Carla Sousa Santos

Carla Sousa Santos

Post-doctoral fellow

Membro da Comissão Executiva do Centro de Biociências



Phone: + 351 21 8811700

Fax: + 351 21 8860954

Personal Website

Curriculum vitae

Interview on CVTV

LinkedIn: carla-sousa-santos/8/421/155

ResearcherID: B-5385-2012

ORCID: 0000-0002-4513-5553 


:: Research interests


Conservation Genetics
Phylogeography and Phylogeny
Ex-situ Conservation
River Restoration
Fish Behaviour


:: Current research projects


My postdoctoral research has been dedicated to the conservation of native freshwater fishes. I was responsible for a project aiming the characterization of the genetic diversity of all the native Portuguese cyprinid species, which resulted in the publication of the first National Genetic Atlas of this fish family. I'm also involved in an ex-situ reproduction program to restock natural populations of endangered cyprinid species in eminent risk of extinction. In 2011 and 2013, more than 6000 captive bred fish were already released in the rivers of origin of their parents, contributing to the reinforcement of wild populations. Currently I am interested in the comparative phylogeography of native freshwater fish, in reconstructing the colonization routes followed by ancient lineages during the colonization of Iberia, and in providing valuable data to establish conservation priorities, especially for endangered fish Mediterranean-type streams subjected to intense droughts.


FISHATLAS  Project - Portuguese Atlantic- and Mediterranean-type Rivers under the effects of climate changes: current and historical demography and comparative phylogeography of fishes as a tool for the conservation of critically endangered species (Project funded by: FCT - PTDC/AAC-CLI/103110/2008)


The Mediterranean region is known to be a hotspot for diversity and endemism, and one of the regions that apparently will be more strongly affected by global climate change. The risk of extinction is particularly high for taxa that are unable to migrate to more favorable regions like primary freshwater fish, which are confined to the rivers where they occur. As our country benefits from a geographic location that is at the transition from an Atlantic climate to a Mediterranean one, the main goal of this project is to exhaustively characterize the genetic structure of populations from Atlantic- and Mediterranean-type rivers with contrasting hydrology and climate (past and present) and to assess the possible effects of future climatic changes. For more information on this project see:


Study of the genetic diversity of the critically endangered endemic freshwater fish species and implementation of ex-situ reproduction programmes (Post-Doctoral Research Grant funded by: FCT - SFRH/BPD/29774/2006)


From the 26 native cyprinid species of the Portuguese freshwater fish fauna, seven are considered to be critically endangered. Most of these species are found in small southern rivers that are subjected to extreme summer droughts and that generally present severe pollution levels and degraded habitats. The persistence of fish populations is thus extremely unstable. Our aim is to conduct a survey of the genetic diversity of the most endangered populations in order to delineate a conservation plan of their valuable biological patrimony. Additionally, major efforts will be conducted to breed the endangered species ex-situ so that, after their rivers of origin have been restored, re-introductions may be possible. Ex-situ reproduction has been recognized by several authors as a necessary measure whenever the habitats of origin of the species are so degraded that the viability of the species is very low or when the numbersof individuals in the populations are so reduced that the risk of extinction is eminent. Ex-situ reproduction programs do not solve the conservation problems menacing the species but represent alternatively a measure to avoid extinction, until habitats are restored.


The ex-situ reproduction program has already been launched for five endangered freshwater fish species - for more information on this project see:


:: Presentations


• Cada rio é uma ilha (pps)

• Conservação ex situ de peixes de água doce criticamente ameaçados 



:: Publications

Sousa-Santos C, Robalo J, Pereira A, I Doadrio. in press. Threatened fishes of the world: Cobitis vettonica. Croatian Journal of Fisheries


Sousa-Santos C, Robalo J, Pereira A, I Doadrio. in press. Threatened fishes of the world: Achondrostoma salmantinum. Croatian Journal of Fisheries


Sousa-Santos, C; Robalo, JI; Francisco, SM; Carrapato, C; Cardoso, AC; Doadrio, I (2014) Metapopulations in temporary streams - the role of drought-flood cycles in promoting high genetic diversity in a critically endangered freshwater fish and its consequences for the future. Molecular Phylogenetics and Evolution doi: 10.1016/j.ympev.2014.08.007 pdf


Sousa-Santos C, Robalo J, Pereira A, I Doadrio. 2014. Threatened fishes of the world: Squalius malacitanus. Croatian Journal of Fisheries 72:209-212.


Sousa-Santos C, Robalo J, Pereira A, I Doadrio. 2014. Threatened fishes of the world: Squalius castellanus. Croatian Journal of Fisheries 72:206-208.


Sousa-Santos C, Robalo J, Pereira A, I Doadrio. 2014. Threatened fishes of the world: Cottus hispaniolensis. Croatian Journal of Fisheries 72:202-205.


Sousa-Santos C, Robalo J, Pereira A, I Doadrio. 2014. Threatened fishes of the world: Cottus aturi. Croatian Journal of Fisheries 72:198-201.


Sousa-Santos C, Robalo J, Almada V  2014. Spawning behavior of a threatened Iberian cyprinid and its implications for conservation. Acta Ethologica DOI 10.1007/s10211-014-0185-5 pdf


Sousa-Santos, C.; Gante, H; Robalo, J; Cunha, PP; Martins, A; Arruda, M; Alves, MJ & Almada, VC. 2014. Evolutionary history and population genetics of a cyprinid fish (Iberochondrostoma olisiponensis) endangered by introgression from a more abundant relative. Conservation Genetics, 15:665–677. DOI 10.1007/s10592-014-0568-1.  pdf


Sousa-Santos, C.; Gil, F.; Almada, V. 2014. Ex-situ reproduction of Portuguese endangered cyprinids in the context of their conservation. Ichthyological Research. 61 (2) 193–198 DOI 10.1007/s10228-013-0383-6 pdf


Robalo JI, Sousa-Santos C, Coelho MM & Almada VC. 2013. Threatened fishes of the world: Iberochondrostoma almacai Coelho, Mesquita e Collares-Pereira, 2005 (Cyprinidae). Croatian Journal of Fisheries, 71, 147-148.


Almada V. & Sousa-Santos C. 2010. Comparisons of the genetic structure of Squalius populations (Pisces, Cyprinidae) from rivers with contrasting histories, drainage areas and climatic conditions. Molecular Phylogenetics and Evolution 57(2): 924-931.


Gil F, Sousa-Santos C & Almada V 2010 A simple and inexpensive technique for the ex-situ reproduction of critically endangered Cyprinids – Achondrostoma occidentale as a case study. Journal of the World Aquaculture Society 41(4):661-664. DOI: 10.1111/j.1749-7345.2010.00408.x


Sousa-Santos C, Robalo JI & Almada V. 2010. Threatened fishes of the world: Squalius torgalensis Coelho, Bogutskaya, Rodrigues & Collares-Pereira, 1998 (Cyprinidae). Environmental Biology of Fishes 87:123-124. DOI 10.1007/s10641-009-9567-3 


Robalo JI, Sousa-Santos C, Almada VC 2009. Threatened fishes of the world: Iberochondrostoma lusitanicum Collares-Pereira, 1980 (Cyprinidae). Environmental Biology of Fishes 86:295-296. 


Robalo JI, Sousa-Santos C, Cabral H, Castilho R & Almada VC. 2009. Genetic evidence fails to discriminate between Macroramphosus gracilis Lowe 1839 and M. scolopax Linnaeus 1758 in Portuguese waters. Marine Biology 156(8):1733-1737. 


Sousa-Santos C, Robalo JI & Almada V (2009) Threatened fishes of the world: Squalius aradensis (Coelho, Bogutskaya, Rodrigues & Collares-Pereira, 1998) (Cyprinidae). Environmental Biology of Fishes. 86(4):457-458. DOI 10.1007/s10641-009-9550-z 


Robalo, J. I., Sousa-Santos, C., Doadrio, I. & Almada, V. C. 2008. Threatened fishes of the world: Achondrostoma occidentale Robalo, Almada, Sousa-Santos, Moreira and Doadrio, 2005. Environmental Biology of Fishes. DOI 10.1007/s10641-008-9348-4


Sousa-Santos, C., Collares-Pereira, M.J. & Almada, V.C. 2007. Fertile triploid males – an uncommon case among hybrid vertebrates. Journal of Experimental Zoology Part A: Physiology and Evolutionary Genetics, 307(4): 220-225.


Vinyoles, D., Robalo, J. I., De Sostoa, A., Almodóvar, A., Elvira, B., Nicola, G. G., Fernández-Delgado, C., Santos, C. S., Doadrio, I., Sardá-Palomera, F. & Almada, V. C. 2007. Spread of the alien bleak Alburnus alburnus (L., 1758) in the Iberian Peninsula: the role of reservoirs. Graellsia, 63(1): 101-110. 


Sousa-Santos, C., Collares-Pereira, M. J. & Almada, V. C. 2007. Reading the history of a hybrid fish complex from its molecular record. Molecular Phylogenetics and Evolution, 45: 981-996. 


Sousa-Santos, C. 2007. Reproductive behaviour and the evolutionary history of the hybridogenetic complex Squalius alburnoides (Pisces, Cyprinidae). PhD thesis. Universidade de Lisboa.


Sousa-Santos C, MJ Collares-Pereira & VC Almada (2006) May a hybridogenetic complex regenerate the nuclear genome of both sexes of a missing ancestor? - First evidence on the occurrence of a nuclear nonhybrid Squalius alburnoides (Cyprinidae) female based on DNA sequencing. Journal of Natural History 40(23-24): 1443-1448


Sousa-Santos C, MJ Collares-Pereira & VC Almada. 2006. Reproductive success of nuclear nonhybrid males of Squalius alburnoides hybridogenetic complex (Teleostei, Cyprinidae): an example of interplay between female choice and ecological pressures? Acta Ethologica 9(1): 31-36


Sousa-Santos C, MJ Collares-Pereira & VC Almada (2006) Evidence of extensive mitochondrial introgression with nearly complete substitution of the typical Squalius pyrenaicus-like mtDNA of the Squalius alburnoides complex (Cyprinidae) in an independent Iberian drainage. Journal of Fish Biology 68 (Supplement B), 292–301


Robalo JI, C Sousa-Santos, VC Almada & I Doadrio (2006) Paleobiogeography of two Iberian endemic cyprinid fishes (Chondrostoma arcasii-Chondrostoma macrolepidotus) inferred from mitochondrial sequence data. Journal of Heredity 97(2): 143-149 


Robalo JI, C Sousa-Santos, A Levy & VC Almada (2006) Molecular insights on the taxonomic position of the paternal ancestor of the Squalius alburnoides hybridogenetic complex. Molecular Phylogenetics and Evolution 39(1): 276-281 


Sousa-Santos C, JI Robalo, MJ Collares-Pereira & VC Almada (2005) Heterozygous indels as useful tools in haplotype reconstruction of DNA nuclear genes of hybrid organisms – an example using cyprinid fishes. DNA Sequence 16(6): 462-467 


Robalo JI, VC Almada, C Sousa-Santos, I Moreira & I Doadrio (2005) Chondrostoma occidentale, a new species of the genus Chondrostoma Agassiz, 1832 (Actynopterigii, Cyprinidae) from western Portugal. Graellsia 61(1): 19-29  


Sousa-Santos C (2001) O género Leuciscus (Pisces: Cyprinidae) na Região Litoral Oeste e suas afinidades biogeográficas. Análise morfológica, genética e do comportamento reprodutor. Relatório de Estágio da Licenciatura em Biologia Aplicada aos Recursos Animais variante Marinhos. Universidade de Lisboa. 56pp.

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