Mammalian turnover as an indicator of climatic and anthropogenic landscape modification: A new Meghalayan record (Late Holocene) in northern Iberia

Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
A ailable online 1 Ma ch 2023
0031-0182/© 2023 The Au ho s. Published by Else ie B.V. This is an open access a icle unde he CC BY license (h p://c ea i ecommons.o g/licenses/by/4.0/).
Mammalian u no e as an indica o o clima ic and an h opogenic
landscape modi ica ion: A new Meghalayan eco d (La e Holocene) in
no he n Ibe ia
Ad i´
an ´
Al a ez-Vena
a
,
*
, Ana B. Ma ín-A oyo
b
, Diego J. ´
Al a ez-Lao
a
, C´
esa Laplana
c
,
Ma ín A iolabengoa
d
, Daniel Balles e os
e
, A an za A anbu u
d
, Pe u Bilbao
d
, ´
Angel As o qui
,
Yolanda Díaz-Casado
a
Depa men o Geology, Uni e si y o O iedo, C/ Jesús A ias de Velasco s/n, 33005 O iedo, Spain
b
E oAdap a G oup, Uni e sidad de Can ab ia, 39005 San ande , Spain
c
Museo A queol´
ogico y Paleon ol´
ogico de la Comunidad de Mad id, Pza. Be na das s/n, 28801, Alcal´
a de Hena es, Mad id, Spain
d
Depa men o Mine alogy and Pe ology, Uni e si y o he Basque Coun y, UPV/EHU, Leioa, Spain
e
Depa men o Geodynamics, Uni e si y o G anada, Campus de Fuen enue a s/n, G anada, Spain
Tanea Documen aci´
on y Conse aci´
on SL, C/ Juan Jos´
e P´
e ez del Molino 16, 39006 San ande , Spain
ARTICLE INFO
Edi o : D . Howa d Falcon-Lang
Keywo ds:
Clima ic change
Chalcoli hic
B onze Age
I on Age
Mic omys minu us
Mus musculus
ABSTRACT
The Pun a Luce o III ca e is a na u al ap whe e abundan e eb a e emains we e accumula ed du ing he
Meghalayan (La e Holocene). To be e unde s and he paleoen i onmen al condi ions in which his eco d was
accumula ed, he mic omammal assemblage, comp ising a minimum numbe o 1396 indi iduals belonging o
19 axa, was s udied using he Mu ual Ecogeog aphic Range and he Habi a Weigh ing Me hod. Th oughou
~2600 yea s, he mic omammal communi y's quick u no e e lec ed a shi om pa chy o es s and humid
meadows o open, sh ubbie g asslands. The La e Holocene The mal Maximum's humid and mild clima ic con-
di ions unde wen a cooling and a idi ica ion phase, coe al wi h he I on Age Cold Epoch. These concluded in a
sligh empe a u e ising, coe al wi h he Roman Wa m Pe iod. Mac omammals expe ienced a shi om wild
popula ions o domes ic he ds. The e o e, his wo k discusses a b oade con ex o his mammalian u no e
om a human cul u al pe spec i e.
1. In oduc ion
The Holocene was conside ed a ela i ely wa m and s able epoch
compa ed o he las glacial pe iod (Dansgaa d e al., 1993). Ne e he-
less, ab up clima ic changes and cooling e en s iden i ied h oughou
he Holocene (Alley e al., 1997; Bond e al., 1997; Mayewski e al.,
2004; Walke e al., 2018), e en i weake han hose o he Las Glacial
Cycle, ha e been in ense enough o cause he ise and collapse o sig-
ni ican cul u es and ci ilisa ions wo ldwide (Van Geel e al., 1996;
deMenocal, 2001; Bün gen e al., 2011).
P incipal wa ming o cooling ends a millennial imescales a e
caused by o bi al o cing (deMenocal e al., 2000; Mayewski e al., 2004;
Wanne e al., 2008) in addi ion o luc ua ions in o al sola i adiance
and peaks o olcanic ac i i y, which a e also ela ed o ab up clima e
shi s (B ay, 1971; Van Geel e al., 1996; Bond e al., 2001; Mayewski
e al., 2004; Wanne e al., 2008; S einhilbe e al., 2009; Mille e al.,
2012; Sigl e al., 2015; Bo zenko a e al., 2015). Howe e , since he end
o he Pleis ocene and, especially du ing he Holocene, human ac i i y
has become an inc easing o ce shaping he e es ial and ma ine eco-
sys ems (Vi ousek e al., 1997; C u zen, 2002; Dough y e al., 2010; Ellis,
2011; Ba nosky, 2008, 2013; Boi in e al., 2016), causing a chemical
inge p in , bo h on a egional and global scale (Ruddiman and Thom-
son, 2001). F om Mid-Holocene onwa ds, human ac i i ies such as an-
imal husband y, ag icul u e, and me allu gy, ha e had a signi ican
impac on he landscapes o he egions inhabi ed by humans, including
he Ibe ian Peninsula, making an h opogenic and clima ic signals o en
indis inguishable in pollen eco ds (Ca i´
on e al., 2010). Due o his
clima ic and an h opogenic in luence, mammals ha e expe ienced sig-
ni ican ange shi s, ex inc ion e en s, ex i pa ions, and domes ica ion
p ocesses (Roseng en e al., 2021). In his sense, he Holocene wa ming
* Co esponding au ho .
E-mail add ess: [email p o ec ed] (A. ´
Al a ez-Vena).
Con en s lis s a ailable a ScienceDi ec
Palaeogeog aphy, Palaeoclima ology, Palaeoecology
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h ps://doi.o g/10.1016/j.palaeo.2023.111476
Recei ed 15 Janua y 2023; Recei ed in e ised o m 23 Feb ua y 2023; Accep ed 24 Feb ua y 2023
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
2
and he de elopmen o human cul u es also led o he u no e o he
Ibe ian mammal communi ies. While domes ic he ds p og essi ely
eplaced wild mac omammals, mic omammal assemblages, whose
geog aphical dis ibu ion was a ec ed by habi a changes (clima ic o
human-d i en), also e lec ed he a i al o alloch honous commensal
species (Al una, 1980; Cuenca-Besc´
os e al., 2009; L´
opez-Ga cía e al.,
2013; Ba˜
nuls-Ca dona e al., 2017a; Domínguez Ga cía e al., 2019;
Domínguez Ga cía e al., 2020; Domínguez-Ga cía e al., 2022; Mocl´
an
e al., 2023). Fo his eason, mac o- and, pa icula ly, mic omammals
a e a aluable sou ce o paleoen i onmen al in o ma ion ha has been
success ully analysed om di e se pe spec i es a key si es o he Can-
ab ian Region (e.g., Al una, 1980; L´
opez-Ga cía, 2008; Cuenca-Besc´
os
e al., 2009, 2012; ´
Al a ez-Lao, 2014; Balles e os e al., 2020;
´
Al a ez-Vena e al., 2021; Jones e al., 2019, 2021).
Ibe ian mac omammal assemblages om Holocene si es a e mainly
associa ed wi h a chaeological con ex s (Al una, 1980; Ma iezku ena,
1990; Cas a˜
nos, 1997; Al una and Ma iezku ena, 2012; Vega-Maeso
e al., 2016). Consequen ly, hey a e biased by an h opogenic ac i i y.
On he o he hand, mic omammal accumula ions usually esul om he
p eda o y ac i i y o ap o s o mammalian ca ni o es (And ews, 1990;
Ga cía-Mo a o e al., 2023). Al hough hese emains can be ound in
assemblages ha do no necessa ily in ol e p eda ion (as in he case o
na u al aps, e.g., ´
Al a ez-Lao e al., 2020), hese kinds o si es a e
sca ce in he li e a u e. In ou s udy case, acciden al alling and
noc u nal bi ds o p ey a e he mos likely accumula ing agen s o he
mic omammals' assemblage. S ill, acco ding o he gene alis habi s o
owls, hey accu a ely ep esen he en i onmen in which hey hun
(And ews, 1990). In his con ex , he si e p esen ed he e is o g ea
ele ance as i yielded a Holocene mammal associa ion mainly o med
in a na u al pi all ap (a leas in he case o mac omammals), i.e.,
wi hou bias by humans o specialised p eda o s, o e ing an excep ional
oppo uni y o unde s and he clima ic change and human-en i onmen
in e ac ions du ing he Meghalayan (La e Holocene) a he Can ab ian
Region.
2. The Pun a Luce o III si e
2.1. Geog aphic con ex
The Pun a Luce o III si e (PL-III) is loca ed a he eas e n end o he
Can ab ian Region, a s ip o land (30–50 km wide) wi h ugged elie
ha ex ends along he no h o he Ibe ian Peninsula. I is a ka s sha
( e ical ca e) si ua ed on he no he n slope o he Pun a Luce o moun
(43◦21′34.44”N, 3◦6′10.31”W, 240 m asl; Fig. 1) in he municipali y o
Zie bena/Cie ana (Bizkaia, Basque Coun y, no he n Spain). This
moun ain lanks he wes bank o he Ne i´
on Ri e mou h, whe e he
Bilbao Po has been buil (Fig. 1C-D). The si e o med wi hin he Lowe
C e aceous limes one (calca eni e), sands one and shale (Ga o e e al.,
1993) and o ms pa o a ka s coas al a ea de eloped in a adose
en i onmen du ing he Qua e na y (A anbu u e al., 2015). Cu en ly,
he ca e is loca ed a he highes poin o a qua y, whose exploi a ion
began du ing he cons uc ion o he Bilbao seapo . Howe e , be o e
hese wo ks, i was loca ed hal way up he slope, be ween he op o he
moun and he Can ab ian Sea (Fig. 1E). Nowadays, his e i o y is pa
o he Eu o-Sibe ian biogeog aphic egion. The si e is in he A lan ic
clima ic domain, cha ac e ised by empe a e summe s, mild win e s,
and no d y season ( ype C b acco ding o he K¨
oppen-Geige classi ica-
ion; Beck e al., 2018). The mean annual empe a u e (MAT) a Zie -
bena is 13.9 ◦C, wi h maximum and minimum mon h-mean
empe a u es o 19.8 and 8.7 ◦C, espec i ely. P ecipi a ion is dis ib-
u ed h oughou he yea , eaching an annual mean o ~1150 mm
(Cou o e al., 2011). Howe e , he plan communi ies in his a ea ha e
30% o Medi e anean species (Pa ino e al., 2002). Some o hese axa
expanded due o human ac i i ies such as he epea ed use o i e o
ob ain pas u e, as is he case o he holm oak (Que cus ilex) and he
Ke mes oak (Que cus cocci e a), adap ed o li e in mo e ex eme
condi ions han he A lan ic species (Pa ino e al., 2002).
Di ec e idence o he human p esence in Pun a Luce o da es o imes
e en be o e he o ma ion o he s udied sequence, as e idenced by he
human emains (Homo sapiens bones) ound a moun idge by G´
omez-
Oli encia e al. (2015) in he si es o Pun a Luce o II (5566–5318 cal y
BP) and Co ach´
on III (4796–4424 cal y BP), a ~200 and ~ 500 m om
he PL-III sha espec i ely. In he nea by si e o Pico Ramos, i has also
been ound a sepulch al con ex o Chalcoli hic age (5862–4295 cal y
BP) con aining human skele al emains o a minimum o 104 indi iduals
(Ba ayba and de la Rúa, 1995; Zapa a, 1995). The palynological s udy
o Pico Ramos Chalcoli hic bu ials depic ed an open landscape wi h
signals o an h opisa ion, in which a mixed-oak g o e o ma ion o
empe a e and humid cha ac e was de eloped (I ia e, 1994).
2.2. S a ig aphy
Be o e he qua y exploi a ion, he s udied sequence consis ed o a
ca e in ill (Fig. 2) whose en ance was abo e he cu en le el. The
homogeneous sedimen a y ill, mainly consis ing o la ge limes one
clas s and loamy sedimen , was sampled a di e en dep h in e als
om i s op owa ds i s base, some imes coinciding wi h s a ig aphic
le els o con as ing cha ac e is ics and, on o he s, wi hin mo e hick
and homogeneous se s. Due o he na ow exca a ion a ea, i was
impossible o p ese e a s a ig aphic p o ile, so we di ided he
sequence in o di e en s a ig aphic sec ions each ime we ex ac ed a
g oup o la ge limes one blocks so ha bone samples could be g ouped.
Sec ions o le els we e labelled F o A, om bo om o op o he p e-
se ed sequence. The dep h was measu ed om he op o he emnan
deposi , pa ially exposed a e he las blas ing wo ks in he qua y. To
iden i y di e ences in he sedimen a y sequence, pe og aphic and
mine alogical analyses we e made in s a ig aphic le els F, D, and C.
O e all, hey disclose ma ix-suppo ed mic o acies, wi h ine o me-
dium sub-angula qua z g ains, mainly o sedimen a y ocks bu also
ew che and me amo phic qua z (Appendix 1). Le el F is a he bo om
o he sequence, lying o e he unexca a ed deposi . This le el was
es ablished by i s high mic ocha coal con en in g eyish sedimen . Le el
F was unde a big limes one block; sedimen con aining his block was
named Le el E. Le el D was de e mined a e inding he i s ce id
emains. Le el C was labelled a e he exca a ion o Le el B, which was
an uns able in ill. Samples collec ed du ing he su ace cleaning and he
emo al o isola ed bones be o e he exca a ion o Le el B we e g ouped
in Le el A since mos o hem ell om he la e al o abo e. The e o e,
Le el A mus be conside ed wi h cau ion as a ewo ked sample.
3. Ma e ials and me hods
The mammal emains s udied in his wo k come om he escue
exca a ions in he Pun a Luce o qua y (Fig. 1D; Fig. 2) in Sep embe
2019. The s abilisa ion blas ing a he qua y was supposed o des oy
he ca e. Howe e , his did no happen, so he emaining s a ig aphic
sequence is s ill p ese ed a he si e.
3.1. S a ig aphic s udy
The s a ig aphic sec ion o he PL-III si e was di ided in o le els
based on he na u e, colou , g anulome y, cohesi i y, sedimen a y
s uc u es, and ela i e abundance o paleon ological emains (e.g.,
A iolabengoa e al., 2015). The op o he PL-III si e was des oyed by
qua ying; hus, le el dep h was measu ed conside ing he exca a ion
su ace in Sep embe 2019. The s a ig aphic sec ion was complemen ed
by mic oscopical obse a ions o unconsolida ed sedimen s and X-Ray
Di ac ion (XRD) analysis o cha ac e ise mine alogy and mic o acies
and o in e he sedimen p o enance. See Appendix 1 o u he de ails.
A. ´
Al a ez-Vena e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
3
Fig. 1. Geog aphical con ex o he Pun a Luce o III si e. (A) Loca ion o he Can ab ian Region (B) in he Ibe ian Peninsula. (B) O he si es men ioned in his wo k:
1 Valda a a-1; 2 Mon e A eo mi e; 3 El Oli o; 4 La Güelga; 5 To ca del Le´
on; 6 El Espinoso; 7 El Se al; 8 Cue o la A ellanosa; 9 Cue a del Cob e; 10 La Molina
pea bog; 11 La Ga ma; 12 Kai e; 13 El Mi ´
on; 14 Cue a Mayo ; 15 Mi ado ; 16 Zalama pea bog; 17 Pico Ramos; 18 A enaza; 19 Pun a Luce o-II and Co ach´
on-III; 20
Kobaede a; 21 Amalda; 22 He iko Ba a. (C) Hypsome ic backg ound wi h 100 m al i ude in e als o he Pun a Luce o su oundings. (D–E) O hopho og aphy o
he Pun a Luce o su oundings: (D) a e he s abilisa ion wo ks in he qua y ha allowed he disco e y o he Pun a Luce o sha , and (E) be o e he cons uc ion o
he Bilbao Po . Ca og aphy and o hopho og aphy om he geoEuskadi online GIS- iewe (Eusko Jau la i za/Gobie no Vasco).
A. ´
Al a ez-Vena e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
4
3.2. Da ing
Th ee bone samples and wo cha coals we e selec ed o AMS
adioca bon da ing and sen o he Be a Analy ic labo a o y (Miami,
USA). Ob ained ages and published da es discussed in his wo k we e
calib a ed h ough he In Cal20 da abase (Reime e al., 2020) in OxCal
.4.4.4 (B onk Ramsey, 2009), conside ing he 2
σ
s anda d de ia ion
(95.4% p obabili y).
Fig. 2. Cha ac e isa ion o he Pun a Luce o III exca a ion. (A) O hopho o o he exca a ion a ea. The p ojec ion o he sec ions in b and c is indica ed. (B) A–B
c oss-sec ion o he sha . (C) C–D c oss-sec ion o he sha . (D) S a ig aphic p o ile o he sha in ill. Small mammal NISPi omi s he Lepus emains. Pho o-
g amme ic econs uc ions o backg ounds om sec ions o B–D p o ided by GIM-Geoma ics.
A. ´
Al a ez-Vena e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
5
3.3. Collec ing and so ing
3.3.1. Mic omammals
A o al o 24 samples o sedimen (~10 kg each) we e collec ed and
wa e -sc eened using wo supe imposed sie es o 0.5 and 2 mm. The
ma e ials in he ine ac ion (0.5–2 mm) we e so ed by checking he
samples wi h a binocula mic oscope unde 10×magni ica ion. The
ob ained bone emains we e hen classi ied and s udied a he Geology
Depa men o he Uni e si y o O iedo (Spain).
Taxonomic iden i ica ions we e ca ied ou employing a s e eomi-
c oscope Nikon SMZ800N, equipped wi h a 16 Mpx digi al came a.
Measu emen s we e aken on he mic oscope pho og aphs using Adobe
Pho oshop CC so wa e. Due o hei diagnos ic alue and po en ial o
p ese a ion, he ana omical elemen s selec ed o axonomic iden i i-
ca ions we e he ollowing: p3 and pos c anial skele on o lagomo phs;
isola ed mola s o mu ids; m1 o a icolines (also conside ing he M3
o he Te icola subgenus, and he M1 and M2 o he species belonging
o he Mic o us (Ag icola) ag es is g oup); mandibles and maxillae o
so icids and e inaceids; and uppe mola s and hume i o alpids. A
compa a i e collec ion o mic omammal skele al emains ob ained om
ecen ba n owl pelle s was also used.
Ha es we e iden i ied ollowing Palacios and L´
opez Ma ínez (1980),
Llo en e (2010), and Pelle ie (2018). The gene al iden i ica ion o o-
den s and sh ews ollowed Rom´
an (2019) and No es (1989), espec-
i ely. De ailed axonomical and mo phological analyses we e based on:
K app and Nie hamme (1982), Nadachowski (1984), and Luzi and
L´
opez-Ga cía (2019) o Mic o us (Ag icola) ex g . ag es is (which in-
cludes he species Mi. (Ag.) la e nedii, Mi. (Ag.) ozianus and Mi. (Ag.)
ag es is) and Mic o us (Mic o us) a alis lowe den i ion; No es e al.
(1982), Pem´
an (1983) and Ba i (2006) o Neomys mandibles; Pasquie
(1974), No es (1988) and Kni lo ´
a and Ho ´
aˇ
cek (2017) o Apodemus
(Syl aemus) den i ion; Da iche e al. (2006) o Mus; and Nie hamme
(1990) and Gu i´
e ez e al. (2019) o Talpa hume i and uppe mola s
espec i ely. The axonomic classi ica ion ollowed he sys ema ics
p oposed by Wilson e al. (2016, 2017) and Wilson and Mi e meie
(2018), also conside ing he wo k o K yˇ
s u ek e al. (2020) o Cle h-
ionomys and K yˇ
s u ek and Shenb o (2022) o Mic o us (Te icola)
py enaicus. The numbe o specimens was ob ained by coun ing he mos
equen axonomically iden i iable elemen pe species (NISPi, see
Lyman, 1984). The ela i e abundance o each mic omammal species
was based on he minimum numbe o indi iduals (MNI), which was
calcula ed conside ing he la e ali y o he mos equen diagnos ic
elemen .
3.3.2. Mac omammals
The main mac omammal emains om each le el we e eco e ed
di ec ly om he exca a ion su ace and coo dina ed indi idually. In
addi ion, sedimen samples o mechanical sie ing o he smalle bones
and agmen s we e collec ed and labelled acco ding o dep h.
Ana omical and axonomical iden i ica ions ollowed he wo ks o Pales
and Lambe (1971), Schmid (1972), Ba one (1976), and Hillson (2005).
The Os eological Collec ion o he E oAdap a g oup a he Uni e si y o
Can ab ia was also used o compa ison pu poses.
The age a dea h has been es ima ed based on he mola de elopmen
and he epiphyseal usion deg ee o he limb bones. Age es ima ions
ollowed he c i e ia published by Noddle (1974), Sil e (1980), Ma -
iezku ena (1983), Azo i e al. (2002), and Tom´
e and Vigne (2003).
Rela i e abundance is based on he numbe o iden i ied specimens
(NISP) wi h p e ious eassembling o he agmen ed bones.
3.4. P elimina y iden i ica ion o p eda o y al e a ions
To e alua e i he likely cause o he mic omammal accumula ion
was o he han he acciden al alling o he ca e, he enamel o a ico-
line mola s was analysed in sea ch o al e a ions caused by diges ion
(and, consequen ly, p eda ion), ollowing And ews (1990) and
Fe n´
andez-Jal o e al. (2016). The obse ed diges ion signals we e
coun ed and classi ied acco ding o he ollowing ca ego ies: absen ,
ligh , mode a e, hea y, and ex eme (And ews, 1990; Fe n´
andez-Jal o
e al., 2016). Mac omammal bones we e also obse ed using a Leica
s e eomic oscope a 10–80×magni ica ion in sea ch o p eda o y o
na u al al e a ions ollowing Bin o d (1981) and Ma ín-A oyo (2010).
3.5. Paleoen i onmen
The landscape su ounding he PL-III sha was in e ed using he
Habi a Weigh ing Me hod (HWM; E ans e al., 1981; And ews, 2006;
Blain e al., 2008; Cuenca-Besc´
os e al., 2009; L´
opez-Ga cía e al., 2014),
which is based on he habi a - ype p e e ences o each mic omammal
axon. The ob ained habi a ypes a e he ollowing: open d y (OD)
comp ises meadows unde seasonal clima e change, d y g asslands, and
sc ublands; open humid (OH) co esponds o e e g een meadows wi h
dense pas u es and sui able opsoil; open woodland (OW) ep esen s
woodland ma gins and o es pa ches wi h mode a e g ound co e ;
woodland (Wo) indica es ma u e o es ; and wa e (Wa) co esponds o
a eas along eshwa e s eams, lakes, and ponds. The en i onmen al
p e e ences o each species we e ob ained om No es (1989), Wilson
e al. (2016, 2017), and Wilson and Mi e meie (2018). Ha e (Lepus
eu opaeus) emains we e ob ained simul aneously by exca a ion
( h oughou he comple e sequence) and sie ing. This makes hem
o e ep esen ed in he sampling compa ed o he o he mic omammal
species, which we e only sampled in speci ic sec ions, so hey ha e been
excluded om landscape econs uc ions. The house mouse (Mus mus-
culus) is a commensal species o humans ha commonly inhabi s an-
h opic en i onmen s; he e o e, i has no been included in he HWM
calcula ions.
3.6. Paleoclima e
The clima ic condi ions a he ime o each le el's accumula ion we e
es ima ed using he Mu ual Ecogeog aphic Range (MER) me hod, based
on mic o e eb a e associa ions (e.g., Ma ínez-Solano and Sanchiz,
2005; Blain e al., 2009, 2016; Fagoaga e al., 2019). This me hod is
equen ly used o calcula e he mean annual empe a u e (MAT), he
mean empe a u e o he wa mes mon h (MTW), he mean empe a u e
o he coldes mon h (MTC), and he mean annual p ecipi a ion (MAP).
In his wo k, we ha e also used his me hod o ob ain mon hly ain all
and empe a u e alues o obse e seasonal pa e ns. These pa ame e s
we e calcula ed by ge ing he common geog aphical a eas ( ep esen ed
in a ne o 10 ×10 km UTM squa es) o each mic omammal species
associa ion (including ha es) om he PL-III sequence. The clima ic da a
o he sha ed UTM squa es o each assemblage and he cu en clima ic
da a om he PL-III si e we e ob ained using he online applica ion
ag oclimap.aeme .es o he Ibe ian Clima e A las (Cou o e al., 2011).
The cu en dis ibu ion o each species was ob ained om Palomo e al.
(2007).
4. Resul s
4.1. Radioca bon da ing
Ob ained da es anged om 4402 o 836 cal y BP (Table 1). Fou
samples yielded an age acco ding o hei s a ig aphical posi ion. In
con as , sample Be a-541,219, a cha coal collec ed a he bo om o he
exca a ed deposi , was likely an ou lie as i p o ided he younges da e
o he se (1055–836 cal y BP).
4.2. P eda o y ac i i y
Tee h wi h signs o diges ion a e sca ce in he mic omammals'
sample (<1%), bu in he ew cases o diges ed mola s, hey eached
hea y o ex eme co osion. Acco ding o he classi ica ion p oposed by
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And ews (1990), his sugges s ha ca ego y-5 p eda o s, mainly
mammalian ca ni o es, inges ed hose ew indi iduals. Howe e , mos
o he specimens obse ed (99%) do no show p eda ion ea u es.
The mac omammal sample does no show bu che y o bu ning
(cooking) ea u es. The only p eda o signals in he PL-III bone accu-
mula ion a e hose p oduced by ca ni o es (Table 5), a ec ing 6.2% o
he specimens. Ca ni o e oo h ma ks a e mo e common a le els C
(12.3%) and B (9%), being sca ce a le els E (5.9%), F (5.3%), and D
(4.6%).
4.3. Mic omammal eco d
The Mic omammal assemblage o he PL-III sequence yielded a
minimum (MNI) o 1396 indi iduals belonging o 19 axa (Table 2;
Figs. 3–6; Appendix 3). Among oden s, a icolines (Fig. 3; Table S3.1)
a e ep esen ed by gene a A icola (A . sapidus), Mic o us (Mi. (Ag icola)
la e nedii, Mi. (Mic o us) a alis, Mi. (Te icola) sp., Mi. (Te icola) py -
enaicus, and Mi. (Te icola) ex g . lusi anicus–duodecimcos a us), and
Cle h ionomys (Cl. gla eolus); addi ionally, mu ids (Fig. 4a–c) include
h ee species belonging o h ee gene a: Mic omys minu us (Fig. 5A;
Table S3.2), Mus musculus, and Apodemus (Syl aemus) syl a icus (Fig.
S3.1; Table S3.3). In he eulipo yphlans o de , he so icids amily
(Fig. 4d–i’) yielded six axa belonging o gene a C ocidu a (C . ussula
and C . gueldens aed ii: Fig. S3.2; Table S3.4), So ex (Soco ona us and So.
minu us) and Neomys (N. anomalus and N. odiens nie hamme i: Fig. 5B;
Table S3.5); alpids and e inaceids a e ep esen ed by one species each:
Talpa aqui ania (Fig. S3.3; Fig. S3.4; Table S3.6) and E inaceus eu opaeus
espec i ely. Las ly, he mo phology and size o he pos c anial skele on
(Fig. 5C; Fig. S3.5a–d’; Table S3.7) and he lowe hi d p emola s (Fig.
S3.5b) allowed us o assign he lagomo ph emains o he species Lepus
eu opaeus. A icolines yielded he mos signi ican numbe o in-
di iduals om le els F, D, and B (Table 2; Fig. 6), being Mi. (Ag.) la -
e nedii he bes - ep esen ed species a hese le els (35.6–42.9%).
Con e sely, he lowes p opo ion o his a icoline, eached a Le el C
(19.4–7.8%), ma ches a conside able inc ease in he abundance o C .
ussula, a so icid ha is sca cely ep esen ed a he bo om o he
sequence (le els F and D: 0.8% and 0% espec i ely).
We iden i ied wo ypes o mic omammal associa ions in he PL-III
sequence acco ding o hei sha ed dis ibu ion ange (Appendix 4;
Fig. 7). The i s one, coming om Le el F, cohabi s nowadays in a eas a
he no he n slope o he Can ab ian moun ains (Fig. 7A), whe e oceanic
in luence is highe . Con e sely, species associa ions om le els C and B,
a p esen , mainly coexis a he sou he n slope o he Can ab ian
moun ains (Fig. 7B–C) in a eas o a mo e con inen al clima e.
4.4. Mac omammal eco d
The s udied sample comp ises 3220 emains (NSP), o which 1569
(48.7%) could be axonomically classi ied (Table 3). Among domes ic
ungula es, bo ids domina e he assemblage (Fig. 8a– ), mainly sheep
and goa s (O is o ien alis a ies: Fig. 8a–a’; and Cap a aegag us hi cus:
Fig. 8b–d) a he op sec ion o he s a ig aphic sequence (le els C–B:
Fig. 9; Table 3), ollowed by ca le (Bos p imigenius au us; Fig. 8e– ).
Con e sely, equids (Equus sp.) a e ba ely ep esen ed, al hough hese
ha e yielded ema kably well-p ese ed emains (Fig. 8i). All he suid
emains co espond o in an ile (mos ly) o young indi iduals
(Fig. 8j–k): den i ions s ill e ain he deciduous ee h in wea ing (M3/m3
a e no ye e up ed a any o he indi iduals), and limb bones a e no ye
ully g own (epiphyses we e no used a he ime o dea h). Conse-
quen ly, i was no possible o asc ibe hese emains o he domes ic pig
o he wild boa , so hey a e classi ied as Sus sc o a ssp. Wild ungula es
comp ise wo ce id species: he ed dee (Ce us elaphus: Fig. 8g) and
he oe dee (Cap eolus cap eolus: Fig. 8h–h’); bo h, especially he ed
dee , a e well ep esen ed a he lowe sec ion o he sequence (le els
F–D: Fig. 9; Table 3). Finally, mos ca ni o e emains belong o he
Eu opean badge (Meles meles). Howe e , emains o canids ha e also
been ound, which, due o hei small size, a e asc ibed o dogs (Canis
lupus amilia is). The age p o ile o he assemblage (Table 4) is domi-
na ed by in an ile (55.8%) and ju enile (24.4%) indi iduals, while
adul s (19.8%) a e he less abundan g oup. In he case o Equus sp.,
Ce us elaphus and Sus sc o a ssp. samples, hese each 80% o in an ile
indi iduals.
4.5. Paleoen i onmen
The di e en mic omammal assemblages ound a each o he s ud-
ied laye s o he PL-III sequence desc ibe an open landscape in an
e ol ing con ex (Table 6, Fig. 10D). Highes p opo ion o mois u e
indica o s (OH: 47.9%) is a he bo om o he sequence (Le el F), whe e
lowes alues o a idi y we e also ob ained (OD: 0.6%). The ma u e
o es signal is weak in mos o he le els (Wo: <1%), eaching he
highes alues also a Le el F (Wo: 2.4%). Howe e , a idi y indica o s
inc ease owa ds de op o he deposi , especially a Le el C (OD:
42.5–52.9%). Rega ding he mic omammal sample om Le el D, his is
no quan i a i ely enough (MNI: 14) o make eliable in e p e a ions,
hough, excep o he highe p opo ion o aqua ic indica o s (ΔWa:
+20%) and he sligh ly inc easing a idi y in his le el (ΔOD: +4.7%), a
ce ain simila i y among le els D and F samples can be obse ed. The
open landscape in e ed om mic omammal assemblage is consis en
wi h he s a ig aphic s udy o he PL-III si e (Appendix 1). This s udy
sugges s sedimen s esul ing om soil e osion and un-o p ocesses in
he su oundings o he paleon ological si e. Bo h p ocesses a e common
in an open landscape wi h li le ege a ion o p o ec he soil om
e osion and imp o e wa e in il a ion a he han uno p ocesses
(Ga cía-Ruiz, 2010).
4.6. Paleoclima e
Acco ding o MER esul s (Appendix 4; Fig. 11; Fig. 12A), mean
empe a u es du ing he o ma ion o Le el F (MAT: 12.5 ±0.7 ◦C),
which a e sligh ly unde he p esen -day da a (MAT: 13.9 ±0.2 ◦C), a e
dec easing owa ds le els C (MAT: 9.9 ±1.7 ◦C) and B (MAT: 10.3 ±
1.3 ◦C), especially he win e empe a u es (Figs. 13–14a). The mean
ain all o Le el F (MAP: 1359 ±298 mm) is sligh ly o e cu en mean
alues (MAP: 1156 ±17 mm), hough i also dec eases in le els C (MAP:
968 ±319 mm) and B (MAP: 893 ±339 mm).
Mon hly empe a u e and ain all es ima ions (Appendix 4; Fig. 11)
ob ained o Le el F display he ypical A lan ic pa e n o he Can a-
b ian coas . Howe e , he climog aphs o le els C and B show an
inc easing con inen al end, wi h mo e ex eme di e ences be ween
Table 1
Radioca bon da es om Pun a Luce o.
Le el Dep h (cm) Ma e ial Me hod Lab Re
14
C De ia ion cal y BP
a
B 77 Bone AMS Be a-629,438 1980 30 1992–1830
C 242 Bone AMS Be a-541,215 2630 30 2837–2723
D 296 Cha coal AMS Be a-541,218 3070 30 3366–3185
F 430 Bone AMS Be a-541,217 3830 30 4402–4097
F 445 Cha coal AMS Be a-541,219 1040 30 1055–836
a
Calib a ed using he so wa e OxCal .4.4.4 (B onk Ramsey, 2009) agains he In Cal20 cu e (Reime e al., 2020). Fu he da a is p o ided in Appendix 2.
A. ´
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win e and summe empe a u es and a conside able dec ease in ain all
h oughou he yea .
5. Discussion
5.1. O igin o he mammal assemblage
P eda ion signals obse ed in he mic omammal sample only a ec a
ew isola ed mola s (<1%), which each a high–ex eme deg ee o
enamel co osion p oduced by diges ion. These specimens could ge he
deposi in he s omach o he apped ca ni o es (Canis lupus amilia is o
Meles meles) since no o he ea u es exclusi ely p oduced by p eda o s
we e obse ed. The aphonomic signals caused by he in ense ock all o
he exca a ion p ocedu es could be masking hose p oduced by
noc u nal bi ds o p ey o o he p eda o s, which canno de ini i ely be
uled ou o ac ing as accumula ing agen s. In any case, no bias is
obse ed ega ding he p ey ype- o size ange. The e o e, he mos
likely causes o he mic omammal sample accumula ion a e acciden al
alling in he ca e, pelle egu gi a ion by noc u nal bi ds o p ey, o a
combina ion o bo h. We do no exclude ha un-o p ocesses deposi ed
some mic omammal emains a e small anspo (<5–10 m) om he
icini y o he PL-III si e (Appendix 1). O he wise, he low anspo was
insu icien o ound o he skele al/den al emains.
Rega ding he mac omammal sample, he p opo ion o ca ni o e
oo h ma ks (6.2%: Table 5) is signi ican ly lowe han ha obse ed in
badge dens by A illa e al. (2020) and Mallye e al. (2008), whose
samples showed his ype o al e a ion in he 40% and he 20–30% o he
bones, espec i ely. The mo ali y p o ile o he mac omammal sample
(Table 4) also displays clea di e ences wi h he ob ained a badge
bu ows by A illa e al. (2020), whe e he 64% o he indi iduals a e
adul s, 28% a e ju enile, and 8% a e in an ile. Con e sely, his pa e n
in e ses a PL-III (adul 15.9%; ju enile 23.8%; in an ile 60.3%). Na -
u al pi all aps, such as he one s udied by ´
Al a ez-Lao (2014), also
p o ided samples domina ed by young indi iduals since hose a e mo e
likely o ge in ol ed in po en ially a al si ua ions. In his case, ca -
casses o dead animals apped in he ca e would ha e been sca enged
by badge s (Meles meles), whose oo h ma ks a e compa ible wi h hose
obse ed in PL-III. The e o e, badge s, he bes - ep esen ed ca ni o e in
he assemblage, could ha e allen in o he ca e while chasing he smell
o he cada e s.
5.2. Paleoen i onmen and paleoclima e
The mic omammal accumula ion o he PL-III sequence s ands ou by
i s g ea abundance, species ichness, and s a e o p ese a ion. Mos o
he iden i ied species inhabi he egion nowadays; howe e , when
compa ed o a sampling on egu gi a ion pelle s pe o med by Gonz´
alez
O eja e al. (1993), some di e ences can be highligh ed: mode n as-
semblages in he ca e su oundings (Fig. 6) include he b own a
(Ra us no egicus) while lacking o he axa such as he bank ole
(Cle h ionomys gla eolus) and he Güldens ¨
ad 's sh ew (C ocidu a guel-
dens aed ii), which a e displaced o less-an h opized inland a eas (Pal-
omo e al., 2007), o he common ole (Mi. (Mi.) a alis), whose p esence
a no h-wes e n Ibe ia is oday limi ed o he sou he n slope o he
Can ab ian moun ains.
Conce ning he e olu ion o he PL-III mic omammal associa ion
(Fig. 6), some axonomic and biogeog aphical ema ks o paleoen i -
onmen al alue can be add essed. Among oden s, species o he ield
ole g oup (Mic o us (Ag icola) ex g . ag es is) a e some o he mos
equen mic omammals in mode n and ossil assemblages o he egion.
The phylogeny o his g oup has been unde con inuous e isions,
leading o he ecogni ion o h ee di e en species unde simila
mo phological ea u es (Paup´
e io e al., 2012; Wilson e al., 2017). A
he ime o he o ma ion o his paleon ological si e, he no he n and
sou he n lineages o he Mi. (Ag.) ag es is g oup (Mi. (Ag.) la e nedii, Mi.
(Ag.) ozianus and Mi. (Ag.) ag es is) had al eady spli since, acco ding o
Table 2
Mic omammal species om Pun a Luce o and hei weigh ed habi a -p e e ences.
Le el F Le el D Le el C-bo om Le el C-middle Le el C-uppe Le el B Habi a ype
(Sample z: 445–420) (Sample z: 296–288) (Sample z: 280–269) (Sample z: 222–208) (Sample z: 208–177) (Sample z: 0–93)
NISP
i
MNI %MNI NISP
i
MNI %MNI NISP
i
MNI %MNI NISP
i
MNI %MNI NISP
i
MNI %MNI NISP
i
MNI %MNI Wa OD OH OW Wo
A icola sapidus 6 4 0.4 4 3 21.4 2 1 1.7 1
Mic o us (Ag icola) la e nedii 746 389 35.6 8 6 42.9 41 21 18.6 6 4 7.8 11 7 19.4 47 24 40.0 0.5 0.5
Mic o us (Mic o us) a alis 1 1 7.1 11 6 5.3 2 1 2.0 1 1 2.8 13 7 11.7 0.75 0.25
Mic o us (Te icola) sp. 266 21 1.9 1 1 7.1 4 2 1.8 1 1 2.0 0.5 0.5
Mi. (Te.) ex g . lusi anicus-duodecimcos a us 176 88 8.1 1 1 1.7 0.5 0.5
Mi. (Te.) py enaicus 57 32 2.9 0.5 0.5
Cle h ionomys gla eolus 16 11 1.0 2 1 0.9 1
Mus musculus 2 1 – 2 1 – – – – – –
Apodemus (Syl aemus) syl a icus 315 161 14.7 2 2 14.3 17 10 8.8 7 4 7.8 1 1 2.8 5 4 6.7 1
Mic omys minu us 10 7 0.6 1
So ex minu us 35 20 1.8 0.25 0.75
So ex co ona us 532 274 25.1 7 5 4.4 7 4 7.8 1 1 2.8 11 5 8.3 0.75 0.25
Neomys odiens nie hamme i 32 16 1.5 0.75 0.25
Neomys anomalus 2 2 1.8 1 1 2.8 0.25 0.75
C ocidu a ussula 15 9 0.8 112 58 51.3 62 35 68.6 45 24 66.7 24 13 21.7 0.75 0.25
C ocidu a gueldens aed ii 96 49 4.5 0.5 0.5
Talpa aqui ania 20 10 0.9 1 1 7.1 10 8 7.1 3 2 3.9 1 1 2.8 5 4 6.7 0.75 0.25
E inaceus eu opaeus 1 1 0.1 1 1 1.7 0.25 0.25 0.5
To al 2325 1093 100 17 14 206 113 90 52 61 36 103 60
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Paup´
e io e al. (2012), hey did a a ound he Las Glacial Maximum
(LGM). The equency o M1 wi h a pos e o-lingual iangle, like he one
obse ed in he M2, is a disc iminan ea u e among Mi. (Ag.) ex g .
ag es is species (K app and Nie hamme , 1982; Wilson e al., 2017). The
low p opo ion o M1 p esen ing his iangle (1.5%: Table S3.1) has
allowed us o asc ibe he ield- ole specimens om PL-III o he Medi-
e anean ield ole (Mi. (Ag.) la e nedii), which is he cu en ep e-
sen a i e o his g oup in he egion. Nowadays, Mi. (Ag.) la e nedii
sp eads h oughou he Can ab ian Region (Palomo e al., 2007).
Con e sely, Mic o us (Mic o us) a alis (common ole), a species o mo e
Fig. 3. Selec ed a icoline specimens om Pun a Luce o III. A icola sapidus: le m1–m3 om Le el F in occlusal (a) iew. Mic o us (Ag icola) la e nedii: le m1–m3
om Le el F in occlusal (b) iew and le M1–M3 om Le el C in occlusal (c) iew. Mic o us (Mic o us) a alis: le m1–m3 om Le el C in occlusal (d) iew, and le
M1–M3 om Le el C in occlusal (e) iew. Mic o us (Te icola) sp.: le m1 om Le el F in occlusal ( ) iew, and le m1–m3 om Le el F in occlusal (g) iew. Mic o us
(Te icola) ex g . lusi anicus–duodecimcos a us: le M3 om Le el F in occlusal (h) iew and le M3 om Le el F in occlusal (i) iew. Mic o us (Te icola) py enaicus:
le M3 om Le el F in occlusal (j) iew and le M3 om Le el F in occlusal (k) iew. Cle h ionomys gla eolus: le m1–m3 om Le el F in occlusal (l) iew. AL:
an e io loop. T: iangle. Scale ba 1 mm.
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con inen al equi emen s (No es, 1989), li es sou h o he wa e shed, in
he no he n hal o he Ibe ian Pla eau (No es, 1989; Palomo e al.,
2007). Conce ning p opo ion among hese species, in Le el C, a
dec ease in he abundance o Mi. (Ag.) la e nedii is obse ed along wi h
he appea ance o Mi. (Mi.) a alis. Howe e , Mi. (Ag.) la e nedii p e ails
a Le el B, coinciding wi h he highes abundance a io o Mi. (Mi.)
a alis in he sequence. I can also be seen ha when Mi. (Ag.) la e nedii
inc eases, also does A icola sapidus, a empe a e-a ini y wa e ole
om sou hwes e n Eu ope. This pa e n, in which he ini ial absence o
Mi. (Mi.) a alis is ollowed by i s inc easing abundance, is also obse ed
a La Güelga (As u ias, ~48–37 ka, ´
Al a ez-Vena e al., 2021), an MIS 3
si e in which an al e na ion o clima ic changes om empe a e o cold
s adials was epo ed in a scena io o inc easing a idi y.
Rega ding he emnan ole axa om PL-III is also ema kable he
occu ence o a leas wo species o he Te icola subgenus: Mi. (Te.)
py enaicus ( he Py enean pine ole) and Mi. (Te.) ex g . lusi ani-
cus–duodecimcos a us. Obse ed mo pho ypes o he M1 poin o he
p esence o bo h Medi e anean (Mi. (Te.) duodecimcos a us) and Lusi-
anian (Mi. (Te.) lusi anicus) pine oles bu o e lapping mo phological
a iabili y did no make us possible o disc imina e be ween hese spe-
cies, which was e en mo e di icul due o he co-occu ence o he
Py enean pine ole. Fo his eason, he o al MNI o he Te icola sub-
genus has been calcula ed based on he m1. Howe e , he p opo ion o
indi iduals o each species belonging o his subgenus has been
Fig. 4. Selec ed mu id, so icid, and alpid specimens om Pun a Luce o III. Mic omys minu us: le mandible om Le el F wi h m1 in lingual (a) and occlusal (a’)
iews. Mus musculus: Le maxilla agmen om Le el F wi h M1–M2 in occlusal (b) iew. Apodemus (Syl aemus) syl a icus: le M1–M3 se ies om Le el F in occlusal
(c) iew. Neomys odiens nie hamme i: igh mandible om Le el F in pos e io (d) and labial (d’) iews. Neomys anomalus: igh mandible om Le el C in pos e io (e)
and labial (e’) iews. So ex co ona us: igh mandible om Le el F in pos e io ( ) and labial ( ’) iews. So ex minu us: igh mandible om Le el F in pos e io (g) and
labial (g’) iews. C ocidu a gueldens aed ii: igh mandible om Le el F in pos e io (h) and labial (h’) iews. C ocidu a ussula: igh mandible om Le el C in
pos e io (i) and labial (i’) iews. Talpa aqui ania: le hume us om Le el C in pos e io (j) iew. Analogous ana omical elemen s ha e been lipped ho izon ally o
show hem om he same side o compa ison pu poses. Scale ba s 1 mm.
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F om ~5 ka, s eppe pas o alis s o he Yamnaya cul u e expanded
eas - and wes wa d, linking Asia and Eu ope (Allen o e al., 2015;
Laza idis e al., 2022). This c ea ed a geog aphic co ido which, aided
by he de elopmen o ho se- iding and cha io y (An hony, 2007; An-
hony and B own, 2011; Wilkin e al., 2021), led o he dispe sal o
c ops, he ds, and commensal species om one con inen o ano he .
Rega ding he Ibe ian Peninsula, an inc easing gene in low om
No he n and Cen al Eu opean human popula ions wi h s eppe ances y
has been de ec ed (Olalde e al., 2019; Villalba-Mouco e al., 2021;
Pa e son e al., 2022), da ing he ea lies e idence om ~4.5–4 ka,
om indi iduals who coexis ed wi h locals wi hou s eppe ances y
(Olalde e al., 2019). This incoming pas o alis peopling o he egion
ag ees wi h he inc easing an h opic p essu e linked o animal hus-
band y obse ed a PL-III and he coe al eco ds men ioned. Howe e ,
he inc ease in domes ic he ds and de o es a ion is no he only way
hose mig a ions a e pe cei ed in he PL-III deposi . The ha es mouse
(Mic omys minu us), iden i ied a Le el F, and he house mouse (Mus
musculus), ound a le els F and C, a e epo ed om PL-III a a signi i-
can ly ea ly ch onology. Acco ding o Ho ´
aˇ
cek e al. (2013), Pliocene
and Pleis ocene Eu opean eco ds p e iously asc ibed o he Mic omys
genus belong o Pa apodemus co onensis Schaub, 1930, which did no
ha e con inui y in he La e Pleis ocene. This ag ees wi h he wo k o
Fig. 11. Tempe a u e and ain all es ima ions based on he Mu ual Ecogeog aphic Range me hod. (A) Box and whiske plo o he mon hly alues ob ained h ough
he Mu ual Ecogeog aphic Range me hod (Appendix 4) o each mic omammal associa ion o he Pun a Luce o III sequence (le els F, C, and B). (B) Mon hly mean
alues o es ima ed empe a u e and ain all om le els F, C, and B compa ed wi h p esen -day da a om he Pun a Luce o III loca ion. (C) Mu ual Ecogeog aphic
Range me hod es ima ions: mean annual empe a u e (MAT), mean empe a u e o he wa mes mon h (MTW), mean empe a u e o he coldes mon h (MTC), mean
annual p ecipi a ion (MAP).
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Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
17
Yasuda e al. (2005), who, based on he phylogeog aphic pa e n o
m DNA, p opose ha he ex an species o igina ed in Eas Asia. The
ossil eco d sugges s ha Mic omys minu us eached Eu ope du ing he
Holocene (e.g., Mis o , 2011, 2013a, 2013b; Se jean son, 2011;
K ajca z e al., 2020; Roye e al., 2021), bu he sca ci y o eliable
eco ds o da e does no allow us o econs uc a p ecise iming. Con-
ce ning he Ibe ian Peninsula, he species has been iden i ied a wo
di e en si es: Le el 2 o Valda a a-1 (Lugo, Galicia; 5305–4978 cal y
BP; L´
opez-Ga cía e al., 2011) and Le el 1 o Amalda (Gipuzkoa, Basque
Coun y; Pem´
an, 1990), which yielded h ee da es anging om 3875 o
1179 cal ky BP ( he oldes conside ed abe an by Al una, 1990).
Conside ing he measu emen s published in L´
opez-Ga cía e al. (2011),
he ma e ial asc ibed o Mic omys minu us in Valda a a-1 alls in size
wi h small specimens o Apodemus syl a icus (Fig. 5A), which could
indica e ha i is his species and no he p e ious one, in he absence o
e iewing he iden i ied ma e ial. This means ha he epo ed in his
wo k could be he oldes eco d o his species in he Ibe ian Peninsula.
The sp ead o he ha es mouse ac oss Eu asia was p obably boos ed
by he de elopmen o ag icul u e, especially by mille c ops, which,
con e sely o o he ce eals, do no belong o he i s se o Neoli hic
c ops om he Middle Eas . Ins ead, his ce eal was ini ially domes i-
ca ed in no h-eas e n China a ~8 ka (Liu e al., 2004; C aw o d e al.,
2013) and la e sp ead o he es o Cen al Eu asia and Eas e n Eu ope
(Mille e al., 2016; Filipo i´
c e al., 2020). Fox ail mille (Se a ia i alica),
among o he emains o ce eals, has been ound a he nea by si es o
Kobaede a (Le el 1) and A enaza (Le el 9) a 5280–4855 cal y BP and
4085–3693 cal y BP espec i ely (Zapa a Pe˜
na, 1999, 2002). The e o e,
he ha es mouse occu ence a PL-III (Le el F: 4402–4097 cal y BP)
and he a i al o mille c ops o he egion sha e simila ch onological
anges. The ea lies e idence o mille consump ion by humans in he
Can ab ian Region (3163–3051 cal y BP) has been documen ed a El
Espinoso (As u ias; Gonz´
alez-Rabanal e al., 2022) a ~100 km wes o
PL-III by indi iduals wi h a signi ican p opo ion o s eppe ances y
(30%, Pa e son e al., 2022). The e o e, s eppe mig a ions could be
ela ed o ha es mouse a i al in he Can ab ian Region.
Abou he house mouse (Mus musculus), i s eco d in sou hwes e n
Eu ope has been e iewed by Cucchi e al. (2005) and Domínguez Ga cía
e al. (2019), who ound ha mos o he epo s o he species lack
published eliable iden i ica ions o come om unclea con ex s. E en
new da a om he Can ab ian Region published by Ma ínez-Villa e al.
(2022) and Ce nadas-Ga ido e al. (2023) may co espond o in usi e
specimens and ma e ials o mixed–ch onology, espec i ely. The ea lies
e idence o Mus musculus domes icus in Ibe ia comes om I on Age a he
Alo da Pa c si e (Valenzuela-Lamas e al., 2011). Domínguez Ga cía
e al. (2019) also place he ea lies eliable p esence o he house mouse
a he Ibe ian Peninsula wi hin he I on Age, a ~3 ka, suppo ing he
hypo hesis o a dispe sal linked o he inc ease in he sea ade be ween
eas e n and wes e n Medi e anean (Cucchi e al., 2005), d i en by he
comme cial expansion o he Phoenicians and G eeks (Cucchi e al.,
2012). The oldes eco ds o he house mouse (Mus musculus musculus)
om inland Eu ope come om Eas e n Eu ope and da e om he end o
he Neoli hic (~6.5 ka; Cucchi e al., 2011, 2020). Based on his unex-
pec ed inding, Cucchi e al. (2012) p oposed a cen e o synan h op-
iza ion o his subspecies in eas e n Eu ope no h o he Black Sea.
The e o e, a ou e om he s eppes h ough inland Eu ope could be an
al e na i e pa hway o he a i al o Mus musculus o he Can ab ian
Region. This is consis en wi h he eliable inding o Mus musculus ssp.
in no he n I aly (Tosina, Milan) da ed a ~6 ka (Bona, 2020). Howe e ,
u he esea ch and di ec adioca bon da ing o he PL-III specimens
need o be done o a oid misin e p e a ion o his eco d.
Fig. 12. Compa ed paleoclima ic and paleoen i onmen al econs uc ions o
he Pun a Luce o III sequence. (A) Mu ual Ecogeog aphic Range es ima ions o
le els F, C, and B: mean annual p ecipi a ion (MAP), mean empe a u e o he
wa mes mon h (MTW), mean annual empe a u e (MAT), mean empe a u e o
he coldes mon h (MTC). Dash lines and as e isk columns indica e he p esen -
day alues ob ained om he Ibe ian Clima e A las (Cou o e al., 2011). (B)
Type o landscape e olu ion a he su oundings o he Pun a Luce o III sha
in e ed h ough he Habi a Weigh ing me hod. Le els and samples a e plo ed
acco ding o dep h. The column wi h an as e isk co esponds o a mode n
mic omammal sample collec ed in he Pun a Luce o a ea om ba n owl
egu gi a ion pelle s by Gonz´
alez O eja e al. (1993).
Table 5
P opo ion (%) o skele al emains showing ca ni o e oo h ma ks.
Le el A Le el B Le el C Le el D Le el E Le el F To al
NSP* 46 188 414 989 707 876 3220
Too h-ma ked specimens 0 17 51 45 42 46 201
F equency (%) 0 9 12.3 4.6 5.9 5.3 6.2
*
Numbe o specimens (includes specimens ha canno be iden i ied o axon).
A. ´
Al a ez-Vena e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
18
6. Conclusions
The mammal assemblage s udied in his wo k was mainly p oduced
by acciden al alling in he PL-III ca e, which ac ed as a na u al ap
h ough ime. Noc u nal bi ds o p ey could also be implied in he
accumula ion o he mic omammal assemblage since ou analyses did
no allow us o ule ou hese p eda o s. The s udied samples p o ided a
la ge numbe o emains co esponding o 18 mic omammal and nine
mac omammal axa. Mic omammal communi y unde wen a quick
u no e which e lec s a shi o he en i onmen al condi ions ha ook
place du ing he o ma ion o he deposi : he mois mosaic o o es and
meadows obse ed in Le el F (4,402–4097 cal y BP) u ned in o an
open and sh ubbie g assland du ing he accumula ion o le els C
(2,837–2723 cal y BP) and B (1,992–1830 cal y BP). The ea lies
Ibe ian eco ds o Mus musculus and, mos likely, o Mic omys minu us
he e epo ed could be ela ed o he dispe sal o he s eppe pas o alis .
S ill, di ec adioca bon da ing needs o be done o ob ain a mo e p ecise
da e o hese emains. Simul aneously, mac omammals expe ienced a
shi om wild popula ions o domes ic he ds, which, along wi h he
abundan cha coal agmen s eco e ed in he sedimen , sugges ha
an h opogenic i es o he de elopmen o li es ock pas u es caused
o es disappea ance and did no allow hem o eco e . Tempe a u e
and ain all es ima ions e eal ha he humid and mild clima ic con-
di ions obse ed a he bo om o he sequence, which occu ed du ing
he La e Holocene The mal Maximum, wen h ough a cooling and a i-
di ica ion phase, coe al wi h he I on Age Cold Epoch, and concluded in
a sligh wa ming, coe al wi h he Roman Wa m Pe iod.
Decla a ion o Compe ing In e es
The au ho s decla e ha hey ha e no known compe ing inancial
in e es s o pe sonal ela ionships ha could ha e appea ed o in luence
he wo k epo ed in his pape .
Da a a ailabili y
No da a was used o he esea ch desc ibed in he a icle.
Acknowledgemen s
AB.M-A. de eloped pa o his esea ch as pa o he ERC Consoli-
da o G an (SUBSILIENCE e . 818299). We hank J.A. Delgado o his
echnical wo k on s udying mac omammal assemblage. Financial sup-
po was p o ided by he Bilbao Po Au ho i y (Au o idad Po ua ia de
Bilbao) wi hin he p ojec “Es abilizaci´
on del sec o occiden al de la
Can e a de Pun a Luce o en el Pue o de Bilbao”. We a e also g a e ul o
Juan Manuel L´
opez-Ga cía and he anonymous e iewe o hei sug-
ges ions and commen s ha s ongly imp o ed he manusc ip .
Appendix A. Supplemen a y da a
Supplemen a y da a o his a icle can be ound online a h ps://doi.
o g/10.1016/j.palaeo.2023.111476.
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Weigh ed (%) habi a -p e e ences
a
wi h 95% con idence in e als
b
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MNI Wa OD OH OW Wo
Zie bena mode n
(Gonz´
alez-O eja e al., 1993) 773 0.00 36.58 13.71 49.61 0.10
95% CI 0.00–0.48 33.21–40.12 11.37–16.34 45.97–53.13 0.00–0.72
Le el B
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95% CI 0.04–8.94 14.72–37.86 21.69–46.69 27.56–53.46 0.00–5.96
Le el C-uppe
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95% CI 0.00–9.74 35.49–69.59 6.37–32.81 16.35–48.11 0.00–9.74
Le el C-middle
(Sample z: 222–208) 51 0.00 52.94 13.73 33.33 0.00
95% CI 0.00–6.98 38.46–67.07 5.70–26.26 20.76–47.92 0.00–6.98
Le el C-bo om
(Sample z: 280–269) 113 0.44 42.48 20.13 36.06 0.88
95% CI 0.00–3.21 33.23–52.13 13.36–28.96 27.45–45.86 0.02–4.83
Le el D
(Sample z: 296–288) 14 21.43 5.36 30.36 42.86 0.00
95% CI 4.66–50.80 0.18–33.87 8.39–58.10 17.66–71.14 0.00–23.16
Le el F
(Sample z: 445–420) 1092 1.47 0.62 47.50 47.99 2.43
95% CI 0.84–2.37 0.26–1.32 44.53–50.54 44.99–51.00 1.64–3.58
a
Wa, wa e ; OH, open humid; OD, open d y; OW, open woodland; Wo, woodland.
b
95% CI: mul iple p opo ions 95% con idence in e als calcula ed using he Cloppe -Pea son me hod (Cloppe and Pea son, 1934) using he PAST 4.0 so wa e
(Hamme e al., 2001).
A. ´
Al a ez-Vena e al.
Palaeogeog aphy, Palaeoclima ology, Palaeoecology 616 (2023) 111476
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