"%#("<"'$*"E *!"#$%&'()&"*+f:*4'': _,c .g2 ?(4*4'':(4<<*"E *!"#$%&'()&"*+f:*4'': _dd*
!#1$511#'( <<; .2 ?(4*^4%<&A*!"#$%&'()&"*+*&:*)(4*E"::&<*"E *8*'%"6A>*A4:)&A'*8A&78<*8A>*A")*"E *8*:6A@* __H B8%B8:: ,2 ]"AO%&'&>*4'':*&A:)48>*"E *4''*E"<<&B<4: __b* d2 ;-)2.07('.0,2"'#"!"#$%&'()&"*+*4'': _cc _2 UKB4#)&"A8<*E"::&<*#%4:4%=8)&"A _cH c2 !"A)4K)68<&h8)&"A*"E *)(4*>&:B"=4%$*"E *A"AO%&'&>*4'':*&A*)(4*0&%>*<&A48'4 _cb g2 R7#<&B8)&"A:*E"%*)(4*"%&'&A*"E *0&%>:*8A>*#%"#":4>*($#")(4:&:*E"%*)(4*"BB6%%4AB4*"E * _g. >&EE4%4A)*4''*B(8%8B)4%&:)&B:*87"A'*B<":4<$*%4<8)4>*8A&78<: H2 ;4+30('.0,2#"&,%".1)")<,3$.0,2",& "&)'.1)%#"'2-"=081. _gd /2 i(8)*(8##4A4>*)"*)(4*"%&'&A8<*E"::&<*"E *)(4*J"A>"A*:#4B&74A*"E *!"#$%&'()&"*+j _gc b2 R7#<&B8)&"A:*E"%*E6)6%4*E"::&<*(8A><&A'C*#%4#8%8)&"A*8A>*:)6>$ _gH .-2 T"D*)"*>&:#<8$*)(4:4*E"::&<: _H.
35&5*"+-'*,+%(!+#(&*'!5$&#'(+&'+&)"+("=&+7'.50"1 <8?
1522."0"(&%*6+0%&"*#%. <88
0%&"*#%.1+%(!+0"&)'!1 ;??
*"3"*"($"1 ;
%5&)'*+#(3'*0%&#'(+%(!+*">5"1&1+3'*+0%&"*#%.1 ;<9
0"0'*#"1+'3+&)#1+*"1"%*$) ;;: The Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage · 15
BOOK SUMMARY
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ʽ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he Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage · 47
INTRODUCTION TO VOLUME I
!""#$"%&'( )"( *+,,*-( .#//#0)"1( 23&( )")2)4/( <4?(54>2)4//?(&D5/4)"()27(>4>&(5>&7&>@42)#"(4"'(23&( 5$6/)%42)#"7()"(*+8+9(4"'(*+8:;(#. (0342(6&%4<&( #%%$>>&"%&(#. (.&24/(&D5#7$>&7()"(23&(.#77)/(4"'-(#"( ="#0"( 47( 23&( 23&#>?( #. ( &@#/$2)#"-( 4"'( 23&( .1)" ,.1)%6" %)#$3.)-" 02" '" ='..)2)-" .%0C-04)2#0,2'3" ')7%#@&>?( #. ( 23&( 54>2)4//?( 4>2)%$/42&'( A#"'#"( E;PF( &11( 0)23( ')..&>&"2( %#>#"4/-( 741)224/( 4"'( 75&%)<&"( #. ( !"#$%&'()&"*+( )"( *:B**-( 23&( C&>/)"( 2>4"7@&>74/(6)Q')<&"7)#"4/(#$2/)"&7G( 75&%)<&"( #. ( !"#$%&'()&"*+( 6&%4<&( 23&( )%#"( #. ( 23&( 23&#>?( #. ( &@#/$2)#"*( .#>( .&42$>)"1( 4( .$//?( H3&( %#<6)"42)#"( #. ( 23&7&( $"&D5&%2&'( &11( 4>2)%$/42&'(4")<4/(2342-($"/)=&(&D24"2(E)"(&D)72&"%&F( .&42$>&7( )"( 477#%)42)#"( 0)23( 23&)>( 7<4//( 7)J&( 4"'( 4")<4/7-( %#<6)"&'( .&423&>&'( 0)"17-( 4( 6#"?( 24)/( 23&( 0)'&75>&4'( 477$<52)#"( 2342( !"#$%&'()&"*+( 4"'( 4( 2##23&'( 6&4=G( H3)7( )7( 23&( >&47#"( 03?( )2( .#77)/)J&'( 42( 23&( 6#22#<( #. ( 4( 734//#0( /41##"( )7( %#<<#"/?( )"2&>5>&2&'( 47( >&5>&7&"2)"1( 4"( 4.2&>( 7)"=)"1( )"2#( )2-( )"72&4'( #. ( 34@)"1( >&7)'&'( &@#/$2)#"4>?( 7241&( 2342( 5>&%&'&'( 23&( 455&4>4"%&( )"(4"(&D5#7&'(542%3(#. (/4"'(2342(047(#@&>%#<&( #. (2#'4?I7(6)>'7G 5@" =,,-0286" )*+3'02" /1@" .1)#)" 72-028#" /)2." $""#2)%&'(.#>(#@&>(*RS(?&4>7('&75)2&(23&(')75/4?( !"( )"@&72)142)#"( )"2#( 23&( ')7>$52)@&( 3?5#23&7)7( #. ( 23&7&( .#77)/7( 6&.#>&( <)//)#"7( #. ( @)7)2#>7( 4"'( 2342( )27( .#77)/)J&'( 5#7&( %#>>&75#"'7( 2#( 4( "&72)"1( 7%>$2)"?(6?(1&"&>42)#"7(#. (>&7&4>%3&>7G 5#72$>&-( >423&>( 234"( K( 47( %#<<#"/?( 6&/)&@&'( )"( .10#"7)3-"F"5)028".1)")2-"%)#$3.",& "'"('%('##",& "'2" J%)<0,$#" %)#)'%(1" /'#" &,($#)-" ,2" .1)" =081." '204'3".1'."-%0&.)-6"=,'.)-"'2-"3'.)%"#$2D"02.,".1)" %4546)/)2?( #. ( 23)7( &D2)"%2( 4")<4/-( 03)%3( 4/7#( 6#22#<(#. (4(L#/"3#.&"(/41##"*-()7('&7%>)6&'(3&>&( %#"2>)6$2&'( 2#( 23&( /4%=( #. ( >&%#1")2)#"( #. ( 23&( )"(M#/$<&(N(#. (H3&(O&')7%#@&>?(#. (!"#$%&'()&"*+G .#77)/)J42)#"( 5#7&( #. ( !"#$%&'()&"*+( 75&%)<&"7( 47( 5#77)6/?( %#>>&75#"')"1( 2#( ')..&>&"2( @)&07( #. ( 4( !"( $"5>&%&'&"2&'( '&1>&&( #. ( &D%&52)#"4/( "&72)"1(5#7)2)#"G( 5>&7&>@42)#"(34'(1#"&($""#2)%&'("#2(#"/?()"(23&( C&>/)"(75&%)<&"I7(.#77)/(6$2(4/7#()"(#23&>(.#77)/7(#. ( ;2"'--0.0,26".1)"72-028#"%)+,%.)-"1)%)"-)4'2-)-" !"#$%&'()&"*+(75&%)<&"7G( 5>#/#"1&'( 4"'( >&5&42&'( 7%>$2)"?( .#>( #@&>( 23>&&( ?&4>7( #. ( 4//( 475&%27( #. ( 6#23( 7/467( #. ( 23&( C&>/)"( >)2#'.0,2'33@6" '" (,2('.)2'.0,2" ,& " 72-028#" 02" 75&%)<&"( .#77)/( 2#( @&>).?( 23&( "&72)"1( 3?5#23&7)7( 7$55#>2( #. ( 1>#$"'( "&72)"1( 6?( 23&7&( 4")<4/7( )7( #. (!"#$%&'()&"*+-()"%/$')"1(23&(64%=(7$>.4%&(#. ()27( 5>&7&"2&'()"(23)7(6##=(@#/$<&-()"%/$')"1(23&(=&?( <4)"(7/46-(03)%3()7(>4>&/?(7&&"(6?(<#72(>&7&4>%3&>7( 72-028",& "'2")88C)2(3,#)-"&).$#"/0.1"'".,,.1)-" '$&(2#()27(&"%47&<&"2()"(4(74>%#5341$7Q/)=&(<#$"2( 6&4=( 4<#"1( #23&>( 5$242)@&( &117( .#$"'( @&"2>4//?( 2342(/&4@&7(#"/?(23&(.>#"2(7$>.4%&(&D5#7&'G 2#(23&(C&>/)"(75&%)<&"I7(6#'?-(47(0&//(47(&D5#7&'( 3&4'( 2$%=&'( .&2$7&7( 0)23( 7$>.4%&( <#>53#/#1?( H3&( ')7%#@&>?( #. ( "#"Q>)1)'( &117( )"( 7&@&>4/( 5>&7&>@42)#"( )"( 6#23( 23&( C&>/)"( 4"'( H&?/&>7( 75&%)<&"7(#. (23)7(6)>'Q>&/42&'(4")<4/(4"'(>&7$/2)"1( !"#$%&'()&"*+(.#77)/7G( )<5/)%42)#"7( 0)//( 6&( 4"4/?J&'( 3&>&T( 342%3)"1( 4")<4/7( 4"'( %/)"1)"1( 342%3/)"17( .>#<( 23&( C&>/)"( L$55#>2)@&( &@)'&"%&( #. ( 4( 23)"( 4"'( '&.#><46/&( 4"'(H&?/&>7(.#77)/7(0)//(6&('&7%>)6&'()"(M#/$<&(NNR( &1173&//( )7( '&7%>)6&'-( 03)%3( #"( 23&( #"&( 34"'( 4"'(23&(C&>/)"(!"#$%&'()&"*+("&72()"(M#/$<&(NNNG The Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage · 543
REFERENCES The Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage · 545
!"# $%&&'()*%+,# -"# ./0"1# !"#$%&'()&"*+# ,$&- .#'/- '0 - !:"#?7H(HC,#B"#$"#Q",#2)C(,#>"#@#T3D4(&3'O%+,#Y"# 12'34)5'/"#.2+3%0+34(#-*%3&,#5%+67'8,#9::;1" T(%# )&0%DC# !"#$%&'()&"*+# .T(%+)E)07V# AF37&7%1V# 7# '%=#DE%436%'#*+)6#C(%#a366%+30O37'bT3C()037'# 9"# <7+=3',# >"# ?"# @# A"# ?"# $7&&74%"# B'# C(%# I)H'07+8# )* # U4(76(7HEC%',# c7F7+37"# 8&&"6# &,# C%'0%'48#)* #DE%43%D#C)#*)+6#F7+3%C3%DG#7'0#)'#C(%# %J!;!#.9:!M1" E%+E%CH7C3)'# )* # F7+3%C3%D# 7'0# DE%43%D# I8# '7CH+7&# 6%7'D# )* # D%&%4C3)'"# 67- 8"'#7- 95//7- :'#7- 9'/;'/7- !!"#<8[%,#5,#]+%63+,#Q",#a73D%+,#5#7'0#<"#`73D("# <''3'=*#!,#JKLK:#.!MKM1" A# 0+)='%0# Q%D)X)34# I3+0# I+%%03'O# 4)&)'8# *+)6# C(%# S7C%# >+%C74%)HD# )* # T+7'D8&F7'37"# N"#<7+=3',#>"#./0"1#>/-)$&->"5=5/-'0 -:(&?#.-%'OH3'# D%)4"I5??&/?#$%0)&/#"",#JNKLJJ9#.9:!91" :,,D#6"KLLMN"7%#."+$530#1)-"02"OPQMIA !9"# $7'O,# d"# %C# 7&"# /OO# 744H6H&7C3)'# =3C(# N<# J"# 5H367+P%D,# Q"# R"# ./0"1# ,$&- @&;5?#'2&"*- '0 - %6I+8)D#E+)F30%D#3'D3O(C#3'C)#C(%#&3*%#(3DC)+8#)* #7# !"#$%&'()&"*+- .A'37- ..1B- !"#$%&'()&"*+C?- $%)#$35/=?"# EC%+)D7H+"-:#5&/#&#!%',#!!;ZL!9:!#.9:!Z1" .AS/?T# S3*%# U43%'4%D# >)6EHC3'O,# -)+CHO7&,# 9:!M1" !N"#?%3DX,#?"#?",#/F7'D,#<"#>",#?)I%+CD,#/"#Q",#UH%D,# Y"L<"#@#A"#Q"#e7C%D"#B&0%DC#[')='#03')D7H+37'# K"# 5H367+P%D,# Q"# R"# ./0"1# ,$&- @&;5?#'2&"*- '0 - '%DC3'O#D3C%#7'0#+%E+)0H4C3F%#I3)&)O8#)* #C(%#/7+&8# !"#$%&'()&"*+#.A'37- ...1V# ,$&- !"#$%&'()&"*+C?- D&?)"# RH+7DD34# D7H+)E)0)6)+E(# J%??'?('/;*34?7- 8"'#7- .AS/?T# S3*%# U43%'4%D# >)6EHC3'O,# -)+CHO7&,# 3'# D%)37-!#%;7-:#57-K:!#$(",#9J9ML9JNN#.9:!91" E+%E7+7C3)'1" !J"#SfE%XLQ7+Cg'%X,#`",#Q)+7C7&&7,#R"R"#7'0#U7'X,# W"# U4(=%3CX%+,# Q"# Y"# U)*C# C3DDH%# E+%D%+F7C3)'# 3'# RASA"T02,#'$%#"2)#.028",2".0-'3"='.#A"8%3%&'=&'="%($*L- C%++%DC+37&#Q%D)X)34#F%+C%I+7C%D"#!//47-@&27#1%")$- 8%3%&'#35F%)'3'=*L-8%3%&'&#'3'=*-$&(,#!KNL!WN#.9:::1" 83%/&)7-:#57#!",#!MZL9!W#.9:!!1" !K"#h(%'O,#d"#&)-%37#-+%D%+F7C3)'#)*#)F7+37'#*)&&34&%D# Z"# ?%3D0)+*,# 5"# @# $HCC[%,# Q"# ?%L%F7&H7C3'O# +%F%7&D# %7+&8# %F)&HC3)'# )* # 7F37'# +%E+)0H4C3F%# Q))03%\D# BE3DC()C)'34L-)DCH+%# Y8E)C(%D3D# I%(7F3)+"#D%)4"&#)"%,#K:ZLK!!#.9:!N1" 3'# 2)DD3&# ]%+C%I+7C%D# -7+C# ^V# ?%EC3&%D# _# C(%# C7E()')68#)* #C(%#I3E%07')D7H+D#E'F(?'=/%)$4?- !W"#Q78+,#5"#@#A"#Q7'%O)&0"#>7'#)F7+37'#*)&&34&%D# 3'/=5(&?- %/;- 64"%2&/%)'"- ?)%"G5# *+)6# C(%# U)&'()*%'# *)DD3&3X%i#D%)4"&#)"",#/!#.9:!N1" A+4(3E%&7O)# .RH+7DD34,# 5%+67'81"# 8%3&'H5';52&"?5)*- %/;-8%3&'&/25"'/F&/)?#"#,#!!;L!WM#.9:!91" !Z"# >(37EE%,# S"# Q"# @# Q"# j3'Ok3'"# ./0D"1# M5";?- '0 -:)'/&7-E$5/&?&-!25%/-N'??53?-N"'F-,$&-!=&-'0 -)$&- M"#]7++344(3),#<"#R"#@#2"#<"#R74[D)'"#?%E+)0H4C3)'# O5/'?%4"?L# E7O%# !:;# .R)('\D# Y)E[3'D# l'3F%+D3C8# 3'#Q%D)X)34#I3+0D#7'0#%F)&HC3)'#)* #C(%#6)0%+'# -+%DD,#c7&C36)+%,#Q7+8&7'0,#lUA,#9:!W1" 7F37'# +%E+)0H4C3F%# 6)0%"#,$&- !4G# $!!,# WKJLWMJ# .9:!W1" !M"# h()'O(%# h()H# 7'0# 2"# h(7'O"# A# E+%4)437&# 7F37'# %6I+8)# *+)6# C(%# S)=%+# >+%C74%)HD# )* # ;"#2)C(,#>",#T3D4(&3'O%+,#Y"#@#B"#$"#Q"#?7H(HC,# >(3'7"#:#5&/#&-!(&,#WKN#.9::J1" `%=#DE%436%'#)* #!"#$%&'()&"*+#E+)F30%D#3'D3O(CD# 3'C)#C(%#%F)&HC3)'#)* #E%''74%)HD#*%7C(%+D"#D%)4"&# !;"#U4(=%3CX%+,#Q"#Y"#%C#7&"#S7C%#>+%C74%)HD#7F37'# %$$,#Z;LM9#.9:!J1" %OOD#=3C(#%6I+8)D#*+)6#A+O%'C3'7"#67-A&")7-8%3&'/)"# ##,#!;!L!;K#.9::91" 546 · The Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage
9:"#2%+'7'0%X,#Q"#U"#&)-%3"#A#&7+O%#744H6H&7C3)'#)* # 9;"# >7=C(+78,# A"# @# R"# Y%+6%D# ./0D"1# E$5#G&/- R- 7F37'#%OOD#*+)6#C(%#S7C%#>+%C74%)HD#)* #-7C7O)'37# 1==B- !/- 1==SE&/)"5#- T45;&- )'- @%5?5/=- 8'43)"*- .^F8# .A+O%'C3'71# +%F%7&D# 7# ')F%&# '%DC3'O# DC+7C%O8# )* # -+%DD,#S%=%D,#l'3C%0#a3'O0)6,#9:!K"# Q%D)X)34#I3+0D"#89>:->D1#'.J1V#%W!:N:#.9:!N1" N:"#>7+E%'C%+,#a",#Y3+D4(,#a"#2"#@#R"#?"#Y)+'%+"# 9!"# B+'%&7D# R"# 2"# `%DCD,# %OOD,# 7'0# 8)H'O# )* # ^'C+)0H4C3)'"#./#>7+E%'C%+,#a,#Y3+D4(,#a"#2"#@#R"# C(%# AXH+%L4+)='%0# YH663'OI3+0# .!F%P533%- ?"#Y)+'%+#./0D"1,#O5/'?%4"-1==?-%/;-M%H5&?,#!L!!"# #*%/'#&($%3%1"#,$&-Q53?'/-6'4"/%3-'0 ->"/5)$'3'=*#$##.N1,# .>76I+30O%#l'3F%+D3C8#-+%DD,#lUA,#!;;J1" K;9LK;Z#.9:!:1" N!"#>(H+%,#<",#TH+'%+,#>"#@#2"##-%C%+D)'"#A'#%6I+8)# 99"# Y)8C,# <"#2"#-+74C347&# 6%C()0D# )* # %DC367C3'O# )* # E%F()'?%4"4?# *+)6# C(%# Q)++3DD)'# *)+67C3)'# F)&H6%#7'0# *+%D(# =%3O(C#)* # I3+0# %OOD"#,$&-!4G# .RH+7DD34,#Q300&%#T3C()'37'1#3'#<3')D7H+#`7C3)'7&# "&,#ZNLZZ#.!;Z;1" Q)'H6%'C,#lC7("#./#>7+E%'C%+,#a,#Y3+D4(,#a"#2"#@# R"#?"#Y)+'%+#./0D"1,#O5/'?%4"-1==?-%/;-M%H5&?,#9;ML 9N"#/+34[D)',#5"#Q",#h%&%'3CD[8,#<"#a",#a78,#<"#^"#@# N!!#.>76I+30O%#l'3F%+D3C8#-+%DD,#lUA,#!;;J1" Q"#A"#`)+%&&"#<3')D7H+#3'4HI7C3)'#E%+3)0D#03+%4C&8# 0%C%+63'%0#*+)6#O+)=C(L&3'%#4)H'CD#3'#%6I+8)'34# N9"# >&7+[%,# R"# @# a"# Q300&%C)'"# c3+0# %F)&HC3)'"# C%%C(# D()=# +%EC3&37'LO+70%# 0%F%&)E6%'C"# 8"'#7- E4""&/)-M5'3'=*-$&,#NK:LNKJ#.9::W1" D%)37-!#%;7-:#57-K:!#$$)#.N1,#KJ:LKJN#.9:!Z1" NN"#c7+)',#Q"#5",#`)+67',#<"#c"#@#c7++%CC,#-"#Q"# 9J"#5+%&&%CLT3''%+,#5"#@#S"#Q"#>(37EE%"#<3')D7H+# A# '%=# (8E)C(%D3D# )* # 03')D7H+# +%&7C3)'D(3ED# 7'0# %OOD#7'0#'%DC3'OV#^6E&347C3)'D#*)+#H'0%+DC7'03'O# %7+&8# 03')D7H+# %F)&HC3)'"# D%)4"&# %)!,# K:!LK:W# C(%#)+3O3'#)* #I3+0D"#./#-"#R"#>H++3%,#/"#c"#a)EE%&(HD,# .9:!Z1" Q"#A"#U(HO7+#@#R"#S#$+3O(C#./0D"1#N&%)$&"&;-O"%='/?,# !MKL9!J#.^'037'7#l'3F"#-+%DD,#lUA,#9::J1" NJ"# Y)+'%+,# R"# @# ?"# Q7[%&7"# `%DC# )* # RHF%'3&%D# E+)F30%D# %F30%'4%# )* # *763&8# DC+H4CH+%# 76)'O# 9K"#h3LaH3,#h"#<3')D7H+D#%OOD#3'#>(3'7V#B'#C(%# 03')D7H+D"#D%)4"&##'#,#9;WLM#.!;Z;1" DC+H4CH+%#7'0#%F)&HC3)'#)* #%OOD(%&&D"#./#>7+E%'C%+,# a,#Y3+D4(,#a"#2"#@#R"#?"#Y)+'%+#./0D"1,#O5/'?%4"- NK"#Y)+'%+,#R"#?"#/OO#4&HC4(%D#7'0#%6I+8)D#)* #C=)# 1==?- %/;- M%H5&?,# !MJL9:N# .>76I+30O%# l'3F%+D3C8# (70+)D7H+37'#03')D7H+D"-67-A&")7-8%3&'"#$",#W:ZLW!!# -+%DD,#lUA,#!;;J1" .!;;;1"
9W"#>7+E%'C%+,#a",#Y3+D4(,#a"#2"#@#R"#?"#Y)+'%+"# NW"#2%0H4437,#A"#./0"1#@5;;3&-'0 -)$&-0&%)$&"&;-;"%='/?# UH667+8# 7'0# E+)DE%4CHD"# ./# >7+E%'C%+,# a,# .e7&%#l'3F%+D3C8#-+%DD,#lUA,#9:!91" Y3+D4(,#a"#2"#@#R"#?"#Y)+'%+#./0D"1,#O5/'?%4"-1==?- %/;-M%H5&?,#NWWLNZ:#.>76I+30O%#l'3F%+D3C8#-+%DD,# NZ"#h%&%'3CD[8,#<"a",#Y)+'%+,#R"?"#@#2"#T(%++3%'"# lUA,#!;;J1" <3')D7H+#%OOD"#./#c+%CCLUH+67',#Q"#a",#Y)&CX,#T"# ?"#@#R"#B"#27+&)=#./0D"1#,$&-E'F(3&)&-O5/'?%4",#W!NL 9Z"# Y%0O%D,# ?"# /"# Q"# c)'%# <37O%'%D3DV# A'# W9:"#.^'037'7#l'3F"#-+%DD,#lUA,#9:!91" BF%+F3%=#)* #-+)4%DD%D"#!"#$%&'F&)"*#))V#N!;_N9M# .9::91" NM"#c)'7E7+C%,#R"#2"#@#]3'4%,#Q"#/&#(7&&7XO)#0%&# E+36%+#'30)#0%#<3')D7H+3)D#C+3nD34)D#.U7H+3D4(37,# 9M"# `8D,# e",# 57HC+)',# R",# 57+437L?H3X,# R"# Q"# @# -+)D7H+)E)071,# T+3nD34)# UHE%+3)+# 0%# -7C7O)'37,# Y3'4[%,# Q"# T"# AF37'# %OOD(%&&# 63'%+7&3X7C3)'V# A+O%'C3'7"#!F&=$5/5%/%#$.91,#!ZNLM9#.!;Z;1" I3)4(%6347&# 7'0# *H'4C3)'7&# 4(7+74C%+3X7C3)'# )* # 67C+3m#E+)C%3'D"-E7-@7-8%3&2'37#!,#KJ;LW9#.9::J1" The Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage · 547
N;"#a3C4(3'O,#R"#$"#-+%&363'7+8#+%E)+C#)'#7#4&HC4(# JM"# UC+7%&%',# ]"#F7'"# T(%# 634+)DC+H4CH+%# )* # C(%# )* #D3m#<3')D7H+37'#%OOD#*+)6#C(%#lEE%+#T+37DD34# 03')D7H+37'#%OOLD(%&&D#*+)6#C(%#>+%C74%)HD#I%0D# /&&3)C# 2)+67C3)',# `)+C(%+'# B+7'O%# 2+%%# UC7C%"# )* #Q)'O)&37"#!F&"5#%/-J4?&4F-D'25)%)&?#$*!,#!LJ# 8%3&'/)'="%($5#%-!0"5#%/%-##,#J!LK#.!;Z;1" .!;9K1"
J:"#?%3DX,#?"#?",#/F7'D,#<"#>",#?)I%+CD,#/"#Q",#UH%D,# J;"#-+3C4(7+0,#-"#>"#Y"#./0"1#1/#*#3'(&;5%-'0 -,4")3&?# Y"L<"#@#A"#Q"#e7C%D"#B&0%DC#[')='#03')D7H+37'# .T"#2"#Y"#-HI&347C3)'D,#`%ECH'%#>3C8,#`%=#R%+D%8,# '%DC3'O#D3C%#7'0#+%E+)0H4C3F%#I3)&)O8#)* #C(%#/7+&8# lUA,#!;Z;1"# RH+7DD34# D7H+)E)0)6)+E(# J%??'?('/;*34?"# 8"'#7- D%)37-!#%;7-:#57-K:!#$(",#9J9ML9JNN#.9:!91" K:"# >))6ID,# $"# RHF%'3&%# 4%+7C)ED37'D# *+)6# Q)'O)&37# _# C(%# D67&&%DC# [')='# 03')D7H+# J!"#Q7C%HD,#^"#Y"#&)-%37#>)HFo%,#)H%*D#%C#%6I+8)'D# DE%436%'D"#D%)4"&##'!,#NM:LNM!#.!;M:1" 0p'#03')D7H+%#C(o+)E)0%#0H#RH+7DD3qH%#DHEo+3%H+# 0%# S)H+3'(P# .-)+CHO7&1"- 9&?# #'F()&?- "&;4?- ;&- K!"# >))6ID,# $"# RHF%'3&%# DE%436%'D# )* # C(%# 3C!#%;&F5&-;&?-?#5&/#&?L-1%")$-%/;-83%/&)%"*-:#5&/#&#!#%,# )+'3C(3D4(37'#03')D7H+#-D3CC74)D7H+HD"#8%3%&'/)'3'=*- Z!LZM#.!;;Z1" #%,#M;L!:Z#.!;M91"
J9"# Y3+D4(,# a"# 2"# RH+7DD34# %OOD(%&&# *+)6# C(%# K9"# c+3CC,# c"# c"# @# c"# 5"# `78&)+"# A'# %6I+8)'34# $%DC%+'#^'C%+3)+"#./#>7+E%'C%+,#a,#Y3+D4(,#a"#2"#@# E%F%"%?%4"4?# .<3')D7H+37,# U7H+)E)071# *+)6# C(%# R"#?"#Y)+'%+#./0D"1,#O5/'?%4"-1==?-%/;-M%H5&?,#!NZL lEE%+#RH+7DD34#Q)++3DD)'#2)+67C3)'#.<+8#Q%D7# !K:#.>76I+30O%#l'3F%+D3C8#-+%DD,#lUA,#!;;J1" jH7++8,# >)&)+70)1"# ./# >7+E%'C%+,# a,# Y3+D4(,# a"# 2"#@#R"#?"#Y)+'%+#./0D"1"#O5/'?%4"-1==?-%/;-M%H5&?,# JN"#`)+%&&,#Q"#A",#>&7+[,#R"#Q",#>(37EE%,#S"#Q"#@#<"# 9KWL9WJ#.>76I+30O%#l'3F%+D3C8#-+%DD,#lUA,#!;;J1" <7D(X%F%O"#A#'%DC3'O#03')D7H+"#D%)4"&-!*',#ZZJ_# ZZW#.!;;K1" KN"#>7+E%'C%+,#a"#@#R"#Q4^'C)D("#lEE%+#RH+7DD34# I7I3%D# *+)6# C(%# Q)++3DD)'# 2)+67C3)'"# ./# JJ"#<)'O,#hLQ"#@#-"#R"#>H++3%"#B'#C(%#03D4)F%+8#)* # >7+E%'C%+,#a,#Y3+D4(,#a"#2"#@#R"#?"#Y)+'%+#./0D"1"# 7'#)F3+7EC)+30#D[%&%C)'#)'#7#'%DC#)* #%OOD#7C#c787'# O5/'?%4"- 1==?- %/;- M%H5&?,# 9KWL9WJ# .>76I+30O%# Q7'07(H,#^''%+#Q)'O)&37,#-%)E&%\D#?%EHI&34#)* # l'3F%+D3C8#-+%DD,#lUA,#!;;J1" >(3'7"#E%/%;5%/-67-1%")$-:#5"#!!,#WN!LWW#.!;;W1" KJ"# c7[%+,# c"# T"# <3')D7H+# (%+%D8# _# 03')D7H+# JK"#Q)+7C7&&7,#R"#R"#@#R"#/"#-)=%&&"#<3')D7H+#'%DC3'O# +%'73DD7'4%"#./#?"#<"#a"#T()67D#@#/"#>"#B&D)'# E7CC%+'D"# ./# >7+E%'C%+,# a,# Y3+D4(,# a"# 2"#@# R"# ?"# ./0D1"# A# E'3;- 9''G- %)- )$&- Q%"FSM3'';&;- O5/'?%4"?# Y)+'%+# ./0D"1,# O5/'?%4"- 1==?- %/;- M%H5&?,# NZLJW# .$%DCF3%=#-+%DD,#c)H&0%+,#>)&)+70),#lUA,#!;M:1" .>76I+30O%#l'3F%+D3C8#-+%DD,#lUA,#!;;J1" KK"#R%'D%',#R"#A"#@#a"#-7037'"#U67&&#EC%+)D7H+D#7'0# JW"# U)4(7F7,# A"# <3')D7H+# %OOD# *+)6# C(%# lEE%+# 03')D7H+D#*+)6#C(%#l'4)6E7(O+%#*7H'7#.c+HD(8# >+%C74%)HD#)* #C(%#5)I3#<%D%+C"#8%3&'/)'37-67-),#K!ZL c7D3'#Q%6I%+,#Q)++3D)'#2)+67C3)'V#iT3C()'37'1,# K9Z#.!;W;1" S7C%# RH+7DD34,# =%DC%+'# >)&)+70)"# 67- 8%3&'/)'3"# &!,# NWJLNZJ#.!;M;1" JZ"# U)4(7F7,# A"# T=)# C8E%D# )* # %OO# D(%&&D# 3'# >%')67'37'# 03')D7H+D"# 8%3&'/)'37- 67# !,# NKNLNW!# KW"#j37'O#R3,#&)-%3"#-C%+)D7H+#%OO#=3C(#7#&%7C(%+8# .!;Z!1" D(%&&"#D%)4"&#)!#,#KZ9#.9::J1" 548 · The Rediscovery of Archaeopteryx · Non-rigid eggs in the bird lineage
KZ"#d37)&3'#$"#@#h"#h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a3''%8,#Q"# S"# @# a"# R"#Q4`767+7"# ./0D"1# K;"# dH,# d"# h()H,# h"# @# d"# $7'O,# T(%# D67&&%DC# V&)&"'#$"'/*B- ,$&- 12'34)5'/- '0 - >/)'=&/*- .UE+3'O%+,# [')='#')'L7F37'#C(%+)E)0#03')D7H+"#D%)4"&#)(',# c)DC)',#Q7DD74(HD%CCD,#lUA,#!;;!1" Z:KLZ:M#.9:::1" WM"# ?%3&&8,# U"# Q"# A'# 3'C%O+7C3F%# 7EE+)74(# C)# W:"#YH,#<",#Y)H,#S",#h(7'O,#S"#@#d"#dH"#A#E+%L 1).)%,(1%,2@X".1)"-0#.02(.0,2"5)./))2"02.)%#+)(07(" !"#$%&'()&"*+# C+))0)'C30# C(%+)E)0# *+)6# >(3'7# '2-"02.%'#+)(07("+1)2,4)2'A"M5'37-67-95//&%/-:'#7# =3C(#&)'O#*%7C(%+D#)'#C(%#6%C7C7+DHD"#D%)4"&#)&$,# &(,#!!;L!JN#.!;;Z1" WJ:LWJN#.9::;1" W;"#a&3'O%'I%+O,#>"#-"#Y%C%+)4(+)'8#7'0#7&&)6%C+8V# W!"#5)0%*+)3C,#-,#>7H,#A",#<)'OLeH,#Y",#/D4H3&&3o,#2"# C(%#7'7&8D3D#)* #%F)&HC3)'7+8#4(7'O%#3'#)'C)O%'8"# $%'(7),#$"#@#5"#<8[%"#A#RH+7DD34#7F37&7'#03')D7H+# M5'37-@&27#*!,#Z;L!9N#.!;;M1" *+)6#>(3'7#+%D)&F%D#C(%#%7+&8#E(8&)O%'%C34#(3DC)+8# )* #I3+0D"#D%)4"&#)"',#NK;LNW9#.9:!N1" Z:"# U63C(,# a"# a"# Y%C%+)4(+)'8# +%F3D3C%0V# C(%# %F)&HC3)'# )* # 0%F%&)E6%'C7&# D%qH%'4%D"# M5'37- 67- W9"# Y)H,# S"LY",# h()H,# h"LY",# Q7+C3',# S"# <"# @# 95//&%/-:'#7#*!,#!W;L!MW#.9::!1" A"#2%0H4437,#A#I%7[%0#I3+0#*+)6#C(%#RH+7DD34#)* # >(3'7"#D%)4"&#!**,#W!WLW!M#.!;;K1" Z!"#Y)EE,#T"#-"#@#Q"#R"#B+D%'"#<3')D7H+#I+))03'O# I%(7F3)+#7'0#C(%#)+3O3'#)* #*%7C(%+D"#./#>H++3%,#-"# WN"#5+%&&%CLT3''%+,#5",#S"#>(37EE%,#Q"#`)+%&'0# R",#a)EE%&(HD,#/"#c",#U(HO7+,#Q"#A"#@#R"#S"#$+3O(C# <"#c)CCk%+"#<3')D7H+#%OOD#7'0#'%DC3'O#I%(7F3)+DV# ./0D"1,# N&%)$&"&;- O"%='/?,# 9NJ# _# 9K:# .^'037'7# A# E7&%)I3)&)O347&# 3'F%DC3O7C3)'"# 8%3%&'=&'="%($*L- l'3F%+D3C8#-+%DD,#9::J1" 8%3%&'#35F%)'3'=*L-8%3%&'&#'3'=*##!#,#9;JLN9!#.9::W1"
WJ"# 2)C(,# >"# @# B"# $"#Q"# ?7H(HC"# ?%L%F7&H7C3)'# )* #C(%#Y77+&%6#!"#$%&'()&"*+#7'0#C(%#+7037C3)'#)* # 67'3+7EC)+7'#C(%+)E)0#03')D7H+D"#MJE-12'37-M5'37- $*,#9NW#.9:!Z1" “What you are claiming only happens once every fifty years. No, it happens once every hundred and fifty years, I responded.” In Preface, The Rediscovery of Archaeopteryx (Volume I)
GROUND BREAKING FINDINGS WENT UNNOTICED FOR OVER 140 YEARS ON THE MONA LISA OF FOSSILS DESPITE ITS DISPLAY BEFORE MILLIONS OF VISITORS AND SCRUTINY BY GENERATIONS OF RESEARCHERS. The Berlin specimen of Archaeopteryx became the icon of the theory of evolution for featuring an animal thought to represent a transitional stage that preceded the appearance of today’s birds. An investigation into the disruptive hypothesis that its fossilized pose corresponds to a nesting posture, rather than being the carcass of an animal that sunk into the bottom of a Jurassic lagoon, is described here and results in the sensational rediscovery of the most famous fossil in the world. In this opening volume of The Rediscovery of Archaeopteryx, non-rigid eggs from bird-related animals are exposed to us all.
The author was the recipient of the Bial Grand Prize in Medicine for developing a new molecular approach to the study of the development of blood cell formation in early embryonic life and having cloned novel genes associated with this process. He also received the Medal of Honor from the Portuguese Business Association for his role as inventor of the ALERT® Electronic Medical Record and founder of ALERT Life Sciences Computing. Jorge Guimarães graduated from the Faculty of Medicine of Porto, where he was an instructor of Physiology; he was also a visiting scientist at the DNAX Research Institute, in Palo Alto, California, where he developed his award-winning work on cloning of differentially expressed genes in embryoid bodies, embryos and cell lines, and a post-doctoral fellow at Stanford University where he worked on gene therapy. His life is now dedicated to investigating the relationship between evolution, reproduction, development and cancer.