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\title{Testing the Dry Refuge Model: paleoecological insights from Late
Pleistocene gomphotheres in Ecuador}



\author[1]{erwin gonzalez}%
\author[2]{Iván Ramírez Pedraza}%
\author[3]{Lidiane Asevedo}%
\author[4]{Natalia Villavicencio}%
\author[5]{Melissa Hidalgo}%
\author[6]{Carlos Tornero}%
\author[7]{Sebastián Escobar}%
\author[8]{Jose Luis Roman}%
\author[8]{Esteban Benalcazar}%
\author[9]{Florent Rivals}%
\affil[1]{Universidad de O'Higgins}%
\affil[2]{Universidad de Barcelona Facultad de Biología}%
\affil[3]{Universidade do Estado do Rio de Janeiro}%
\affil[4]{Pontificia Universidad Catolica de Chile}%
\affil[5]{Terra Ignota}%
\affil[6]{Universidad Autonoma de Barcelona}%
\affil[7]{Pontificia Universidad Catolica del Ecuador}%
\affil[8]{Escuela Politécnica Nacional}%
\affil[9]{Institut Català de Paleoecologia Humana i Evolució Social }%


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  \date{April 25, 2026}


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\begin{abstract}
Aim: Paleoenvironmental records have indicated that neotropical areas
near the equatorial area experienced a decrease in rainfall during the
Last Glacial Maximum. However, some records show contradictory
paleoenvironmental signs. To resolve these differences, previous studies
have proposed two demographic/genetic scenarios for cloud forests based
on opposing precipitation regimes. We use a multiproxy approach on
extinct proboscideans (so-called gomphotheres) from Ecuador, to
determine the magnitude of the influence of the wet and dry forest model
on the coast and in the Inter-Andean Valley of Ecuador. Location:
Ecuador. Taxon: Notiomastodon platensis. Methods: Fossil molars were
analyzed using stable isotope analysis, dental microwear analysis, and
radiocarbon dating. Results: The gomphotheres that were analyzed by
radiocarbon dating indicated a range between 23,830 and 31,820 cal yr
BP. The \selectlanguage{greek}δ\selectlanguage{english}13Cbioapatite results (--14.82 to --0.78 %0) indicate that
proboscideans lived in a wide variety of environments; however, the
values indicate a tendency towards more open and relatively dry
environments. We also used dental microwear analysis, which points in
the same direction: mostly grass consumption. The \selectlanguage{greek}δ\selectlanguage{english}18O meteoric water
values calculated from tooth enamel indicate a clear altitudinal effect
on the distribution of proboscideans. Furthermore, the \selectlanguage{greek}δ\selectlanguage{english}18O meteoric
water values are similar to those observed at present-day meteorological
stations in Ecuador. Main conclusions: At least during the lifetime of
proboscideans, we interpret that there were no significant differences
in the \selectlanguage{greek}δ\selectlanguage{english}18O values of atmospheric circulation between the Late
Pleistocene and the present in the Ecuador area. Although there are
still large chronological gaps in paleoenvironmental reconstruction, the
effects of the ''dry refuge model'' probably had the greatest impact on
the landscape of Ecuador during the Late Pleistocene; namely, cloud
forests underwent a process of displacement and compression into refuges
due to the opposing effects of aridity and cooling.%
\end{abstract}%



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\verb`Table S1.docx`  available at \url{https://authorea.com/users/1014210/articles/1382721-testing-the-dry-refuge-model-paleoecological-insights-from-late-pleistocene-gomphotheres-in-ecuador}

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\verb`Table S3.docx`  available at \url{https://authorea.com/users/1014210/articles/1382721-testing-the-dry-refuge-model-paleoecological-insights-from-late-pleistocene-gomphotheres-in-ecuador}

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\verb`Table S4docx.docx`  available at \url{https://authorea.com/users/1014210/articles/1382721-testing-the-dry-refuge-model-paleoecological-insights-from-late-pleistocene-gomphotheres-in-ecuador}

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\verb`Table S5.docx`  available at \url{https://authorea.com/users/1014210/articles/1382721-testing-the-dry-refuge-model-paleoecological-insights-from-late-pleistocene-gomphotheres-in-ecuador}

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