Holocene summer temperature reconstruction based on a chironomid record from Sierra Nevada, southern Spain

Qua e na y Science Re iews 319 (2023) 108343 A ailable online 29 Sep embe 2023 0277-3791/© 2023 The Au ho s. Published by Else ie L d. This is an open access a icle unde he CC BY-NC license (h p://c ea i ecommons.o g/licenses/by- nc/4.0/). Holocene summe empe a u e econs uc ion based on a chi onomid eco d om Sie a Ne ada, sou he n Spain Gonzalo Jim´ enez-Mo eno a , * , Oli e Hei i b , An onio Ga cía-Alix a , R. Sco Ande son c , F ancisco J. Jim´ enez-Espejo d , Cha o L´ opez-Blanco a , Lau a Jim´ enez a , Ca men P´ e ez-Ma ínez e , Ma a Rod igo-G´ amiz a , Alejand o L´ opez-A il´ es a , Jon Camue a d a Depa men o S a ig aphy and Paleon ology, Uni e si y o G anada, 18002, G anada, Spain b Geoecology, Depa men o En i onmen al Sciences, Uni e si y o Basel, 4056, Basel, Swi ze land c School o Ea h & Sus ainabili y, No he n A izona Uni e si y, Flags a , 86011, AZ, USA d Andalusian Ea h Sciences Ins i u e (IACT), Spanish Resea ch Council – Uni e si y o G anada (CSIC-UGR), 18100, A milla, (G anada), Spain e Depa men o Ecology, Uni e si y o G anada, 18071, G anada, Spain ARTICLE INFO Handling Edi o : P Rioual Keywo ds: Chi onomids Summe empe a u e Holocene Las glacial pe iod Sie a Ne ada Wes e n Eu ope ABSTRACT Ob aining accu a e empe a u e econs uc ions om he pas is c ucial in unde s anding he consequences o changes in ex e nal clima e o cings, such as o bi al-scale insola ion o mul idecadal o cen ennial-scale a i- abili y on he clima e sys em and he en i onmen . In addi ion, hese econs uc ions help in comp ehending he ampli ude o na u al empe a u e changes in he pas , which can assis in e alua ing he ampli ude and a e o ecen an h opogenic global wa ming. He e we p esen he i s de ailed Holocene mean July ai empe a u e econs uc ion based on chi onomid assemblages om sedimen s e ie ed om Laguna de Río Seco, an alpine lake in Sie a Ne ada, sou he n Spain. Coldes clima e condi ions a e eco ded du ing he las glacial maximum and he las deglacia ion. Wa ming occu ed in he Ea ly Holocene and wa mes summe empe a u e condi ions and he Holocene he mal maximum (HTM) occu ed in he in e al oughly be ween 9000 and 7200 cal y BP, concu en wi h summe insola ion maxima. Rapid cooling o ~1.5 ◦C occu ed a e he wa mes maximum and be ween ~7200 and 6500 cal y BP, and empe a u es s abilized be ween ~6500 and 3000 cal y BP. A u he cooling began ~3000 cal y BP and culmina ed wi h coldes summe condi ions du ing he Da k Ages (DA) and Li le Ice Age (LIA) a ~1550 cal y BP (~400 CE) and ~200 cal y BP (~1750 CE), espec i ely. This cooling empe a u e end was in e up ed by wa me condi ions du ing he Ibe ian-Roman Humid Pe iod (IRHP) ~2000 cal y BP and du ing he Medie al Clima e Anomaly (MCA) a ~1000 cal y BP. Ou econs uc ion shows a g ea e han wo-deg ee cooling du ing he Middle and La e Holocene, ag eeing wi h global mean su ace empe a u e (GMST) econs uc ions. Mode n clima e wa ming (MCW) du ing summe exceeds he wo-deg ee Celsius o ecas ed o he u u e due o an h opogenic g eenhouse gases, sugges ing ha ecen wa ming is ampli ied a high ele a ions. Alpine en i onmen s and he biodi e si y con ained he e a e hus in dange i he obse ed empe a u e end con inues in he nex decades. 1. In oduc ion The In e go e nmen al Panel on Clima e Change (IPCC e al., 2022) p edic s a possible inc ease in global empe a u es o ~2 ◦C (highe han he pe iod 1850–1900 CE) by he end o he 21s cen u y. Impac s o inc easing empe a u es in moun ains ha e been documen ed in ecen decades wi h obse able and se ious consequences o socie ies, biodi- e si y, and ecosys ems. These include, o example, changing seasonal wea he pa e ns, educ ions in snow co e ex en and du a ion a low ele a ion, loss o glacie mass, inc eased pe ma os haw, changes in he iming o glacie - and snow-mel discha ge, enhanced i e ac i i y, and changing na u al dis ibu ion o plan and animal species shi ing o highe ele a ions (IPCC e al., 2022). P e ious s udies show ha wa ming will be mo e p onounced in high al i ude alpine a eas (Pepin e al., 2022) causing d ama ic impac s o hose en i onmen s. Some a eas o sou he n Eu ope a e he mos esponsi e egions o MCW, * Co esponding au ho . E-mail add ess: [email p o ec ed] (G. Jim´ enez-Mo eno). Con en s lis s a ailable a ScienceDi ec Qua e na y Science Re iews jou nal homepage: www.else ie .com/loca e/quasci e h ps://doi.o g/10.1016/j.quasci e .2023.108343 Recei ed 23 July 2023; Recei ed in e ised o m 25 Sep embe 2023; Accep ed 26 Sep embe 2023 Qua e na y Science Re iews 319 (2023) 108343 2 especially in summe (Lo enzo and Al a ez, 2022). In addi ion, he Medi e anean side o sou he n Ibe ia appea s o be especially sensi i e as he occu ence o summe hea wa es has inc eased by 4% pe decade be ween 1950 and 2020 CE (Díaz-Poso e al., 2023). Unde s anding he long- e m ela ionship be ween en i onmen al change caused by clima e change in moun ain a eas om he Medi e - anean basin is pa amoun o comp ehend ecosys em esponse o inc easing empe a u es in his clima e-sensi i e a ea. To accomplish his, de ailed paleoecological and paleoclima ic s udies a e necessa y, especially o pe iods ha a e wa me han p esen . In his espec , he sedimen a y eco d om he alpine lake Laguna de Río Seco (LdRS; Fig. 1) has al eady p o ided de ailed insigh s in o en i onmen al change in he Sie a Ne ada ange o sou he n Spain and he wes e n Medi e - anean a ea in gene al (sedimen a ion, eolian inpu , ege a ion and lake en i onmen al geochemis y) including in o ma ion on pas clima e a ia ions and human impac du ing he Holocene (~las 12,000 yea s; Ande son e al., 2011; Ga cía-Alix e al., 2013, 2018, 2020; Jim´ enez-Espejo e al., 2014; Jim´ enez e al., 2018, 2019; Toney e al., 2020). Howe e , mos o hese p e ious paleoen i onmen al s udies we e only in e p e ed in e ms o quali a i e paleoclima ic change and he only a ailable empe a u e econs uc ion comes om algae-de i ed long-chain diol analysis (Ga cía-Alix e al., 2020; Toney e al., 2020). Accu a e quan i a i e empe a u e econs uc ions a e c ucial o cali- b a e esponses o he clima e sys em o ex e nal o cing such as o bi al-scale changes in insola ion o mul idecadal o cen ennial-scale clima e a iabili y and o help modele s pa ame e ize hei clima e p ojec ions unde di e en scena ios in he pas , he eby con ibu ing o mo e accu a e clima e p edic ions o he u u e (e.g., Hei i e al., 2014; PAGES 2k-PMIP3 g oup, 2015; Sama in e al., 2017). He e we show a de ailed ossil chi onomid analysis o he Laguna de Río Seco eco d (LdRS-01, co e ing he pas ~21,000 cal y BP; Fig. 2). Remains o chi onomid la ae, mainly hei chi inous head capsules, a e p ese ed well in Pleis ocene and Holocene lake sedimen s (Bolland e al., 2020, 2021; Bennike e al., 2023). The head capsules o he di e en species possess unique mo phological cha ac e is ics pe mi - ing hei iden i ica ion a he gene ic, species-g oup, o mo pho ype le els (B ooks e al., 2007). Chi onomid assemblages a e known o be sensi i e o summe empe a u es in con inen al en i onmen s (B ooks, 2006; Egge mon and Hei i, 2012) and can be used o es ima e pas summe empe a u e a iabili y based on changes in species assem- blages (e.g., B ooks and Bi ks, 2001; Sama in e al., 2017). The LdRS-01 chi onomid eco d, he e o e, allows he de elopmen o a quan i a i e summe empe a u e econs uc ion o he Sie a Ne ada egion since he end o he Las Glacial Maximum (LGM). The compa ison o his summe empe a u e eco d wi h o he paleoclima e eco ds will help e ine ou unde s anding o egional clima e dynamics du ing he deglacia ion and he Holocene, he ex en ha ecen clima e wa ming exceeds his baseline a iabili y, and he impac s o clima ic change on he en i onmen in a moun ain a ea in sou he n Spain. 2. S udy si e and egional se ing LdRS (37◦03 ′ 7 ′′ N, 3◦20 ′ 44 ′′ W, 3029 m a.s.l.) is in a sou h- acing glacial ci que basin loca ed in he Sie a Ne ada ange, sou he n Spain (Fig. 1). The lake has a maximum dep h o ~3 m, a su ace a ea o 0.42 ha, and a d ainage basin o 9.9 ha (Mo ales-Baque o e al., 1999). Clima e in he a ea is Medi e anean, wi h ho and d y summe s. Me eo ological eco ds a 2500 m a.s.l. show mean annual empe a u e o ~4.4 ◦C, anging om ~18 ◦C in summe o ~ −4 ◦C in win e . The annual p ecipi a ion a his ele a ion is be ween ~700 and 750 mm and mainly alls as snow be ween Oc obe and May (Obse a o io del cam- bio global de Sie a Ne ada, 2020; Spanish Na ional Wea he Agency – Fig. 1. Loca ion (A and B) and pho os (C and D) o LdRS in Sie a Ne ada, sou he n Spain. (A) Map o he Sie a Ne ada (SN in A) moun ain ange, sou he n Ibe ian Peninsula (modi ied om Jim´ enez-Mo eno e al., 2022). The ec angle shows he magni ied a ea shown in (B). (C) Sa elli e pho o o he LdRS a ea om Google Ea h. (D) Field pho o o LdRS wi h he loca ion o he co e aken in Sep embe 2006. The Mulhac´ en, he highes peak in he Ibe ian Peninsula wi h 3479 m a.s.l., is in he backg ound. G. Jim´ enez-Mo eno e al. Qua e na y Science Re iews 319 (2023) 108343 3 AEMe Open Da a, 2022). The LdRS lake wa e empe a u e has been moni o ed since 2011 and oscilla es be ween ~0 ◦C in Feb ua y–Ma ch and ~16 ◦C in July–Augus , wi h a mean empe a u e in July o 14.7 ◦C and a mean empe a u e o he summe mon hs (June-July-Augus ) o 12 ◦C. Su ace ai empe a u es a ound LdRS ha e been measu ed om 2019 o 2022 and ange om −5 o 15 ◦C, wi h a mean annual ai empe a u e o 2.7 ◦C, mean July empe a u e o 14 ◦C and a mean empe a u e o he summe mon hs o 13 ◦C. LdRS is meso-oligo ophic (TP =16 μ g/L, TN =403 μ g/L; Chlo o- phyll: ~0.5-2 μ g/L) (Jim´ enez e al., 2019; Mo ales-Baque o and Conde-Po cuna, 2000) and i s e y na ow sho eline is co e ed by b yophy es. I has in e mi en wa e in lows, p o iding wa e o he lake du ing la e sp ing and ea ly summe , and a small ou le ha is only ac i e du ing he highes lake wa e le el (in June–July). The lake is co e ed in ice and snow om a ound Oc obe –No embe un il May–June, depending on he yea . The pH alues ange om 6 o 7.4, conduc i i y does no exceed 80 μ S/cm, and he dissol ed o ganic ca - bon alues ange om 0.74 o 3.40 mg/L. I does no he mally s a i y (see he mis o da a in Ga cía-Ju ado e al., 2011) so he bo om is highly oxygena ed, and du ing he ice- ee pe iod, Secchi disk isibili y exceeds he wa e dep h. I is a ishless lake wi h low plank onic di- e si y (Jim´ enez e al., 2019). Mo e de ailed limnological da a a e compiled by e.g. S´ anchez-Cas illo e al. (1989), Mo ales-Baque o e al. (1999) and Reche e al. (2005). LdRS (37◦03 ′ N, 3◦20 ′ W) is si ua ed abo e he eeline (abo e ~2500 m a.s.l.), whe e ex eme clima e condi ions occu . Vege a ion in hese alpine en i onmen s is ypically c yo omedi e anean, cha ac e ized by und a-like open g assland and plan s wi h basal ose es (Valle, 2003; Ande son e al., 2011). The LdRS ca chmen a ea is pa ially ege a ed (~15%) by alpine meadows, p ima ily made up o sedges (Cype aceae) and g asses (Poaceae), and xe ophy ic sh ubland (Ande son e al., 2011). The es o he ca chmen a ea is cha ac e ized by ba e ocks ha like he lake bed ock a e mainly composed o mica schis s (Díaz de Fede ico, 1980). 3. Ma e ials and me hods Two adjacen sedimen co es we e collec ed in Sep embe 2006 a he depocen e o LdRS om a loa ing pla o m ancho ed o he sho e. These included a 150 cm-long co e (LdRS-06-01) using a Li ings one squa e- od pis on co e , and a 37 cm-long co e (LdRS-06-02) using a uni e sal co e wi h a plexiglass ube (Aqua ic Resea ch, Inc.) o eco e he unconsolida ed uppe sedimen s. Maximum dep h o he lake when co ed was 1.7 m. Sedimen cha ac e is ics o bo h co es we e desc ibed in he labo- a o y (see Ande son e al., 2011 o mo e de ails; Fig. 2). Magne ic suscep ibili y (MS) o LdRS-06-01 was de e mined using a Ba ing on MS2E me e wi h measu emen s aken e e y 5 mm h oughou he leng h o he co e (Fig. 2). MS was no done on co e LdRS-06-02 as he sedimen s we e sampled in he ield and we e pu in whi lpack bags. Nine bulk sedimen samples in he LdRS-06-01 co e we e p e iously analyzed o adioca bon da ing and 11 samples om he uppe 16 cm o co e LdRS-06-02 we e p e iously analyzed o 210 Pb and 137 Cs da ing (Ande son e al., 2011). In his s udy, a u he ba ch o wel e bulk sedimen samples we e aken om he lowe mos and uppe mos pa s o he LdRS-06-01 co e o adioca bon analysis (Table 1; Fig. 2). All he esul s om adioca bon analyses (21 adioca bon da es in o al) we e con e ed o calib a ed adioca bon yea s be o e p esen (cal y BP, wi h p esen aken a 1950 CE by con en ion) using he In Cal20 calib a ion cu e (Reime e al., 2020) and a e ised Bayesian age-model has been calcula ed using he R-based Bacon package, e sion 3.1.0 (Blaauw and Ch is en, 2011, Fig. 2). Two seemingly old samples (in ed in Table 1) we e no included in he age-dep h model. The age-dep h model ha p o ided he lowes unce ain ies was ob ained by compu ing Ma ko Chain Mon e Ca lo (MCMC) in e ela ions in 30 sec ions o 5 cm wi h he accumula ion p io s 2.25 (shape) and 100 (mean). Low alues we e se o he memo y p io s (memo y s eng h =5 and memo y mean = 0.1) due o he la ge shi s in he accumula ion a es a he op and a he bo om o he co e (Fig. 2). Fig. 2. Li hology, Bayesian age-dep h model, and MS o he LdRS eco d. Li hology and MS we e p e iously desc ibed in Ande son e al. (2011). On he le , SC s ands o sho co e RS eco e ed in 2008 (see de ails in Jim´ enez e al., 2019). Blue a eas in he age-dep h model ep esen he calib a ed adioca bon ages along wi h hei unce ain ies, he ed line indica es he weigh ed median ages and he do ed line a ound he g ey shaded a ea shows he 95% con idence in e al o he modelled ages. The di e en pa ame e s and p io s o he model a e explained in he ex . G. Jim´ enez-Mo eno e al. Qua e na y Science Re iews 319 (2023) 108343 4 Fo y-eigh samples o 2 cm 3 om co e LdRS-06-01 we e p ocessed o chi onomid analysis, which in ol ed chemical ea men wi h so- dium hexame aphospha e, sie ing a 100- μ m, so ing o chi onomids om sie e esidue a ca. 30x magni ica ion, and iden i ica ion o chi- onomids on slides wi h Eupa al medium wi h a ligh - ansmi ed mi- c oscope a 100x - 400x magni ica ion (B ooks e al., 2007). The axonomical iden i ica ions mos ly ollowed B ooks e al. (2007) (Fig. 3). These chi onomid coun s we e combined wi h p e iously published chi onomid da a om en samples om ano he sho co e aken om he depocen e o he LdRS basin, RS, co e ing 155 o −55 cal y BP (Jim´ enez e al., 2019). This way, a composi e chi onomid e- co d o chi onomid assemblage change in LdRS was p oduced om wo co es, LdRS-06-01 and RS, which we e independen ly da ed and we e co ela ed by ch onos a ig aphy. Chi onomid ela i e abundances (%) we e calcula ed wi h espec o he o al chi onomid coun in each sample and we e plo ed using TILIA (G imm, 1992) in a de ailed dia- g am shown in Fig. 4 whe e all he iden i ied axa a e shown. A con- s ained inc emen al sum o squa es clus e analysis (CONISS, G imm, 1987) was pe o med o de e mine pe iods wi h simila chi onomid assemblages (Fig. 4). A De ended co espondence analysis (DCA) was applied o he chi onomid da a o de e mine whe he linea (PCA) o unimodal s a- is ical (CA) echniques a e he mos app op ia e o model he species esponse, es ima ed as uni s o composi ional u no e in s anda d de- ia ion (SD) o he i s majo g adien o he DCA. As he leng h o he explo a o y DCA o he squa ed- oo ans o med da a o chi onomids was 2.5 SD, he assemblage a ia ion is wi hin a ela i ely na ow ange and linea me hods such as PCA a e app op ia e (Lepˇ s and ˇ Smilaue , 2003; Legend e and Bi ks, 2012). PCA was un on he chi onomid pe - cen age da a using he Pas 4 so wa e (Hamme e al., 2001). PCA de- e mines a iables (componen s) ha ep esen , as much as possible, he a iance o he mul i a ia e da a (Hamme e al., 2001). These new componen s a e linea combina ions o he o iginal a iables. The PCA can be used o summa ize he da a o, o example, only wo a iables ( he i s wo componen s) o compa ison in g aphs. A PCA co ela ion loading and sca e biplo diag ams a e shown in Fig. 5. The mos clima e-sensi i e indica o s om chi onomid (Mic opsec a adialis ype), pollen (A emisia) and MS da a a e plo ed oge he o isual compa ison on Fig. 6. Based on he ossil chi onomid pe cen age da a mean July ai em- pe a u e es ima ions we e p oduced using a 274-lake chi onomid- empe a u e calib a ion da ase and ans e unc ion om he Swiss Alps and No way ha co e s a mean July ai empe a u e g adien om 4 o 18.4 ◦C and a wide ange o a c ic, alpine, subalpine and empe a e lakes (Hei i e al., 2011). This econs uc ion was de eloped using he p og am C2 (Juggins, 2007) and he applied ans e unc ion was based on weigh ed a e aging pa ial leas squa es eg ession (WA-PLS; Te B aak and Juggins, 1993; Te B aak e al., 1993). The model ea u ed a c oss- alida ed 2 o 0.9 and a oo -mean-squa e e o o p edic ion (RMSEP) o 1.39 ◦C. The RMSEP, coe icien o de e mina ion (R 2 ) and sample-speci ic e o s o p edic ion (eSEPs) we e calcula ed based on 9999 boo s apping cycles in C2. Chi onomid assemblage pe cen age da a we e squa e oo ans o med be o e he calcula ion o WA-PLS and dis ance me ics. A o al o 19 si es wi h unusual hyd ological o ecological condi ions and high p edic ion esiduals we e dele ed om he model as ou lie s as desc ibed in Hei i e al. (2011). The quan i a i e summe (June-July-Augus ) empe a u e om LdRS was also econs uc ed using he pollen eco d (Ande son e al., 2011), which includes 69 ossil pollen samples om co es LdRS-06-01 and LdRS-06-02. In o de o ge an accu a e econs uc ion o a wes - e n Eu opean high-al i ude moun ain a ea, we used he Eu asian Mod- e n Pollen Da abase 2 (EMPD 2) (Da is e al., 2020) delimi ed o la i ude 25◦-50◦N and longi ude 15◦W-5◦E, and excluding low-al i ude (below 1500 m a.s.l.) mode n pollen si es. The pollen axonomy was ha monized using he Plan s o he Wo ld online da abase (www.plan so hewo ldonline.o g) and he In eg a ed Taxonomic In o ma ion Sys em (www.i is.go ). Aqua ic plan s we e emo ed, assuming ha he dis- ibu ion o hese plan s could be a ec ed by o he ac o s no ela ed o clima e. The e o e, he aining se is based on 300 mode n pollen si es and 194 ha monized pollen axa. The ecen empe a u e o each mode n pollen si e was ob ained om he Wo ldClim 2.1 da abase unde a 30 s esolu ion (www.wo ldclim.o g) (Fick and Hijmans, 2017). The pollen-based ans e unc ion om LdRS was de eloped using he C2 so wa e unde e sion 1.7.7 (Juggins, 2007). We used he wo-componen WA-PLS me hod unde lea e-one-ou c oss- alida ion. The WA-PLS me hod assumes ha each axon has a unimodal dis ibu- ion wi h espec o clima ic pa ame e s and i is obus o spa ial au oco ela ion (Tel o d and Bi ks, 2005). The squa e- oo species ans o ma ion was used o educe he noise o he da a. The WA-PLS o he summe empe a u e p o ided a lea e-one-ou , c oss alida ed R 2 alue o 0.71 and a RMSEP o 1.8 ◦C. 4. Resul s 4.1. New ch onology o LdRS eco d The wel e new adioca bon analyses, oge he wi h he nine p e- ious analyses, indica e ha he LdRS-06-01 co e con ains a eco d o he pas ~21,000 cal y BP (Fig. 2; Table 1). The e ised ch onology shows ha he LdRS sedimen a y eco d is olde han p e iously hough (~12,000 cal y BP; Ande son e al., 2011), now da ing back o a leas he end o he LGM. This new ch onology mos ly conce ns he Table 1 Radiome ic ages om Laguna de Río Seco co es 06-01 and 06-02. In bold he wel e new adioca bon analyses ca ied ou in his s udy. In ed he da es ha we e ejec ed o he age model cons uc ion. Radioca bon ages we e cali- b a ed using he In Cal 20 calib a ion cu e (Reime e al., 2020). 6 G. Jim´ enez-Mo eno e al. Qua e na y Science Re iews 319 (2023) 108343 5 Fig. 3. Pho os o chi onomid emains om he LdRS sedimen a y eco d: A - Mic opsec a adialis ype- LdRS D .03 10–11 cm; B - Chi onomus plumosus ype- LdRS D .02 10–11 cm; C - Co ynoneu a a c ica ype- LdRS D .01a 20–21 cm; D - Co ynoneu a a c ica ype LdRS D 01a 20–21, head capsule su ace o namen a ion de ail; E − Me iocnemus- LdRS D .01 64–65 cm; F - Psec ocladius so didellus ype LdRS D .01a 20–21 cm; Psec ocladius (Allosep ocladius) la us ype- LdRS D .03 6–7 cm; Mic opsec a insignilobus ype- LdRS D .01 84–85 cm; He e o issocladius ma cidus ype- LdRS D .02 30–31 cm. G. Jim´ enez-Mo eno e al. Qua e na y Science Re iews 319 (2023) 108343 6 bo ommos pa o he LdRS eco d, dila ing he p e ious eco d om Ea ly Holocene (EH) o he La es Pleis ocene age. The accumula ion a e o he bo ommos pa o he eco d is e y low 0.0007 cm/y , con as ing wi h he mean sedimen a ion a e o he uppe mos 135 cm (0.014 cm/y ). As he new adioca bon da ings o he bo ommos pa o he co e a e somehow ambiguous (25,862 y BP a 136 cm, 19,803 y BP a 140 cm and 20,930 y BP a 143 cm), we use a conse a i e app oach o he p e-Holocene sec ion and used gene al e ms as “Deglacia ion sensu la o” o “La e Pleis ocene” (see below). 4.2. Chi onomid esul s Fi een mo pho ypes o chi onomid axa we e iden i ied in he composi e chi onomid eco d ha consis s o 48 analyzed samples om co e LdRS-06-01 and 10 p e iously published chi onomid samples om Jim´ enez e al. (2019). Chi onomid concen a ions in 2 cm 3 o sandy sedimen s om he bo ommos pa o he sedimen a y eco d we e low, wi h an a e age numbe o 50 iden i ied head capsules, due o hei e y ino ganic na u e (Fig. 4). Howe e , chi onomid abundance inc eased conside ably in he EH and emained high (a e age o 130) du ing he es o he Holocene. Pho og aphs o he mos common and clima ically signi ican axa a e shown in Fig. 3. Chi onomid di e si y in Fig. 4. De ailed diag am o chi onomid assemblages changing o e ime in he LdRS eco d. Shadings show 5x exagge a ion. On he igh a e he six chi onomid zones iden i ied using cons ained inc emen al sum o squa es (CONISS; G imm, 1987) (see ex o explana ion). Fig. 5. P incipal componen analyses (PCA) om he LdRS chi onomid da a (A and B). A PCA co ela ion loading ( o componen 1) is shown in (A) and sca e biplo diag am is shown in (B). The analysis was ca ied ou using PAST 4.10 (Hamme e al., 2001). In e p e ed PCA g oups a e shown (see ex o explana ion). G. Jim´ enez-Mo eno e al. Qua e na y Science Re iews 319 (2023) 108343 7 he s udied samples is ela i ely low, wi h 3–9 mo pho ypes pe sample. Mic opsec a adialis ype, Psec ocladius so didellus ype and Co ynoneu a a c ica ype a e he mos abundan axa in he sedimen a y eco d (Figs. 3 and 4). P onounced changes a e obse ed in hese and o he axa and wi h he help o he clus e analysis and isual obse a ion we we e able o dis inguish he ollowing pe iods (zones). 4.2.1. Deglacia ion sensu la o (DG) - (146.5–132 cm; ~20,900–9000 cal y BP) The assemblages du ing his pe iod including he la es Pleis ocene and pa o he EH a e cha ac e ized by he abundance o Mic opsec a adialis ype, he dominan axon wi h abundances be ween 80 and 100%, and Psec ocladius (Allopsec ocladius) la us ype. 4.2.2. Ea ly Holocene sensu la o (EH) – (132-107 cm; ~9000-7100 cal y BP) Psec ocladius so didellus ype and Co ynoneu a a c ica ype appea ed and inc eased du ing he EH and ea ly MH, eaching a ound 50 and 40%, espec i ely. Chi onomus plumosus ype also appea ed a he beginning o his pe iod and disappea ed a he end o i , showing a e age ela i e abundance o a ound 5%. Mic opsec a adialis ype dec eased conside ably o pe cen ages lowe han 5% and dec eases o Psec ocladius (Allopsec ocladius) la us ype a e also obse ed du ing his zone. Chi onomus an h acinus ype occu ed a he beginning o his pe iod. This pe iod is also cha ac e ized by he occu ence o He e o- issocladius ma cidus ype and Me iocnemus ype, bo h eaching a maximum abundance o a ound 1%. 4.2.3. Middle Holocene (MH) - (107-59 cm; ~7100-4200 cal y BP) Psec ocladius so didellus ype displays an inc easing end du ing his pe iod, eaching maxima highe han 85% a ~5800 and 4500 cal y BP. Co ynoneu a a c ica ype seems o show a dec easing end, wi h ma ked a iabili y opposi e o he abundances o Psec ocladius so didellus ype, and ea u es minima a ~5800 and 4500 cal y BP o a ound 12%. Psec ocladius (Allopsec ocladius) la us ype con inued dec easing in his zone and disappea ed a he end o i . Mic opsec a adialis ype occu ed wi h pe cen ages a ound 5% o lowe in his pe iod. 4.2.4. La e Holocene 1 (LH1) - (59-26 cm; ~4200-2000 cal y BP) Psec ocladius so didellus ype shows a dec easing end a he beginning o his zone, eaching minima wi h pe cen ages o 55% a 4000 and 3600 cal y BP and an inc easing end a he end o he zone wi h alues a ound 70%. Co ynoneu a a c ica ype shows he opposi e ends, inc easing i s , eaching maxima (42%) a ound 3600 and 3100 cal y BP and hen dec easing down o 20% a he end o he pe iod. Mic opsec a adialis ype inc eased a he end o his zone, eaching a peak o 10% be ween ~2700-2600 cal y BP. 4.2.5. La e Holocene 2 (LH2) - (26–5.75 cm; ~2000-0 cal y BP) Psec ocladius so didellus ype depic s an inc easing end in his zone, eaching alues abo e 85% a he end o i a ~900 and 0 cal y BP. Co ynoneu a a c ica ype dec eased conside ably, eaching a minimum o almos 0% a ~900 and be ween 100 and 50 cal y BP. Mic opsec a adialis ype con inued inc easing in his pe iod o a peak o ~30% a ~1600 cal y BP, subsequen ly dec eased o ~12% a ~900 cal y BP and inc eased again o 25% a he end o his zone a ~150 and 78 cal y BP. Fig. 6. Compa ison o indi idual chi onomid (Mic opsec a adialis ype), and pollen (A emisia) axa wi h sedimen a ion da a (MS) om LdRS. MS and A emisia da a we e p e iously published in Ande son e al. (2011). No e he isual co a ia ion be ween he independen LdRS en i onmen al indica o s and chi onomid da a. La e Pleis ocene and Ea ly, Middle and La e Holocene bounda ies a e shown wi h dashed lines. IRHP, DA, MCA and LIA s and o Ibe ian Roman Humid Pe iod, Da k Ages, Medie al Clima e Anomaly and Li le Ice Age. G. Jim´ enez-Mo eno e al. Qua e na y Science Re iews 319 (2023) 108343 8 4.2.6. Mode n clima ic wa ming (MCW) - (5.75–0 cm; 0 o −58 cal y BP; 1950–2008 CE) The mos no ewo hy cha ac e is ic o his pe iod consis s o he enewed a i al o Chi onomus plumosus ype and He e o issocladius ma cidus ype, axa ha we e p e iously abundan in he EH, as well as o Mic opsec a insignilobus ype (Fig. 4). M. insignilobus ype inc eased in he pas 50 yea s and eaches a ound 20% a p esen . He e o issocladius ma cidus ype and Chi onomus plumosus ype, which las occu ed a ~8500 and 7000 cal y BP in he LdRS eco d, espec i ely, eappea ed again since 1970 CE. 4.3. PCA analysis o chi onomid da a PCA indica es ha p incipal componen axis 1 (PC1) accoun s o much o he a iabili y o he da a, explaining 73.5% o he a iance. PC2 explains 24.9% o he a iance. The PCA co ela ion loadings and sca e diag am shown in Fig. 5 illus a e o wha deg ee he di e en axa co ela e wi h he di e en componen s. PCA shows wo main g oups o dis inc i e axa (Fig. 5). One g oup (A), cha ac e ised by posi i e co ela ion o PC1, is mos ly made up o Co ynoneu a a c ica ype and Psec ocladius so didellus ype bu also by Me iocnemus, Mic opsec a insignilobus ype, M. con ac a ype and Chi onomus plumosus ype. A second g oup (B) is cha ac e ised by nega i e co ela ion alues o PC1 and is mos ly cha ac e ized by Mic opsec a adialis ype bu also by Psec ocladius (Allopsec ocladius) la us ype. A subdi ision o g oup A would be possible be ween axa ha show posi i e co ela ions o PC2 (A+), mos ly cha ac e ized by Co ynoneu a a c ica ype, and axa ha show a nega i e co ela ion o PC2 (A-) domina ed by Psec ocladius so didellus ype (Fig. 5B). Clus e analysis o he chi onomid da a ag ees wi h he PCA wi h chi onomid zone DG cha ac e ized by high ela i e abundances o chi onomids in he PCA B g oup (domina ed by Mic opsec a adialis ype), zones EH, MH and LH1 by maxima o axa belonging o he PCA A g oup, such as Co ynoneu a a c ica ype and Psec ocladius so didellus ype, wi h a ying abundances o axa in he A+and A-g oups. LH2 is cha ac e ized by an inc ease in chi onomid axa om PCA B g oup and MCW wi h highe abundances o hose in he A g oup (Figs. 4 and 5). 4.4. Chi onomid-in e ed empe a u e esul s Chi onomid-in e ed July ai empe a u es du ing zone DG oscilla e be ween 4.5 and 7.5 ◦C (mean empe a u e a ound 6 ◦C), he lowes alues in he eco d, and inc ease dis inc ly a he end o his pe iod and onse o he Holocene (Fig. 7). The inc easing empe a u e end con- inues du ing he EH, eaching empe a u e maxima o 10.5–10.8 ◦C be ween ~8600 and 7200 cal y BP. A minimum empe a u e o ~9.2 ◦C is eco ded be ween hose wo maxima a ~7350 cal y BP. Dec easing empe a u es a e obse ed du ing he beginning o he MH, eaching a pla eau o a ound 9 ◦C be ween ~6500 and 3000 cal y BP. Tempe a- u es emained s able du ing he i s pa o LH1 un il ~3000 cal y BP when hey dec ease down o a ound 8.5 ◦C a ~2700 cal y BP. A sligh inc ease in empe a u es o 9 ◦C occu s a he end o LH1 and he beginning o LH2 be ween ~2250 and 1800 cal y BP. A dec ease in empe a u es occu s subsequen ly, eco ding a minimum below 8 ◦C a ~1550 cal y BP. An inc ease occu s a e wa ds and a peak sligh ly abo e ~8 ◦C is obse ed a ~1200 cal y BP (~700 CE) (Fig. 8). Tem- pe a u es dec ease la e on and a minimum a ound 8 ◦C is eached be- ween ~1450-1800 and 1900 CE, sepa a ed by an inc ease o 8.5 ◦C a ~1850 CE. The chi onomid-based ai summe empe a u e econs uc- ion sugges s an inc easing end om 1950 CE un il he p esen eaching ~10.6 ◦C, one o he highes empe a u e alues eached du ing he Holocene (Figs. 8 and 9). Fig. 7. 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