The complex case of Macaronichnus trace fossil affecting rock porosity

1 Vol.:(0123456789) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s The complex case o Maca onichnus ace ossil a ec ing ock po osi y Ja ie Do ado 1*, F ancisco J. Rod íguez‑To a 2 & Olmo Miguez‑Salas2 Bio u ba ion is an impo an ac o o ese oi quali y due o he modi ica ion o hos ock pe ophysical p ope ies (i.e., po osi y, pe meabili y, and connec i i y). Howe e , he e is no p edic able ela ionship be ween bio u ba ion and i s e ec on ock p ope ies, due o he a iabili y o he in ol ed ichnological ea u es. A de ailed ichnological analysis is necessa y o de e mine how bio u ba ion a ec s pe ophysical p ope ies in a bio u ba ed ese oi . T adi ionally, ichnological ea u es such as densi y, ie ing, size, o ien a ion, a chi ec u e, and ill, ha e been conside ed. Howe e , o he p ope ies ha e been unde alued as is he case o lining. He e, we p esen a de ailed s udy on he e ec s o Maca onichnus bu ows, an ichno axon usually ela ed o hyd oca bon explo a ion due o i s high concen a ion in ock no ably a ec ing pe ophysical p ope ies. Maca onichnus, a subho izon al cylind ical bu ow, is cha ac e ized by a well‑de ined and de eloped ou e im su ounding he ube co e. Ou da a indica es a clea zona ion in po osi y acco ding o bu ow s uc u e, wi h he lowes po osi y in he ube co e and highe alues associa ed wi h he su ounded im. This duali y is de e mined by he acemake g ain selec i e eeding ac i i y and he consequen concen a ed cemen a ion. The o ganism concen a es he ligh e mine als in he ube co e ill du ing eeding, a o ing pos ‑deposi ional cemen a ion du ing diagenesis and his esul s in lowe po osi y han he hos ock. Howe e , hea y mine als, mainly glauconi e, a e loca ed in he im, showing highe po osi y. Ou esul s suppo he iew ha ichnological analyses a e essen ial o de e mine ese oi quali y in bio u ba ed ese oi s, e idencing ha o he ichnological p ope ies in addi ion o hose adi ionally conside ed mus be e alua ed. Bio u ba ion is an impo an ac o o ese oi quali y es ima ion in he oil and gas indus y. Biogenic ac i i ies may a ec some pe ophysical p ope ies such as po osi y, pe meabili y, and connec i i y1–3. This in luence on ock p ope ies can be ei he posi i e o nega i e (i.e., ei he inc easing o dec easing po osi y, espec i ely, in compa ison o he hos ock). De ailed ichnological analyses should be conduc ed in e e y bio u ba ed ese oi o quan i y i s e ec and es ima e he ese oi quali y2,4. T adi ionally, a en ion has been paid o pa icula cha ac e is ics o bu ows impac ing ock p ope ies and ese oi quali y such as densi y, ie ing, size, o ien a- ion, a chi ec u e, ill, and lining2,5. Wi h espec o lining, analyses ha e ocused on i s p esence o absence6,7, and a ely on he pe ophysical p ope ies8,9. Howe e , hickness and composi ion o he lining/man le can be a majo ea u e o be conside ed o some bu ows, as is he case o Maca onichnus10. Maca onichnus has been commonly associa ed wi h ese oi exploi a ion and hen i s e ec has been ana- lyzed in de ail o some deposi s1,9,11–13. Thus, Ging as e al.11 b ie ly poin ed ou ha po osi y dis ibu ion in sands ones domina ed by Maca onichnus is complex and may a ec he ese oi quali y, bu hey sugges mo e esea ch had o be done. La e , Go don e al.13 analyzed he p esence o Maca onichnus in he Bluesky Fo ma ion (Albe a, Canada) o de e mine ha pe meabili y was enhanced in hose zones whe e Maca onichnus we e abun- dan compa ed o simila non-bio u ba ed in e als. Then, G eene e al.8 poin ed o di e ences be ween lining and illing, and Quaye e al.9 ound ha po osi y and pe meabili y in he bu ow ill o Maca onichnus we e highe han in he hos sedimen in he Funing Fo ma ion (Subei basin, China). These s udies ha e shown he e ec o Maca onichnus and i s posi i e impac in spi e o he gene al idea ha bio u ba ion educes pe meabili y4. Mo eo e , Maca onichnus usually occu s in dense concen a ions, commonly occupying mo e han 70% o he ock4, and i jus akes e y li le ime o p oduce s (i.e., polychae es) o do i 14. A ha concen a ion, i is an impo an biogenic s uc u e a ec ing pe ophysical p ope ies, luid low, and, consequen ly, oil p oduc ion. Fo all hese easons, we decided o analyze he e ec o Maca onichnus in mo e de ail. Maca onichnus seg ega is15 can be ound om Pe mian o Holocene2, mos ly in shallow o esho e and shal- low sub idal deposi s, bu i has been also obse ed in deep-sea sedimen s such as shel o slope deposi s16–18. Maca onichnus is usually iden i ied as non-b anching, cylind ical, and sub-ho izon al bu ows om jus a ew OPEN 1Depa men o Ea h Sciences, Royal Holloway Uni e si y o London, Egham TW20 0EX, UK. 2Depa amen o de Es a ig a ía y Paleon ología, Uni e sidad de G anada, 18002 G anada, Spain. *email: Ja ie [email p o ec ed] 2 Vol:.(1234567890) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s/ millime e s up o 15mm diame e , cha ac e ized by a clea mine alogical seg ega ion be ween he cylinde ube co e and he su ounding im15. Rim composi ion shows a concen a ion o da ke and hea y mine als e-so ed by he p oduce du ing eeding, while ligh e mine als commonly cha ac e ize he ube co e2,11. This im can be ela i ely hick, especially in he subichnospecies Maca onichnus seg ega is degibe i16,17. Howe e , he incidence o he im on pe ophysical p ope ies has a ely been conside ed sepa a ely bu as pa o he en i e in illing ma e ial ( ube co e and im)9. He e, we analyze in de ail he composi ion o in illing ma e ial om Maca onichnus seg ega is degibe i di e en ia ing be ween ube co e and im, ocusing on i s espec i e incidence on po osi y and hen on hei pa icula impac in ese oi quali y. This will help us o unde s and be e he ela i e e ec o e e y single pa in po osi y, which is essen ial o p o ide an accu a e es ima ion o po osi y in bio u ba ed ese oi s. Fo his s udy we in es iga e ock samples including specimens o Maca onichnus seg ega is degibe i ha we e collec ed om wo ou c ops in no he n Mo occo: El Ade gha (34°4′34.19″N, 4°51ʹ33.43″W) and Ki - m a (34°10ʹ15.07″N, 5°14ʹ21.43″W)18–20 (Fig.1A). These ou c ops a e in e p e ed as clas ic con ou i e sands ones associa ed o he paleo-Medi e anean Ou low Wa e and he Ri ian co ido ha was connec ing he A lan ic Ocean and he Medi e anean Sea du ing La e Miocene18–20. The ou c ops belong o he Sou h Ri ian co ido (Saiss Basin) and a e La e Miocene in age19–23. These loca ions we e selec ed as sampling allows us o e alua e nice examples whe e Maca onichnus specimens a e clea ly iden i ied wi h di e en deg ees o bio u ba ion and o e lapping be ween bu ows (Fig.1B). Resul s Pe og aphic analyses e eal ha s udied clas ic con ou i es, de ined as sands ones in he ield, ha e simila p o- po ions o clas ic and ca bona e componen s (Table1). S ic ly, hey a y be ween bioclas ic limes ones and li hic sands ones, bo h wi h abundan glauconi e. Ca bona e clas s consis o bioclas s and some ewo ked ca bona e ock agmen s. Bioclas s a e mainly bi al e agmen s oge he wi h o he agmen s o o amini e a, echinoids, calca eous algae, and a ely b yozoa. Mos bioclas s a e ounded and clea ly ewo ked. Qua z g ains a e abun- dan , eldspa and mica almos absen . Pelle s o glauconi e a e abundan and ecognized in all he samples; hei Figu e1. (A) Loca ion map o he samples. (B) Ou c ops and ock samples con aining Maca onichnus wi h di e en bio u ba ion in ensi y and o e lapping be ween hem. Whi e a ows poin o Maca onichnus in ock samples. BS Bio u ba ed Su ace. Scale ba s 1cm. Table 1. Mean alues o composi ion and po osi y om analyzed samples conside ing he whole sample (WS), ube co e (TC), im (R) and hos ock (HR). T . T aces, Feld. Feldspa , Glauc. Glauconi e, Ca b. Ca bona es, R.F. Rock agmen s, A gill. A gilliceous, Biocl. Bioclas s, Dolom. Dolomi e, Opaq. Opaques, Po . Po osi y. Qua z Feld Mica Glauc Ca b. R.F O he R.F A gill. ma ix Hea y min Fo ams O he biocl Calci e cemen Dolom I on ox./ hyd ox Opaq Po (%) El Ade gha WS 20.6 0.6 0.0 20.5 15.8 0.6 0.7 0.0 5.0 10.9 21.0 0.2 4.0 0.0 1.5 TC 21.2 0.0 0.0 5.2 22.6 0.0 1.7 0.0 9.7 8.0 28.5 0.0 3.1 0.0 1.3 R – – – – – – – – – – – – – – 3.0 HR 19.0 0.7 0.0 18.3 15.8 0.7 1.1 0.0 6.5 9.7 23.7 0.7 3.6 0.4 1.5 Ki m a WS 15.1 0.0 0.0 22.5 20.6 0.0 1.1 0.0 3.4 15.6 9.6 0.2 11.9 0.2 1.6 TC 23.0 0.0 0.0 2.7 18.7 0.7 0.0 0.0 5.7 12.3 26.0 1.3 9.3 0.3 0.0 R 22.6 0.0 0.0 29.3 6.6 0.0 1.0 T . 1.7 18.1 13.2 0.0 7.0 0.3 5.3 HR 17.5 0.0 0.0 18.9 21.6 0.0 0.0 0.0 2.1 13.4 16.5 0.3 9.3 0.3 2.4 3 Vol.:(0123456789) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s/ dis ibu ion is no andom and is de e mined by bu ow con igu a ion (Fig.2, Table1). Glauconi e is mainly concen a ed in he im o Maca onichnus bu ows; meanwhile he ube co e is mos ly composed on ca bona e componen s (i.e., ca bona e ock agmen s and calci e cemen ) and qua z (Fig.2, Table1). Visually, bu ow ims a e mos ly de ined by a ma ked concen a ion o glauconi e pelle s. Elonga e clas s lie pa allel o he ma gin. When compa ing he da a o po osi y om he whole sample (ma ix and bu ows) and he hos ock (i.e., he ma ix), which ep esen he p ima y po osi y (Table1), alues a e sligh ly a ec ed (2.4% dec eases o 1.6% in Fig.2H) o e en emain unal e ed (1.5% in El Ade gha; Fig.2D). Howe e , po osi y is no homogeneously dis ibu ed and i is clea ly de e mined by zona ion o he bu ows. Tube co es show lowe alues han hos sedimen in El Ade gha (1.3% s 3.0%), and hey a e being e en comple ely non-po ous in he case o Ki m a (0.0% s 5.3%). The highes po osi y is loca ed in ims, doubling alues eco ded in he hos sedimen in bo h cases and showing up o 8.8% po osi y in some a eas (Table1, Fig.2). Discussion I is well es ablished ha bio u ba ion may a ec ese oi quali y1–3,11,12,21. P e ious s udies ha e demons a ed ha Maca onichnus is one o he ace ossils ha can play an impo an ole in bio u ba ed ese oi s9,11,13, gene - ally ha ing a posi i e impac . Howe e , he p e ious s udies mos ly compa ed pe ophysical p ope ies om he hos sedimen and he en i e bu ow, wi hou dis inguishing be ween bu ow ube co e and ou e im. Ou esul s demons a e ha he p esence o Maca onichnus seg ega is degibe i ha e, in gene al, a sligh ly nega i e impac on po osi y, educing alues less han 1%. Howe e , going deepe , ou da a show a ele an ac , ha is he ma ked di e ence be ween ube co e and im, no only in e ms o composi ion, as is well-known, bu also in po osi y. Rega ding composi ion, he ube co e is illed wi h ligh mine als ha a e highly cemen ed, and da k mine - als, mainly glauconi e, a e concen a ed in he ims. This composi ional zona ion is a diagnos ic c i e ion o Maca onichnus15 and is de e mined by ace-make s du ing selec i e eeding ac i i y11,15,17. In ega ds o po osi y, a clea zona ion is e ealed. Mean po osi y in he ims is wo imes highe han in he hos sedimen , eaching up o 8.8% po osi y in some a eas o he ock samples; howe e , po osi y is e y low in ube co es. This po osi y zona ion could be caused by o iginal mine alogical he e ogenei y combined wi h he associa ed pos -deposi ional cemen a ion1. Du ing p oduce eeding ac i i y, ligh mine als (mos ly qua z and ca bona es) a e concen a ed in he ube co e and da ke mine als (e.g., glauconi e) a e placed on he su ounded im. This zona ion causes he concen a ion o deposi ional cemen a ion in he ube co e. This concen a ed cemen a ion p oduced du ing diagenesis, gene a es a po osi y zona ion, which plays a de e minan ole in he e ec on pe ophysical p ope ies. Then, ube co es a e cha ac e ized by lowe po osi y alues han he su ounded ims. Ging as e al.11 epo ed ha po osi y dis ibu ion in Maca onichnus bio u ba ed sands one is complex and ac s like a dual po osi y–pe meabili y sys em. Tha means he e would be a p e e en ial low h ough he bu - ows and a seconda y low om su ounding sedimen in o he bu ows. Dual pe meabili y sys ems a e ound in ese oi s whe e pe meabili y con as be ween ma ix and bu ows is high, a ound h ee o de s o magni ude4. Howe e , when pe meabili y con as is lowe , a dual-po osi y sys em is de eloped and he e is no a p e e en ial low h ough he ace ossils4. Based on ou esul s, his can be mo e complex o Maca onichnus as he e is no a single low sys em o he whole bu ow due o he obse ed zona ion. Pe ophysical p ope ies om ube co e and im a e comple ely di e en and hen bo h pa s o he bu ow should be conside ed sepa a ely when e alua ing he e ec on ese oi quali y. Then, luids would p e e en ially low h ough he im and, o iden i y he low sys em, pe meabili y con as be ween hos sedimen and im has o be e alua ed. Maca onichnus bu - ow diame e s ha e been epo ed om jus a ew millime e s11,14,22 o la ge sizes up o 15mm diame e 16–18,23,24. Howe e , mos o he p e ious s udies do no p o ide im hick measu emen s. Sa da and Uddin10 eco ded bu ow ube co es and im measu emen s and no ed ha ube co e diame e s anged om 4 o 19mm in la ge specimens. They epo ed ha he im hickness was a iable along he bu ow, anging om abou 1 o 5mm, being hicke a he bo om pa o he bu ow. This e eals ha he im occupies 25–30% o he olume o he Figu e2. Close-ups om El Ade gha (A–D) and Ki m a (E–H). (A,E) Close-ups om hos ock (HR). (B,F) Tube co es (TC). (C,G) Rims (R). (D,H) Thin sec ions. Pe cen age alues show he po osi y in e e y case; in (C) and (G) his is jus e e ed o he im. In hin sec ions (D) and (H), alues measu ed in ube co es (black), ims ( ed), and hos sedimen (blue). 4 Vol:.(1234567890) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s/ bu ows10. This ac can also be obse ed in some o he illus a ed igu es om p e ious pape s. Fo example, Rod íguez-To a and Agui e16 show la ge specimens o Maca onichnus seg ega is degibe i wi h ~ 15mm diam- e e whe e im was a ound 2–3mm hickness, and Na a and Seike17 illus a ed ha he im can be e en hicke ep esen ing hal he olume o he ac ual bu ow (Fig.3 om Na a and Seike17). The e o e, im hickness mus no be unde es ima ed, e ealing as a key poin o con ol Maca onichnus po osi y e ec . Especially conside ing ha im po osi y can be wo o h ee imes highe han hos sedimen and ep esen an impo an olume po ion o he whole bu ow (up o 50%). Assuming ha Maca onichnus a e equen ly in dense concen a ions, commonly ep esen ing mo e han 70% o bio u ba ion4, we can es ima e ha in some cases bu ow ims can ep esen 35% o he o al olume in a bio u ba ed sample. Acco dingly, o conduc a p ope ese oi quali y es ima ion in Maca onichnus-bea ing ocks, po osi ies om bo h zones ( im and ube co e) and hei ela i e olumes need o be conside ed. Ano he aspec ha needs o be conside ed is he equen bu ow c osscu ing be ween specimens. Se e al heo e ical si ua ions in ol ing Figu e3. Theo e ically modeled po osi y e ec by Maca onichnus. (A) Model scena io o be compa ed wi h 16% o bio u ba ed su ace (BS) and po osi y (ϕ) 2.1%. (B) Si ua ion wi h same BS, bu di e en im/ ube co e a io ( / c), causing an inc ease in ϕ. (C) Scena io wi h same ϕ (2.1%) in spi e o he lowe BS, due o a highe / c. (D) O e lapped bu ows wi h a e y simila BS, bu a dec ease in im su ace due o c osscu ing, educing ϕ. Bold numbe s ep esen po osi y om hos sedimen (blue), im ( ed), and ube co e (black). 5 Vol.:(0123456789) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s/ di e en a iables such as im/ ube co e olume a ios, he deg ee o bio u ba ion and he o e lapping be ween bu ows a e illus a ed in Fig.3, assuming po osi y alues o 2.0%, 0.0% and 6.0% in he hos ock, im, and ube co e, espec i ely. We es ablished a model scena io conside ing an example wi h 16% o bio u ba ed su ace (BS) by isola ed bu ows, wi h a im/ ube co e ( / c) olume a io o 0.77 and po osi y (ϕ) o 2.1% o (Fig.3A). We conside ed ano he scena io whe e bu ows occupy he same su ace bu wi h hicke im ( / c = 2.2), which esul ed in highe po osi y (ϕ = 2.4%) (Fig.3B). Whe eas in a less bio u ba ed example (Fig.3C), we ob ained he same po osi y alue compa ed o he model scena io (ϕ = 2.1%) due o a highe / c olume a io ( / c = 3.5). We simula ed a inal scena io wi h o e lapped bu ows (Fig.3D) whe e he c osscu ing o bu ows esul ed in a sligh ly lowe su aces alue and / c a io which caused a small educ ion o he po osi y alue compa ed o he model scena io (ϕ = 2.0%). These ou scena ios show how he ela i e / c olume a ios and o e lapping o bu ows can a ec he esul ing po osi y alues. Al hough he esul s om hese scena ios did no show sig- ni ican changes in po osi y alues, bu conside ing ha Maca onichnus is commonly ound ep esen ing mo e han 70% o ock olumes4, whe e ims could ep esen up o 50% o he bu ows17 and o e lapped, hey could signi ican ly modi y he esul ing po osi y alues o he hos ock. Conclusions Maca onichnus has been shown o be a ele an ace ossil o ese oi quali y in bio u ba ed deposi s, mainly ela ed o i s occu ences as dense concen a ions. This s udy has demons a ed ha he po osi y e ec by Maca- onichnus is no homogenous in he en i e bu ow and i s dis ibu ion is clea ly associa ed wi h he ace ossil s uc u e. Po osi y in he bu ow ube co e is di e en o ha om he ou e im. P oduce s o Maca onichnus gene a e a g ain eso ing du ing eeding when hey concen a e ca bona e componen s and qua z g ains in he ube co e ill, and hea y and da ke mine als, mainly glauconi e, in he im. Rega ding ese oi quali y, esul s e eal ha Maca onichnus is a complex case and has a dual beha io . Bu ow ims ha e inc eased po osi y wi h espec o he hos sedimen and p o ide posi i e e ec s. Fillings in ube co es ha e lowe po osi y and p o ide nega i e e ec s. On his basis, he ela i e olume occupied by ims and ube co es should be de e mined o es ima e he eal e ec o Maca onichnus on po osi y. Any e alua ion o he impac o Maca onichnus on pe ophysical p ope ies, and hen on ese oi exploi a ion, based on jus he in ensi y o bio u ba ion o he s udy o he whole bu ow mus induce signi ican misin e p e a ions o ele an scien i ic and economic implica ions. This complex dual beha io has been obse ed o he i s ime in Maca onichnus, bu we could no disca d ha his could also be iden i ied in some o he ace ossils wi h ma ked in e nal zona ion. Ma e ials and me hods Twen y- ou ock samples o con ou i e sands ones including specimens o Maca onichnus seg ega is degibe i we e collec ed om wo ou c ops in no he n Mo occo: El Ade gha (34°4ʹ34.19″N, 4°51ʹ33.43″W) and Ki m a (34°10ʹ15.07″N, 5°14ʹ21.43″W)18–20, (Fig.1A). Ten samples we e selec ed encompassing di e en deg ees o bio- u ba ion and a iable o e lapping be ween bu ows (Fig.1B). F om he 10 samples, 30 hin sec ions con aining Maca onichnus we e p epa ed in he Depa men o Ea h Sciences a Royal Holloway Uni e si y o London and imp egna ed wi h blue dyed esin o highligh po osi y. S ue s EpoFix epoxy mixed wi h Sudan Blue II powde was used o he esin. Composi ion o all hin sec ions was desc ibed and eigh o he hin sec ions, whe e Maca onichnus was be e de ined, we e selec ed o conduc modal analyses. Hos ock, ube co e, and im we e di e en ia ed o hese analyses. Ele en quan i a i e modal analyses we e conduc ed by de e mining he composi ion a 300 poin s on each hin sec ion using a s epping s age and associa ed PETROG (Conwy Valley Sys ems Limi ed, UK), a so wa e commonly used in quan i a i e pe og aphy25,26, conside ing he la g- es in ac ec angula a ea a oiding any ma ginal al e a ion. Mine al composi ional analyses o ube co es and ims we e unde aken by de ining an ellip ical a ea wi hin hese bu ow pa s. Rim modal analyses we e no possible in samples om El Ade gha due o im size limi a ions. Po osi y was also measu ed du ing modal analyses in hos ock, ube co e, and im. Addi ionally, po osi y om all he hin sec ions was also ob ained by blue pixels coun ing using Pho oshop27, ob aining a ep esen a i e mean alue o he whole samples and om e e y single pa (i.e., hos ock, ube co e, and im). Blue pixels we e quan i ied using Colo Range Selec ion Me hod28. This me hod allows selec ing all he blue pixels om an a ea o in e es by clicking on some blue pixels and ex ending he selec ion o all he pixels wi h simila alues. Once all he blue pixels a e quan i ied, po osi y can be calcula ed conside ing he numbe o pixels composing he a ea o in e es . In ensi y o bio u ba ion was quan i ied conside ing he bio u ba ed su ace (BS), using Ichnological Digi al Images Analysis Package (IDIAP) quan i a i e me hod28. Theo e ical scena ios we e simula ed conside ing di e en im/ ube co e a ios, deg ee o bio u ba ion and o e lapping be ween bu ows (Fig.3). Fo ha , we selec ed some easonable po osi y alues (2.0% o hos ock, 0.0% o ims and 6.0% o ube co e bu ows) based on hin sec ions obse a ions. Resul ing po osi y was calcula ed using he selec ed po osi y alues o e e y zone conside ing he ela i e su ace o e e y single pa in e e y scena io. Da a a ailabili y All da a analysed du ing he p esen s udy a e summa ized in his published a icle. Analyzed hin sec ions a e a ailable unde eques . Recei ed: 7 Oc obe 2020; Accep ed: 6 Janua y 2021 6 Vol:.(1234567890) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s/ Re e ences 1. Pembe on, S. G. & Ging as, M. K. Classi ica ion and cha ac e iza ions o biogenically enhanced pe meabili y. AAPG Bull. 89, 1493–1517 (2005). 2. Knaus , D. A las o T ace Fossils in Well Co e (Sp inge , New Yo k, 2017). 3. El om, H. A., Rankey, E. C., Hasio is, S. T. & Ba a i, R. E ec o bio u ba ion on pe ophysical p ope ies: Insigh s om geos a is i- cal and low simula ion modeling. Ma . Pe . Geol. 104, 259–269 (2019). 4. Ging as, M. K. e al. Po osi y and pe meabili y in bio u ba ed sedimen s. In T ace Fossils as Indica o s o Sedimen a y En i onmen s (eds Knaus , D. & B omley, R. G.) 837–868 (Else ie , Ams e dam, 2012). 5. Knaus , D. Bio u ba ion and ese oi quali y: owa ds a gene ic app oach. AAPG Annual Con en ion and Exhibi ion. Sea ch and Disco e y A icle #50900. (2013). 6. Pembe on, S. G. & F ey, R. W. T ace ossils nomencla u e and he Planoli es-Palaeophycus dilemma. J. Palaeon ol. 56, 843–881 (1982). 7. Pede sen, G. K. & B omley, R. G. Ophiomo pha i egulai e, a e ace ossil in shallow ma ine sands ones, C e aceous A ane Fo ma ion, Wes G eenland. C e . Res. 27, 964–972 (2006). 8. G eene, T. J., Ging as, M. K., Go don, G. S. & McKeel, D. R. The signi icance o deep-wa e c yp ic bio u ba ion in slope-channel massi e sand deposi s o he lowe Rio Dell Fo ma ion, El Ri e basin, Cali o nia. Ma . Pe . Geol. 29, 152–174 (2012). 9. Quaye, J. A., Jiang, Z. & Zhou, X. Bio u ba ion in luence on ese oi ock quali y: A case s udy o Well Bian-5 om he second membe Paleocene Funing Fo ma ion in he Jinhu sag, Subei basin, China. J. Pe . Eng. 172, 1165–1173 (2019). 10. Sa da, C. E. & Uddin, A. La ge Maca onichnus and hei beha io al implica ions (C e aceous Eu aw Fo ma ion, Alabama, USA). Ichnos 12, 1–9 (2005). 11. Ging as, M. K., MacMillan, B., Balcom, B. J., Saunde s, T. & Pembe on, S. G. Using magne ic esonance imaging and pe og aphic echniques o unde s and he ex u al a ibu es and po osi y dis ibu ion in Maca onichnus-bu owed sands one. J. Sed. Res. 72, 552–558 (2002). 12. Knaus , D. Ichnology as a ool in ca bona e ese oi cha ac e iza ion: A case s udy om he pe mian: T iassic khu o ma ion in he middle eas . GeoA abia 14, 17–38 (2009). 13. Go don, J. B., Pembe on, S. G., Ging as, M. K. & Konhause , K. O. Biogenically enhanced pe meabili y: A pe og aphic analysis o Maca onichnus seg ega us in he lowe c e aceous bluesky o ma ion Albe a Canada. AAPG Bull. 94, 1779–1795 (2010). 14. Da oe, L. T., Ging as, M. K. & Pembe on, S. G. De e mining euzonus muc ona a bu owing a es wi h applica ion o ancien Maca onichnus seg ega is ace-make s. Ichnos 15, 78–90 (2008). 15. Cli on, H. E. & Thompson, J. K. Maca onichnus Seg ega is: A eeding s uc u e o shallow ma ine polychae es. J. Sed. Res. 48, 1–10 (1978). 16. Rod íguez-To a , F. J. & Agui e, J. Is Maca onichnus an exclusi ely small, ho izon al and unb anched s uc u e? Maca onichnus seg ega is degibe i isubsp. no .. Spanish J. Paleon ol. 29, 131–142 (2014). 17. Na a, M. & Seike, K. Palaeoecology o Maca onichnus seg ega is degibe i: Recons uc ing he in aunal li es o he a isiid poly- chae es. Palaeogeog . Palaeocl. Palaeoecol. 516, 284–294 (2019). 18. Miguez-Salas, O., Rod íguez-To a , F. J. & De Wege , W. Maca onichnus and con ou i e deposi ional se ings: Bo om cu en s and nu ien s as coupling ac o s. Palaeogeog. Palaeocl. Palaeoecol. 545, 109639 (2020). 19. Capella, W. e al. Sandy con ou i e d i in he la e Miocene Ri ian Co ido (Mo occo): Recons uc ion o deposi ional en i on- men s in a o eland-basin seaway. Sed. Geol. 355, 31–57 (2017). 20. De Wege , W. e al. La e Miocene con ou i e channel sys em e eals in e mi en o e low beha io . Geology 48, 1. h ps ://doi. o g/10.1130/G4794 4.1 (2020). 21. Knaus , D., Do ado , J. & Rod íguez-To a , F. J. Bu owed ma ix powe ing dual po osi y sys ems: A case s udy om he Maas- ich ian chalk o he Gull aks Field, No wegian No h sea. Ma . Pe . Geol. 113, 104158 (2020). 22. Qui oz, L. I., Bua ois, L. A., Gab iela Mángano, M., Ja amillo, C. A. & San iago, N. Is he ace ossil Maca onichnus an indica o o empe a e o cold wa e s? Explo ing he pa adox o i s occu ence in opical coas s. Geology 38, 651–654 (2010). 23. Cu an, H. A. The ace ossil assemblage o a C e aceous nea sho e en i onmen : English own Fo ma ion o Delawa e, USA. in Biogenic S uc u es: Thei Use in In e p e ing Deposi ional En i onmen s (ed. Cu an, H.A.), 261–276 (1985). 24. Agui e, J., de Gibe , J. M. & Puga-Be nabéu, A. P oximal-dis al ichno ab ic changes in a siliciclas ic shel , Ea ly Pliocene, Gua- dalqui i Basin, sou hwes Spain. Palaeogeog. Palaeocl. Palaeoecol. 291, 328–337 (2010). 25. Pan opoulos, G. & Zelilidis, A. Pe og aphic and geochemical cha ac e is ics o Paleogene u bidi e deposi s in he sou he n Aegean (Ka pa hos Island, SE G eece): Implica ions o p o enance and ec onic se ing. Geochemis y 72, 153–166 (2012). 26. Ca oll, E. L. & Squi es, K. E. Bu ning by numbe s: A pilo s udy using quan i a i e pe og aphy in he analysis o hea -induced al e a ion in bu ned bone. In . J. Os eoa chaeol. 30, 691–699 (2020). 27. Zhang, X. e al. Adobe pho oshop quan i ica ion (PSQ) a he han poin -coun ing: A apid and p ecise me hod o quan i ying ock ex u al da a and po osi ies. Compu . and Geosci. 69, 62–71 (2014). 28. Do ado , J., Rod íguez-To a , F. J. & IODP Expedi ion 339 Scien is s. Quan i a i e es ima ion o bio u ba ion based on digi al image analysis. Ma . Geol. 349, 55–60 (2014). Acknowledgemen s The esea ch by JD was unded h ough a EU’s Ho izon 2020 esea ch and inno a ion p og am unde he Ma ie Skłodowska-Cu ie G an Ag eemen No. 792314 (ICON-SE). OM-S is unded h ough a p e-doc o al g an om he Minis e io de Educación, Cul u a y Depo e, Spain. The p esen con ibu ion was also unded by p ojec s CGL2015-66835-P and PID2019-104625RB-100 (Sec e a ía de Es ado de I+D+I, Spain), B-RNM-072-UGR18, P18-RT-4074 (Jun a de Andalucía), and he Scien i ic Excellence Uni UCE-2016-05 (UGR). The esea ch was conduc ed wi hin he “The D i e s RG” (RHUL) and he “Ichnology and Palaeoen i onmen RG” (UGR). We a e mos g a e ul o he cons uc i e commen s o wo anonymous e e ees who helped us o imp o e he cla - i y o he manusc ip . Au ho con ibu ions J.D., F.J.R.-T. and O.M.-S. designed he idea, collec ed and selec ed he samples. J.D. coo dina ed he composi- ion and conduc ed he po osi y analysis. All he au ho s discussed he esul s, in e p e a ions and con ibu ed o he inal e sion o he manusc ip . Compe ing in e es s The au ho s decla e no compe ing in e es s. 7 Vol.:(0123456789) Scien i ic Repo s | (2021) 11:1975 | h ps://doi.o g/10.1038/s41598-021-81687-6 www.na u e.com/scien i ic epo s/ Addi ional in o ma ion Co espondence and eques s o ma e ials should be add essed o J.D. Rep in s and pe missions in o ma ion is a ailable a www.na u e.com/ ep in s. Publishe ’s no e Sp inge Na u e emains neu al wi h ega d o ju isdic ional claims in published maps and ins i u ional a ilia ions. Open Access This a icle is licensed unde a C ea i e Commons A ibu ion 4.0 In e na ional License, which pe mi s use, sha ing, adap a ion, dis ibu ion and ep oduc ion in any medium o o ma , as long as you gi e app op ia e c edi o he o iginal au ho (s) and he sou ce, p o ide a link o he C ea i e Commons licence, and indica e i changes we e made. The images o o he hi d pa y ma e ial in his a icle a e included in he a icle’s C ea i e Commons licence, unless indica ed o he wise in a c edi line o he ma e ial. I ma e ial is no included in he a icle’s C ea i e Commons licence and you in ended use is no pe mi ed by s a u o y egula ion o exceeds he pe mi ed use, you will need o ob ain pe mission di ec ly om he copy igh holde . To iew a copy o his licence, isi h p://c ea i eco mmons .o g/licen ses/by/4.0/. © The Au ho (s) 2021