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"Science knows no country, because knowledge belongs to humanity,

and is the torch which illuminates the world"

Louis Pasteur

20. Frade PR, Bongaerts P, Englebert N, Rogers A, Gonzalez-Rivero M and Hoegh-Guldberg O (2018) Deep reefs of the Great Barrier Reef offer limited thermal refuge during mass coral bleaching.

Nature Communications 9 (1). doi: 10.1038/s41467-018-05741-0

19. Benzoni F, Arrigoni R, Berumen M, Taviani M, Bongaerts P and Frade PR (2018) Morphological and genetic divergence between the Mediterranean and Caribbean populations of Madracis pharensis (Heller 1868): too much for one species?

Zootaxa 4471 (3): 473–492. doi: 10.11646/zootaxa.4471.3.3

18. van Oppen MJH,  Frade PR, Bongaerts P, Peplow LM, Boyd S, Nim H and Bay LK (2018) Adaptation to reef habitats through selection on the coral animal and its associated microbiome.

Molecular Ecology 27: 2956-2971. doi: 10.1111/mec.14763

17. Glasl B, Bourne DG, Frade PR and Webster NS (2018) Establishing microbial baselines to identify indicators of coral reef health.

Microbiology Australia 10.1071/MA18011

16. Glasl B, Bongaerts P, Elisabeth NH, Hoegh-Guldberg O, Herndl GJ and Frade PR (2017) Microbiome variation in corals with distinct depth distribution ranges across a shallow-mesophotic gradient (15-85 m).

Coral Reefs 36 (2): 447-452. doi: 10.1007/s00338-016-1517-x

15. Goldammer H, Hollergschwandtner E, Elisabeth NH, Frade PR, and Reipert S (2016) Automated freeze substitution of algae accelerated by a novel agitation module.

Protist 167: 369-376. doi: 10.1016/j.protis.2016.06.002

14. Glasl B, Herndl GJ and Frade PR (2016) The microbiome of coral surface mucus has a key role in mediating holobiont health and survival upon disturbance.

The ISME Journal, advance online publication, 8 March 2016. doi: 10.1038/ismej.20169

13. Frade PR, Roll K, Bergauer K and Herndl GJ (2016) Archaeal and Bacterial communities associated with the surface mucus of Caribbean corals differ in their degree of host specificity and community turnover over reefs.

PLoS ONE 01/2016, 11(1): e0144702. doi: 10.1371/journal.pone.0144702

12. Frade PR, Schwaninger V, Glasl B, Sintes E, Hill RW, Simó R and Herndl GJ (2015) Dimethylsulfoniopropionate in corals and its interrelations with bacterial assemblages in coral surface mucus.

Environmental Chemistry 13(2) 252-265. doi: 10.1071/EN15023

11. Bongaerts P, Carmichael M, Hay KB, Tonk L, Frade PR and Hoegh-Guldberg O (2015) Prevalent endosymbiont zonation shapes the depth distributions of scleractinian coral species.

Royal Society Open Science 2: 140297. doi: 10.1098/rsos.140297

Peer-review articles


10. Bongaerts P, Frade PR, Hay KB, Englebert N, Latijnhouwers K, Bak RPM, Vermeij MJA and Hoegh-Guldberg O (2015) Deep down on a Caribbean reef: lower mesophotic depths harbor a specialized coral-endosymbiont community.

Scientific Reports 5: 7652. doi: 10.1038/srep07652

9. Bongaerts P, Frade PR, Ogier JJ, Hay KB, van Bleiswijk J, Englebert N, Vermeij MJA, Bak RPM, Visser PM and Hoegh-Guldberg O (2013) Sharing the slope: depth partitioning of Agariciid corals and their associated Symbiodinium across shallow and mesophotic habitats (0-60 m) on a Caribbean reef.

BMC Evolutionary Biology 13: 205. doi: 10.1186/1471-2148-13-205

8. Filatov MV, Frade PR, Bak RPM, Vermeij MJA and Kaandorp JA (2013) Comparison between colony morphology and molecular phylogeny in the Caribbean scleractinian coral genus Madracis.

PLoS ONE 8 (8): e71287. doi: 10.1371/journal.pone.0071287

7. Vermeij MJA, Frade PR and Bak RPM (2013) Zooxanthellae presence acts as a settlement cue for aposymbiotic planulae of the Caribbean coral Montrastraea faveolata.

Caribbean Journal of Science 47: 31-36

6. Frade PR, Reyes-Nivia MC, Faria J, Kaandorp JA, Luttikhuizen PC and Bak RPM (2010) Semi-permeable species boundaries in the coral genus Madracis: The role of introgression in a brooding coral system.

Molecular Phylogenetics and Evolution 57: 1072-1090

5. Hill RW, Li C, Jones AD, Gunn JP and Frade PR (2010) Abundant betaines in reef-building corals and ecological indicators of a photoprotective role.

Coral Reefs 29: 869-880

4. Frade PR, de Jongh F, Vermeulen F, van Bleijswijk J and Bak RPM (2008) Variation in symbiont distribution between closely related coral species over large depth ranges.

Molecular Ecology 17: 691-703

3. Frade PR, Bongaerts P, Winkelhagen AJS, Tonk L and Bak RPM (2008) In situ photobiology of corals over large depth ranges: a multivariate analysis on the roles of environment, host, and algal symbiont.

Limnology and Oceanography 53: 2711-2723

2. Frade PR, Englebert N, Faria J, Visser PM and Bak RPM (2008) Distribution and photobiology of Symbiodinium types in different light environments for three colour morphs of the coral Madracis pharensis: is there more to it than total irradiance?

Coral Reefs 27: 913-925

1. Vermeij MJA, Frade PR, Jacinto RIR, Debrot AO and Bak RPM (2007) Habitat-related differences in population structure are associated with coral reproductive mode: a comparison between fringing reef and inland bay coral communities.

Marine Ecology Progress Series 351: 91-102

Thesis / Books
2. Frade PR, Bongaerts P, Baldwin CC, Trembanis AC, Bak RPM and Vermeij MJA (in press) The mesophotic coral ecosystems of Bonaire and Curaçao. In: Loya Y, Puglise K and Bridge T (in press) Coral Reefs of the World Series: Mesophotic Coral Ecosystems. Springer
1. Frade PR (2009) Corals through the light - Phylogenetics, functional diversity and adaptive strategies of coral-symbiont associations over a large depth range.
PhD Thesis, University of Amsterdam, 176 pp. (ISBN: 978-90-9024362-7) (

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