bioRxiv preprint doi: https://doi.org/10.1101/2021.04.19.440491; this version posted June 21, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. A FOSSIL FISH ASSEMBLAGE FROM THE MIDDLE MIOCENE OF THE COCINETAS BASIN, NORTHERN COLOMBIA GUSTAVO A. BALLEN1,2,* ORCID 0000-0001-5424-8608 CARLOS JARAMILLO2,3,4 ORCID 0000-0002-2616-5079 !ERNANDO C. P. DAGOSTA5 ORCID 0000-0001-7163-296# MARIO C. C. DE PINNA1 ORCID 0000-0003-1711-4816 *C$%%&'($)*+), a./0$%, ,-1-22&)304,5-+2.c$5, ,-1-22&)304-2.5)+.usp.br 1M.'&u de Z$$2$,+- da U)+7&%'+*-*& de S8o P-.2$, S8o P-.2$, S", Bra9+2 2S5+/0'$)+-) T%$(+3-2 R&'&-%30 Ins/+/./&, P-)-5:, P-)-5: 3ISEM, U. M$)/(&22+&%, CNRS, E";E, IRD, M$)/(&22+&%, F%-)3& 4D&(-%/5&)/ of G&$2$,=, F-3.2/y of S3+&)3&', U)+7&%'+/y of S-2-5-)3-, S-2-5-)3-, S(-+) 5!-3.2*-*& de Ci>)3+-' Bi$2?,+3-' e A51+&)/-+', U)+7&%'+*-*& F&*&%-2 da G%-)*& D$.%-*$', D$.%-*$', MS, Bra9+2 Abstract: F%&'0@-/&% f$''+2 fi'h f-.)-' ha7& be&) l$)g us&d to i)<&% pa'/ d%-+)-,& 3$))&3/+$)', a' t0&= a%& bounded by ph='+3-2 fre'0@-/&% ba%%+&%'. H&%& w& s/.*y a 5+**2& M+$3&)& (15.0-–15.5 M-) fre'0@-/&% fi'h f$''+2 f-.)- (M-E-%-+(-$D fr$5 t0& C-'/+22&/&' F$%5-/+$) in nor/0&%n C$2$51+-, no@-*-=' w&'/ of t0& A)*&'. W& r&3$%d /0& pre'&)3& of l.),A'0&' (Lepidosiren), pa3.' (Mylossoma and Piaractus), a%5$%&* 3-/A'0&' (C-22+30/0=+*-&), a)* r&d-t-+2 c-/A'0&' (Phractocephalus). EG/-)/ m&51&%' of a22 t0$'& g%$.(' (eG3&(/ t0& C-22+30/0=+*-&, due to l-3E of t-G$)$5+3 r&'$2./+on) a%& <ound i) A5-9$)+-) f-.)-' e-'/ of t0& A)*&' and a%& a1'&)/ fr$5 f-.)-' w&'/ of t0& 1 bioRxiv preprint doi: https://doi.org/10.1101/2021.04.19.440491; this version posted June 21, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. A)*&', i)*+3-/+), t0-/ t0& r+7&%+)& s='/&5' of t0& G.-H+%- P&)+)'.2- w&%& c$))&3/&d t$ A5-9$)+- dur+)g t0& m+**2& M+$3&)&. T0& s+5+2-%+/y of L- V&)/- (w&'/ of t0& A)*&'D -nd Rio A3%& (e-'/ of t0& A)*&') fi'h f-.)-' dur+)g t0& l-/& M+$3&)& f.%/0&r i)*+3-/&' /0-/ t0& nor/0&%n A)*&-n upl+</ w-' no/ a c$5(2&/& ba%%+&r a/ l&-'/ unt+2 ∼ 11 Myr a,$. ;$@&7&%, t0&%& i' a c$)/+)&)/-2-@+*& s/%.3/.%+), of t0& M+$3&)& fi'h f-.)-' t0-/ i' a2'$ <ound i) t0& eG/-)/ f-.)-', s.,,&'/+)g t0-/ o/0&% f-3/$%' s.3h a' e3$2$,+3-2 c$)*+/+$)', +n a**+/+on t$ t0& upl+</ of t0& A)*&', ha7& s0-(&d t0& bi$,&$,%-(0+3 e7$2./+$) of S$./h A5&%+3-n fi'h f-.)-'. Keywords: M+$3&)&, C&)$9$+3, C$3+)&/-', S$./h A5&%+3-, P-2&$+30/0=$2$,=. INTROD CTION !%&'0@-/&% fi'0&' a%& t0& r+30&'/ v&%/&1%-/& c$)/+)&)/-2 c$5($)&)/ of t0& N&$/%$(+3', @+/0 ∼ 7,000 s(&3+&' (A21&%/ & R&+' 2011). T0&y a%& h+,02= d+7&%'&, ra),+)g fr$5 1%-3E+'h w-/&%' to e2&7-/+$)' a1$7& 3000 m in t0& A)*&' (S30-&<&% 2011). D&'(+/& t0+' &)$%5$.' di7&%'+/=, fr&'0@-/&% fi'0&' a%& no/ a' pr$5+)&)/ i) t0& f$''+2 r&3$%d of S$./0 A5&%+3- a' m-55-2' or c%$3$*=2+-)', a)* t0&y a%& of/&) r&3$%*&* fr$5 bone <%-,5&)/' of l+5+/&* di-,)$'/+3 v-2.& (L.)*1&%, et al. 2010). T0& pa.3+/= of w+*&- %-),+)g c$5(-%-/+7& m$%(0$2$,+3-2 a)-2='&' f$3.'&d on di-,)$'/+3 c0-%-3/&%' (%&'&%7&d i) f$''+2 fi'h s(&3+5&)' l+5+/' t0& i*&)/+A3-/+$) to only c$-%'& t-G$)$5+3 2&7&2' s.30 a' o%*&% or f-5+2=. S.30 d%-@1-3E' l+5+/ po/&)/+-2 us& of N&$/%$(+3-2 <%&'0@-/&r f$''+2 fi'0&' i) pa2&$&3$2$,+3-2, s='/&5-/+3, bi$'/%-/+,%-(0+3, and 1+$,&$,%-(0+3 s/.*+&'. !%&'0@-/&% fi'0&' ha7& pr$7+*&* e7+*&)3& of a)3+&)/ dra+)-,& c$))&3/+$)' be/@&&n %+7&r s='/&5' e-'/ a)* w&'/ of t0& A)*&', a' t0& di'(&%'-2 po/&)/+-2 of fr&'0@-/&r fi'0&' -3%$'' m$.)/-+n ra),&' i' l+5+/&d for l$@2-nd t-G-, t0.' he2(+)g t$ c$)'/%-+n t0& &7$2./+on of t0& A)*&-n or$,&)y (L.)*1&%, 1997; L.)*1&%, et al. 2010). For &G-5(2&, eG/-)/ fr&'0@-/&r fi'h t-G- c.%%&)/2= r&'/%+3/&* to c+'-A)*&-n d%-+)-,&' (e.,., 2 bioRxiv preprint doi: https://doi.org/10.1101/2021.04.19.440491; this version posted June 21, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. O%+)$3o-A5-9on) a%& found in M+$3&)& t%-)'-A)*&-n s+/&' (+.e., w&'/ of m$*&%) A)*&' r-),& a' L- V&)/-, Ca'/+22&/&' and U%.5-3o), i)*+3-/+), a)3+&)/ h=*%$2$,+3-2 3$))&3/+7+/y t0-/ i' s&7&%&d no@-*-=' (D+-9 de G-5&%o 1996; G%&,$%=-F$*9+3E+ 2000; M$)/&' et al. 2021). A num1&r of s/.*+&' ha7& i5(%$7&d our unde%'/-)*+), of t0& g&$2$,y a)* (-2&$)/$2$,y of t0& C$3+)&/-' s&*+5&)/-%y ba'+) in t0& G.-H+%- P&)+)'.2- of no%/0&%) C$2$51+- (M$%&no et al. 2015). S&7&%-2 v&%/&1%-/& g%$.(' a%& r&(%&'&)/&d i) t0&'& '&*+5&)/', bo/h m-%+)& and c$)/+)&)/-2 (A,.+2&%- et al. 2013aJ F$%-'+&(+ et al. 2014; C-*&)- & J-%-5+22o 2015a,bJ A5'on et al. 2016; M$%&no-Be%)-2 et al. 2016; S.-%&9 et al. 2016; A,.+2&%- et al. 2017; C-%%+22o-Br+3&K$ et al. 2019); in c$)/%-'/, fr&'0@-/&r A'0&' r&5-+)' l-%,&2= unr&($%/&* in t0&'& a''&512-,&' (but s&& A,.+2&%- et al. 2013b). ;&%&+n, w& r&($%/ t0& fr&'0@-/&r f$''+2 fi'0&' of t0& M+$3&)& Ca'/+22&/&' F$%5-/+on i) /0& M-E-%-+(-o l$3-2+/=, nor/0&%n C$2$51+-, and t0&+r r&2&7-)/ a)-/$5+3-2 c0-%-3/&%'. F& c$5(-%& t0& f$''+2 a''&512-,& w+/h S$./0 A5&%+3-) fi'h f-.)-' a' w&22 a' w+/h /0& eG/-)/ c$)/+)&)/-2 fi'h d+7&%'+/y i) o%*&% to unde%'/-nd t0& hi'/$%+3-2 c$5($)&)/ of '+5+2-%+/y pa//&%)' in t+5& and s(-3&. MATERIALS AND METHODS Geological setting T0& C$3+)&/-' s&*+5&)/-%y ba'+) pr&'&%7&' a s.33&''+on of c$)/+)&)/-2 t$ s0-22$@ 5-%+)& s/%-/- on t0& nor/0&%n m-%,+n of t0& S$./h A5&%+3-) P2-/&. M$%&no et al. (2015) r&*&A)&* t0& s/%-/+,%-(0y of t0& ba'+n a)* s0$@&d t0-/ t0& C-'/+22&/&' !$%5-/+on i' c$5($'&d of s&*+5&)/' of M+$3&)& a,& fr$5 m$'/2= s0-22$@ m-%+)& &)7+%$)5&)/'. It' l$@&r bounda%= i' c$)3$%*-)/ w+/h t0& J+5$2 F$%5-/+on w0&%&-' i/' .((&r bounda%= i' s&/ by an unc$)<$%5+/y w+/0 t0& W-%& F$%5-/+on. T0& C-'/+22&/&' 3 bioRxiv preprint doi: https://doi.org/10.1101/2021.04.19.440491; this version posted June 21, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder, who has granted bioRxiv a license to display the preprint in perpetuity. It is made available under aCC-BY-NC 4.0 International license. !$%5-/+on ha' be&) da/&* a' l$@&r to m+**2& M+$3&)& us+)g Sr g&$30%$)$2$,y (H&)*= et al. 2015). M-E-%-+(-$ (l$3-2+/y STRI 390093, 11º54’32.0“N 71º20’24.4”F) i' a m$*&%-/& (2-/&-. l=+)g to t0& w&'/ of t0& T.3-3-' ba=, m.)+3+(-2+/y of U%+1+-, G.-H+%- *&(-%/5&)/, no%/0&%n C$2$51+- (F+,.%& 1--c). T0& l$3-2+/y i' r&3$%*&* in t0& '/%-/+,%-(0+3 s&3/+$) c-22&* “L$)g s&3/+$)O (M$%&)$ et al. 2015). T0& l$3-2+/y l+&' -1$./ 127 s/%-/+,%-(0+3 m&/&%' a1$7& t0& ba'& of t0& s&3/+$) (F+,.%& 1*) and ∼ 279 m -1$7& t0& ba'& of t0& C-'/+22&/&' f$%5-/+on (S.-%&9 et al. 2016). S(&3+5&)' w&%& 3$22&3/&d fr$5 a s-)*'/$)& and c$),2$5&%-/+3 s-)*'/$)& l&7&2. T0& Sr i'$/$(+3 da/- 3$)'/%-+)' t0& M-E-%-+(-o l$3-2+/y t$ a 15.0–-15.5 M- a,& (C-%%+22o-Br+3&K$ et al.