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
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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´
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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´
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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.
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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. Chi onomid-in e ed July ai empe a u es om he LdRS composi e eco d (solid lines) and associa ed empe a u e es ima e e o (eSEP; dashed lines) o
each sample. IRHP, DA, MCA, LIA and MCW s and o Ibe ian Roman Humid Pe iod, Da k Ages, Medie al Clima e Anomaly, Li le Ice Age and Mode n
Clima e Wa ming.
G. Jim´
enez-Mo eno e al.
Qua e na y Science Re iews 319 (2023) 108343
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(cap ion on nex page)
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enez-Mo eno e al.
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Ga cía-Alix, A., Toney, J.L., Jim´
enez-Mo eno, G., P´
e ez-Ma ínez, C., Jim´
enez, L.,
Rod igo-G´
amiz, M., Ande son, R.S., Camue a, J., Jim´
enez-Espejo, F.J., Pe˜
na-
Angulo, D., Ramos-Rom´
an, M.J., 2020. Algal lipids e eal unp eceden wa ming
a es in alpine a eas o SW Eu ope du ing he Indus ial Pe iod. Clim. Pas 16,
245–263.
Ga cía-Alix, A., Camue a, J., Ramos-Rom´
an, M.J., Toney, J.L., Sachse, D., Sche uß, E.,
Jim´
enez-Mo eno, G., Jim´
enez-Espejo, F.J., L´
opez-A il´
es, A., Ande son, R.S.,
Yanes, Y., 2021. Paleohyd ological dynamics in he Wes e n Medi e anean du ing
he las glacial cycle. Global Plane . Change 202, 103527.
Ga cía-Ju ado, F., Jim´
enez-G´
omez, F., Gue e o, F., 2011. E ec s o a d y pe iod on he
limnological cha ac e is ics o a Medi e anean high moun ain lake. Limn´
e ica 30,
5–16.
Ga cía-La uen e, J., Samma ino, S., Hue as, I.E., Flecha, S., S´
anchez-Leal, R.F.,
Na anjo, C., Nadal, I., Bellanco, M.J., 2021. Ho e and weake Medi e anean
ou low as a esponse o basin-wide al e a ions. F on . Ma . Sci. 8, 613444.
Gonz´
alez T ueba, J.J., Mo eno, R.M., Ma ínez De Pis´
on, E., Se ano, E., 2008. “Li le ice
age” glacia ion and cu en glacie s in he Ibe ian Peninsula. Holocene 18, 551–568.
G imm, E.C., 1987. CONISS: a Fo an 77 p og am o s a ig aphically cons ained
clus e analysis by he me hod o inc emen al sum o squa es. Compu . Geosci. 13,
13–35.
G imm, E.C., 1992. Tilia, Ve sion 2. Sp ing ield. IL 62703. Illinois S a e Museum.
Resea ch and Collec ion Cen e , USA.
Hamme , Ø., Ha pe , D.A.T., Ryan, P.D., 2001. PAST: Paleon ological s a is ics so wa e
package o Educa ion and da a analysis. Palaeon ol. Elec on. 4 (1), 9pp.
Hei i, O., Lo e , A.F., 2003. 9000 yea s o chi onomid assemblage dynamics in an alpine
lake: long- e m ends, sensi i i y o dis u bance, and esilience o he auna.
J. Paleolimnol. 30 (3), 273–289.
Hei i, O., Lo e , A.F., 2005. Holocene and La eglacial summe empe a u e
econs uc ion in he Swiss Alps based on ossil assemblages o aqua ic o ganisms: a
e iew. Bo eas 34, 506–516.
Hei i, O., Lo e , A.F., 2010. How does axonomic esolu ion a ec chi onomid-based
empe a u e econs uc ion? J. Paleolimnol. 44, 589–601.
Hei i, O., B ooks, S.J., Bi ks, H.J.B., Lo e , A.F., 2011. A 274-lake calib a ion da a-se
and in e ence model o chi onomid-based summe ai empe a u e econs uc ion
in Eu ope. Qua . Sci. Re . 30, 3445–3456.
Hei i, O., B ooks, S.J., Renssen, H., Bed o d, A., Hazekamp, M., Ilyashuk, B., Je e s, E.S.,
Lang, B., Ki ilo a, E., Kuipe , S., Mille , L., Sama in, S., To h, M., Ve b uggen, F.,
Wa son, J.E., an Asch, N., Lamme sma, E., Amon, L., Bi ks, H.H., Bi ks, H.J.B.,
Mo ensen, M.F., Hoek, W.Z., Magya i, E., Sob ino, C.M., Seppa, H., Tinne , W.,
Tonko , S., Veski, S., Lo e , A.F., 2014. Valida ion o clima e model-in e ed
egional empe a u e change o la e-glacial Eu ope. Na u e Communica ions, 2015.
In: Hei i, O., Ilyashuk, B., Mille , L., Sama in, S., Lo e , A.F. (Eds.), S acking o
Discon inuous Regional Palaeoclima e Reco ds: Chi onomid-Based Summe
Tempe a u es om he Alpine Region, ol. 5, p. 5914. Holocene 25(1), 137-149.
Hou, M., Wu, W., Cohen, D.J., Zeng, Z., Huang, H., Zheng, H., Ge, Q., 2023. De ec ion o
a mid-Holocene clima e e en a 7.2 ka BP based on an analysis o globally-
dis ibu ed mul i-p oxy eco ds. Palaeogeog . Palaeoclima ol. Palaeoecol. 618,
111525.
Ilyashuk, E.A., Koinig, K.A., Hei i, O., Ilyashuk, B.P., Psenne , R., 2011. Holocene
empe a u e a ia ions a a high-al i ude si e in he Eas e n Alps: a chi onomid
eco d om Schwa zsee ob S¨
olden, Aus ia. Qua . Sci. Re . 30, 176–191.
Ilyashuk, E.A., Hei i, O., Ilyashuk, B.P., Koinig, K.A., Psenne , R., 2019. The Li le Ice Age
signa u e in a 700-yea high- esolu ion chi onomid eco d o summe empe a u es
in he Cen al Eas e n Alps. Clim. Dynam. 52, 6953–6967.
IPCC, 2022. In: P¨
o ne , H.-O., Robe s, D.C., Tigno , M., Poloczanska, E.S.,
Min enbeck, K., Aleg ía, A., C aig, M., Langsdo , S., L¨
oschke, S., M¨
olle , V.,
Okem, A., Rama, B. (Eds.), Clima e Change 2022: Impac s, Adap a ion, and
Vulne abili y. Con ibu ion o Wo king G oup II o he Six h Assessmen Repo o
he In e go e nmen al Panel on Clima e Change. Camb idge Uni e si y P ess.
Jim´
enez, L., Ruhland, K.M., Jezio ski, A., Smol, J.P., P´
e ez-Ma ínez, C., 2018. Clima e
change and Saha an dus d i e ecen cladoce an and p ima y p oduc ion changes in
emo e alpine lakes o Sie a Ne ada, Spain. Global Change Biol. 28, 139–158.
Jim´
enez, L., Conde-Po cuna, J.M., Ga cía-Alix, A., Toney, J.L., Ande son, R.S., Hei i, O.,
P´
e ez-Ma ínez, C., 2019. Ecosys em esponses o clima e- ela ed changes in a
Medi e anean alpine en i onmen o e he las ~ 180 yea s. Ecosys ems 22,
563–577.
Jim´
enez-Espejo, F.J., Ma ínez-Ruiz, F., Roge son, M., Gonz´
alez-Donoso, J.M.,
Rome o, O., Lina es, D., Sakamo o, T., Gallego-To es, D., Rueda Ruiz, J.L., O ega-
Hue as, M., Pe ez Cla os, J.A., 2008. De i al inpu , p oduc i i y luc ua ions, and
w´
a e mass ci cula ion in he wes e nmos Medi e anean Sea since he Las Glacial
Maximum. Geochem. Geophy. Geosy. 9, Q11U02.
Jim´
enez-Espejo, F.J., Ga cía-Alix, A., Jim´
enez-Mo eno, G., Ma ínez-Ruiz, F.,
Ande son, R.S., Rod íguez-To a , F.J., Gi al , S., Rod igo-G´
amiz, M., Delgado
Hue as, A., Pa do-Igúzquiza, E., 2014. Saha an aeolian inpu and e ec i e humidi y
a ia ions o e Wes e n Eu ope du ing he Holocene. Chem. Geol. 374–375, 1–12.
Jim´
enez-Mo eno, G., Ande son, R.S., 2012. Holocene ege a ion and clima e change
eco ded in alpine bog sedimen s om he Bo eguiles de la Vi gen, Sie a Ne ada,
sou he n Spain. Qua . Res. 77, 44–53.
Jim´
enez-Mo eno, G., Ga cía-Alix, A., He n´
andez-Co bal´
an, M.D., Ande son, R.S.,
Delgado-Hue as, A., 2013. Vege a ion, i e, clima e and human dis u bance his o y
in he sou hwes e n Medi e anean a ea du ing he la e Holocene. Qua . Res. 79,
110–122.
Jim´
enez-Mo eno, G., Ga cía-Alix, A., Ande son, R.S., Ramos-Rom´
an, M.J., Camue a, J.,
Mesa-Fe n´
andez, J.M., Toney, J.L., Jim´
enez-Espejo, F.J., Ca i´
on, J.S., L´
opez-
A il´
es, A., Rod igo-G´
amiz, M., Webs e , C.E., 2022. Recons uc ion o pas
en i onmen and clima e using We land sedimen eco ds om he Sie a Ne ada.
In: Zamo a, R., Oli a, M. (Eds.), The Landscape o he Sie a Ne ada. Sp inge
Na u e Swi ze land AG, pp. 95–114.
Joannin, S., B ugiapaglia, E., de Beaulieu, J.-L., Be na do, L., Magny, M., Pey on, O.,
Go ing, S., Vanni`
e e, B., 2012. Pollen-based econs uc ion o Holocene ege a ion
and clima e in sou he n I aly: he case o Lago T i oglie i. Clim. Pas 8, 1973–1996.
Juggins, S., 2007. C2 Ve sion 1.5: So wa e o Ecological and Palaeoecological Da a
Analysis and Visualiza ion. Newcas le Uni e si y, Newcas le upon Tyne, UK.
Kau man, D.S., Age , T.A., Ande son, N.J., Ande son, P.M., And ews, J.T., Ba lein, P.J.,
B ubake , L.B., Coa s, L.L., Cwyna , L.C., Du all, M.L., Dyke, A.S., Edwa ds, M.E.,
Eisne , W.R., Gajewski, K., Gei sdo i , A., Hu, F.S., Jennings, A.E., Kaplan, M.R.,
Ke win, M.N., Lozhkin, A.V., MacDonald, G.M., Mille , G.H., Mock, C.J., Oswald, W.
W., O o-Bliesne , B.L., Po inchu, D.F., Ruhland, K., Smol, J.P., S eig, E.J., Wol e, B.
B., 2004. Holocene he mal maximum in he wes e n A c ic(0-180 deg ees W). Qua .
Sci. Re . 23, 529–560.
Kau man, D., McKay, N., Rou son, C., E b, M., D¨
a wyle , C., Somme , P.S., Hei i, O.,
Da is, B., 2020a. Holocene global mean su ace empe a u e, a mul i-me hod
econs uc ion app oach. Sci. Da a 7, 201.
Kau man, D., McKay, N., Rou son, C., E b, M., Da is, B., Hei i, O., Jacca d, S., Tie ney, J.,
D¨
a wyle , C., e al., 2020b. A global da abase o Holocene paleo- empe a u e
eco ds. Sci. Da a 7, 115.
Ko hola, A., Wecks ¨
om, J., Holms ¨
om, L., E ¨
as ¨
o, P., 2000. A quan i a i e Holocene
clima ic eco d om dia oms in no he n Fennoscandia. Qua . Res. 54, 284–294.
La ocque, I., Hall, R.I., G ahn, E., 2001. Chi onomids as indica o s o clima e change: a
100-lake aining se om a suba c ic egion o no he n Sweden (Lapland).
J. Paleolimnol. 26, 307–322.
Laska , J., Robu el, P., Jou el, F., Gas ineau, M., Co eia, A.C.M., Le a d, B., 2004.
A long- e m nume ical solu ion o he insola ion quan i ies o he Ea h. As on.
As ophys. 428, 261–285.
Legend e, P., Bi ks, H.J.H., 2012. F om classical o canonical o dina ion. In: Bi ks, H.J.B.,
Lo e , A.F., Juggins, S., Smol, J.P. (Eds.), T acking En i onmen al Change Using
Lake Sedimen s Volume 7. Sp inge , Do d ech , The Ne he lands.
Lepˇ
s, J., ˇ
Smilaue , P., 2003. Mul i a ia e Analysis o Ecological Da a Using Canoco.
Camb idge Uni e si y P ess, Camb idge.
L´
opez-A il´
es, A., Jim´
enez-Mo eno, G., Ga cía-Alix, A., Ga cía-Ga cía, F., Camue a, J.,
Ande son, R.S., Sanju jo-S´
anchez, J., Chamo o, C.A., Ca i´
on, J.S., 2022. Pos -
glacial e olu ion o alpine en i onmen s in he wes e n Medi e anean egion: he
Laguna Seca eco d. Ca ena 211, 106033.
L´
opez-A il´
es, A., Ga cía-Alix, A., Jim´
enez-Mo eno, G., Ande son, R.S., Toney, J.L., Mesa-
Fe n´
andez, J.M., Jim´
enez-Espejo, F.J., 2021. La es Holocene paleon i onmen al and
paleoclima e econs uc ion om an alpine bog in he Wes e n Medi e anean
egion: he Bo eguil de los La ade os de la Reina eco d (Sie a Ne ada).
Palaeogeog . Palaeoclima ol. Palaeoecol. 573, 110434.
Lo enzo, M.N., Al a ez, I., 2022. Fu u e changes o ho ex emes in Spain: owa ds
wa me condi ions. Na . Haza ds 113, 383–402.
Luo o, T.P., 2009. Sub ossil Chi onomidae (Insec a: Dip e a) along a la i udinal g adien
in Finland: de elopmen o a new empe a u e in e ence model. J. Qua . Sci. 24,
150–158.
Ma ín-Pue as, C., Vale o-Ga c´
es, B.L., B aue , A., Ma a, M.P., Delgado-Hue as, A.,
Dulski, P., 2009. The Ibe ian-Roman Humid Pe iod (2600-1600 cal y BP) in he
Zo˜
na lake a e eco d (Andalucía, sou he n Spain). Qua . Res. 71, 108–120.
Ma ín-Pue as, C., Ma hes, K., B aue , A., Muschele , A., Hansen, F., Pe ick, C.,
Aldahan, A., Possne , G., an Geel, B., 2012. Regional a mosphe ic ci cula ion shi s
induced by a g and sola minimum. Na . Geosci. 5, 397–401.
Mesa-Fe n´
andez, J.M., Jim´
enez-Mo eno, G., Rod igo-G´
amiz, M., Ga cía-Alix, A.,
Jim´
enez-Espejo, F.J., Ma ínez-Ruiz, F., Sco Ande son, R.S., Camue a, J., Ramos-
Rom´
an, M.J., 2018. Vege a ion and geochemical esponse o Holocene apid clima e
change in he Sie a Ne ada (sou heas e n Ibe ia): he Laguna Honde a eco d. Clim.
Pas 14, 1687–1706.
Mo ales-Baque o, R., Ca illo, P., Reche, I., Sanchez-Cas illo, P., 1999.
Ni ogenephospho us ela ionship in high moun ain lakes: e ec s o he size o
ca chmen basins. Can. J. Fish. Aqua . Sci. 56, 1809–1817.
Mo ales-Baque o, R., Conde-Po cuna, J.M., 2000. E ec o he ca chmen a eas on he
abundance o zooplank on in high moun ain lakes o Sie a Ne ada (Spain).
Ve handlungen de In e na ionale Ve einigung ü heo e ische und angewand e
Limnologie 27, 1–5.
Mo cillo-Mon alb´
a, L., Rod igo-G´
amiz, M., Ma ínez-Ruiz, F., O ega-Hue as, M.,
Schou en, S., Sinninghe Dams ´
e, J.S., 2021. Rapid clima e changes in he
wes e nmos Medi e anean (Albo an Sea) o e he las 35 ky : new insigh s om
ou lipid paleo he mome e s (UK’ 37, TEXH 86, RI-OH’, and LDI). Paleoceanog .
Paleoclima ol. 36, e2020PA004171.
Mo eno, A., P´
e ez, A., F igola, J., Nie o-Mo eno, V., Rod igo-G´
amiz, M., Ma a , B.,
Gonz´
alez-Samp´
e iz, P., Mo ell´
on, M., Ma ín-Pue as, C., Co ella, J.P., Belmon e, ´
A.,
Sancho, C., Cacho, I., He e a, G., Canals, M., G imal , J.O., Jim´
enez-Espejo, F.,
Ma ínez-Ruiz, F., Vegas-Vila úbia, T., Vale o-Ga c´
es, B.L., 2012. The medie al
clima e Anomaly in he Ibe ian Peninsula econs uc ed om ma ine and lake
eco ds. Qua . Sci. Re . 43, 16–32, 2012.
Obse a o io del cambio, 2020. Global de Sie a Ne ada. In: Lina ia 1.0. iEcolab e
Labo a o io de Ecología Te es e e Uni e sidad de G anada. h p://lina ia.obsne .
es.
Oli a, M., G´
omez-O iz, A., 2012. La e-Holocene en i onmen al dynamics and clima e
a iabili y in a Medi e anean high moun ain en i onmen (Sie a Ne ada, Spain)
in e ed om lake sedimen s and his o ical sou ces. Holocene 22 (8), 915–927.
Oli a, M., G´
omez-O iz, A., Palacios, D., Sal ado -F anch, F., And ´
es, N., Tana o, L.N.,
Fe n´
andez-Fe n´
andez, J.M., Ba iocanal, C., 2020. Mul ip oxy econs uc ion o
Holocene glacie s in Sie a Ne ada (sou h Spain). Medi . Geosci. Re . 2, 5–19.
G. Jim´
enez-Mo eno e al.

Qua e na y Science Re iews 319 (2023) 108343
17
PAGES 2k-PMIP3 g oup, 2015. Con inen al-scale empe a u e a iabili y in PMIP3
simula ions and PAGES 2k egional empe a u e econs uc ions o e he pas
millennium. Clim. Pas 11, 1673–1699.
Palacios, D., G´
omez-O iz, And ´
es, N., Sal ado , F., Oli a, M., 2016. Timing and new
geomo phologic e idence o he las deglacia ion s ages in Sie a Ne ada (sou he n
Spain). Qua . Sci. Re . 150, 110–129.
Palacios, D., Oli a, M., G´
omez-O iz, A., And ´
es, N., Fe n´
andez-Fe n´
andez, J.M.,
Schimmelp ennig, I., e al., 2020. Clima e sensi i i y and geomo phological esponse
o ci que glacie s om he la e glacial o he Holocene, Sie a Ne ada, Spain. Qua .
Sci. Re . 248, 106617.
Pędziszewska, A., Tylmann, W., Wi ak, M., Pio owska, N., Maciejewska, E.,
La ałowa, M., 2015. Holocene en i onmen al changes e lec ed by pollen, dia oms,
and geochemis y o annually lamina ed sedimen s o Lake Suminko in he
Kashubian Lake Dis ic (N Poland). Re . Palaeobo . Palynol. 216, 55–75.
Pepin, N.C., Lundquis , J.D., 2008. Tempe a u e ends a high ele a ions: pa e ns ac oss
he globe. Geophys. Res. Le . 35 (14), L14701.
Pepin, N.C., A none, E., Gobie , A., Haslinge , K., Ko la ski, S., No a nicola, C., e al.,
2022. Clima e changes and hei ele a ional pa e ns in he moun ains o he wo ld.
Re . Geophys. 60, e2020RG000730.
Ramos-Rom´
an, M.J., Jim´
enez-Mo eno, G., Ande son, R.S., Ga cía-Alix, A., Toney, J.L.,
Jim´
enez-Espejo, F.J., Ca i´
on, J.S., 2016. Cen ennial-scale ege a ion and No h
A lan ic oscilla ion changes du ing he la e Holocene in he wes e n medi e anean.
Qua . Sci. Re . 143, 84–95.
Ramos-Rom´
an, M.J., Jim´
enez-Mo eno, G., Camue a, J., Ga cía-Alix, A., Ande son, R.S.,
Jim´
enez-Espejo, F.J., Sachse, D., Toney, J.L., Ca i´
on, J.S., Webs e , C., Yanes, Y.,
2018. Millennial-scale cyclical en i onmen and clima e a iabili y du ing he
Holocene in he wes e n Medi e anean egion deduced om a new mul ip oxy
analysis om he Padul eco d (Sie a Ne ada, Spain). Global Plane . Change 168,
35–53.
Raposei o, M.P., Saez, A., Gi al , S., Cos a, A.C., Gonçal es, V., 2018. Causes o spa ial
dis ibu ion o sub ossil dia om and chi onomid assemblages in su ace sedimen s o
a emo e deep island lake. Hyd obiol. (So ia) 815, 141–163.
Reche, I., Pulido-Villena, E., Mo ales-Baque o, R., Casamayo , E.O., 2005. Does
ecosys em size de e mine aqua ic bac e ial ichness? Ecology 86, 1715–1722.
Reime , P., Aus in, W., Ba d, E., Bayliss, A., Blackwell, P., B onk Ramsey, C., Talamo, S.,
2020. The In Cal20 no he n hemisphe e adioca bon age calib a ion cu e (0–55
cal kBP). Radioca bon 62 (4), 725–757.
Renssen, H., Sepp¨
a, H., Hei i, O., Roche, D.M., Goosse, H., Fiche e , T., 2009. The spa ial
and empo al complexi y o he Holocene he mal maximum. Na u e Geosci. 2,
411–414.
Rie ade all, M., Bonada, N., P a , N., 1999. Subs a e and Dep h p e e encs o
mac oin e eb a es along a ansec in a Py enean high moun ain lake (Lake Red´
o,
NE Spain). Limn´
e ica 17, 127–134.
Rod igo-G´
amiz, M., Ga cía-Alix, A., Jim´
enez-Mo eno, G., Ramos-Rom´
an, M.J.,
Camue a, J., Toney, J.L., Sachse, D., Ande son, R.S., Sinninghe Dams ´
e, J.S., 2022.
Paleoclima e econs uc ion o he las 36 ky based on b anched glyce ol dialkyl
glyce ol e ae he s in he Padul palaeolake eco d (Sie a Ne ada, sou he n Ibe ian
Peninsula). Qua . Sci. Re . 281, 107434.
Sama in, S., Hei i, O., Joos, F., Renssen, H., F anke, J., B ¨
onnimann, S., Tinne , W.,
2017. Wa m Medi e anean mid-Holocene summe s in e ed om ossil midge
assemblages. Na . Geosci. 10, 207–212.
S´
anchez-Cas illo, P.M., C uz-Piza o, L., Ca illo, P., 1989. Ca ac e izaci´
on del
i oplanc on de las lagunas de al a mon a˜
na de Sie a Ne ada (G anada, Espa˜
na) en
elaci´
on a las ca ac e ís icas ísico-químicas del medio. Limn´
e ica 5, 37–50.
Sche e , S.C., Ceppi, P., C oci-Maspoli, M., Appenzelle , C., 2012. Snow-albedo eedback
and Swiss sp ing empe a u e ends. Theo . Appl. Clima ol. 110 (4), 509–516.
Sic e, M.-A., Jalali, B., Ma a , B., Schmid , S., Basse i, M.-A., Kallel, N., 2016. Sea
su ace empe a u e a iabili y in he No h wes e n medi e anean Sea (Gul o
lion) du ing he common E a. Ea h Plane Sci. Le . 456, 124–133.
Sie o, F.J., Hodell, D.A., Ande sen, N., Azibei o, L.A., Jimenez-Espejo, F.J., Bah , A.,
e al., 2020. Medi e anean o e low o e he las 250 ky : F eshwa e o cing om
he opics o he ice shee s. Paleoceanog . Paleoclima ol. 35, e2020PA003931.
Spanish Na ional Wea he Agency, 2022. AEMe Open Da a. AEMe Open Da a. h p://
www.aeme .es/es/da os_abie os/AEMET_OpenDa a.
Ta a s, P., Hei i, O., Vale o-Ga c´
es, B., Ca˜
nedo-A güelles, M., P a , N., Rie ade all, M.,
Gonz´
alez-Samp´
e iz, P., 2018. Chi onomid-in e ed Holocene empe a u e
econs uc ion in Basa de la Mo a Lake (Cen al Py enees). Holocene 28 (11),
1685–1696.
Tel o d, R.J., Bi ks, H.J.B., 2005. The sec e assump ion o ans e unc ions: p oblems
wi h spa ial au oco ela ion in e alua ing model pe o mance. Qua . Sci. Re . 24,
2173–2179.
Te B aak, C.J.F., Juggins, S., 1993. Weigh ed a e aging pa ial leas squa es eg ession
(WA-PLS): an imp o ed me hod o econs uc ing en i onmen al a iables om
species assemblages. Hyd obiol 269, 485–502.
Te B aak, C.J.F., Juggins, S., Bi ks, H.J.B., an de Voe , H., 1993. Weigh ed a e aging
pa ial leas squa es eg ession (WA-PLS): de ini ion and compa ison wi h o he
me hods o species-en i onmen calib a ion. Chap e 25. In: Pa il, G.P., Rao, C.R.
(Eds.), Mul i a ia e En i onmen al S a is ics, No h-Holland, Ams e dam.
Toney, J.L., Ga cía-Alix, A., Jim´
enez-Mo eno, G., Ande son, R.S., Moossen, H., Seki, O.,
2020. New insigh s in o Holocene hyd ology and empe a u e om lipid bioma ke s
in wes e n Medi e anean alpine we lands. Qua . Sci. Re . 240, 106395.
To h, M., Magya i, E.K., Buczko, K., B aun, M., Panagio opoulos, K., Hei i, O., 2015.
Chi onomid-in e ed Holocene empe a u e changes in he Sou h Ca pa hians
(Romania). Holocene 25 (4), 569–582.
Valle, F., 2003. Mapa de Se ies de Vege aci´
on de Andalucía. Edi o ial Rueda, S.L.,
Mad id.
Va gas-Y´
a˜
nez, M., Mo a, F., Ga cía-Ma ínez, M.C., Tel, E., Zunino, P., Plaza, F., Sala , J.,
Pascual, J., L´
opez-Ju ado, J.L., Se a, M., 2010. Clima e change in he wes e n
Medi e anean Sea 1900-2008. J. Ma . Sys . 82, 171–176.
Walke , I.R., 2001. Midges: Chi onomidae and ela ed Dip e a. In: Smol, J.P., Bi ks, H.J.
B., Las , W.M. (Eds.), T acking En i onmen al Change Using Lake Sedimen s.
Volume 4: Zoological Indica o s. Kluwe Academic Publishe s, Do d ech , pp. 43–66.
G. Jim´
enez-Mo eno e al.