Recuerdo de Ramón Margalef - Encuentros en la Biología

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Primavera 2014
!!
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F. Xavier Niell
Catedrático de Ecología de la Universidad de Málaga
[email protected] !
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Vol.7 ¦ Nº 147
Primavera 2014
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!
Referencias:
! Margalef, R. 1997. Our Biosphere. Excellence in Ecology 10. Ecology Institute. Oldendorf/Luhe.
! Peters, F. 2010. Ramón Margalef, the curiosity driven life of a self-taught Naturalist. Limnologia and Oceanography Bulletin
19 (1): 2-15.
Vol.7 ¦ Nº 147
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