José Ricardo Paula, PhD

Principal Investigator

I am a behavioral ecologist who is fascinated by the intricacies of social interactions in nature. I'm an Assistant Researcher at the University of Lisbon/MARE and a visiting researcher at the Hawai'i Institute of Marine Biology - University of Hawai'i (at Liz Madin lab).  I've done my PhD at the University of Lisbon with Rui Rosa and a Post-doc at the University of Hong Kong with Celia Schunter

Most of my research has been focused on cleaner fish, small aquatic animals that provide a valuable service to other species by removing parasites and dead tissue from their skin. 

I'm particularly interested in the relevance of animal cognition at a ecosystem level, how species interact, and how their behavior and cognition have evolved over time.

My research also focuses on understanding the neural basis of fish cognition and how it has evolved over time.  For this, I use a combination of techniques, including neurogenomics, neuroanatomy, and behavioral analysis. 

I love to hike, cook, travel, and obviously scuba dive! I can also be really clumsy with coffee mugs next to keyboards (accidents have happened).


Google Scholar:


51)  B. P. Pereira, S. Neff, F. O. Borges, E. Otjacques, G. Barreto, M. Ranucci, M. Court, R. Rosa, T. Repolho, J. R. Paula (2023) Transgenerational exposure to deoxygenation and warming disrupts mate-detection in Gammarus locusta. Behavioral Ecology (in press)

50)  J. L. Röckner; V.M. Lopes; J.R. Paula; M.R. Pegado; M.C. Seco; M. Diniz; T. Repolho; R. Rosa (2023) Octopus “crawling" on land: physiological and biochemical responses of Octopus vulgaris to emersion. Marine Biology (in press)

49) M. Gomes, V.M. Lopes, M.G. Mai, J.R. Paula, R. Bispo, H. Batista, C. Barraca, N. Baylina, R. Rosa, M.S. Pimentel (2023) Impacts of acute hypoxia on the short-snouted seahorse metabolism and behaviour. Science of The Total Environment, 904, 166893. DOI: 10.1016/j.scitotenv.2023.166893

48) J. Kang, S Ramirez-Calero, J.R. Paula, Y. Chen, C. Schunter (2023) Gene losses, parallel evolution and heightened expression confer adaptations to dedicated cleaning behaviour. BMC Biology 21, 180 DOI: 10.1186/s12915-023-01682-3

47) S. Ramirez-Calero, J.R.Paula, E. Otjacques, T. Ravasi, R. Rosa, C. Schunter (2023) Neuromolecular responses in disrupted mutualistic cleaning interactions under future environmental conditions. bioRxiv 2023.07.20.549851 ; DOI: 10.1101/2023.07.20.549851 

46) J.R. Paula, X. Wang, C. Xu, D.C. Moreira (2023) Editorial: The defense responses of aquatic animals to the environment. Frontiers in Physiology 14, 1231014.DOI: 10.3389/fphys.2023.1231014

45) M. Court, M. Macau, M. Ranucci, T. Repolho, V. M. Lopes, R. Rosa, J.R. Paula (2023) Oxygen loss compromises the survival and cognition of a coastal cephalopod. bioRxiv 2023.06.03.543560; DOI: 10.1101/2023.06.03.543560

44) R. Rosa, E. Nunes, V. Pissarra, C.P. Santos, J Varela, M.   Baptista, J. Castro, J.R. Paula, T. Repolho, T. Marques, R. Freitas, C. Frazão Santos (2023) Evidence for the first multi-species shark nursery area in Atlantic Africa (Boa Vista Island, Cabo Verde) Frontiers in Marine Science, 10,1077748. DOI: 10.3389/fmars.2023.1077748

43) M Dias, J.R Paula, P Pousão-Ferreira, S Casal, R Cruz, S Cunha, R. Rosa, A. Marques, P. Anacleto, A. Maulvaut (2023) Combined effects of climate change and BDE-209 dietary exposure on the behavioural response of the white seabream, Diplodus sargus. Science of the Total Environment 881:13400. DOI:10.1016/j.scitotenv.2023.163400

42) J. Varela, S. Martins, M. Court, CP Santos, JR Paula, IJ Ferreira, M. Diniz, T. Repolho, R. Rosa (2023) Impacts of Deoxygenation and Hypoxia on Shark Embryos Anti-Predator Behavior and Oxidative Stress. Biology 12(4):577. DOI:10.3390/biology12040577

41) J.R. Paula, L. Cascalheira, R. Oliveira, E. Otjacques, C. Frazão-Santos, R. Beldade, S. C. Mills, R. Rosa (2023) GABAergic role in the disruption of wild cleaner fish behaviour under high CO2. Animal Behaviour 195, 77-84. DOI: 10.1016/j.anbehav.2022.11.0

40) J. Coelho, M. Court, E. Otjacques, V.M. Lopes, J.R. Paula, T. Repolho, M. Diniz, R. Rosa (2023) Effects of tidal emersion and marine heatwaves on cuttlefish early ontogeny. Marine Biology  170, 3DOI: 10.1007/s00227-022-04150-8

39) F.O. Borges, M. Guerreiro, C. P. Santos, J.R. Paula, R. Rosa (2022) Projecting future climate change impacts on the distribution of the ‘Octopus vulgaris species complex’. Frontiers in Marine Science,  9, 1018766. DOI: 10.3389/fmars.2022.1018766

38) M. Court, J. R. Paula, M. Macau, E. Otjacques, T. Repolho, R. Rosa, V. M. Lopes (2022) Camouflage and exploratory avoidance of newborn cuttlefish under warming and acidification. Biology 11, 1394. DOI: 10.3390/biology11101394

37) M. Deleja, J.R. Paula, T. Repolho, M. Franzitta, M. Baptista, V. Lopes, S. Simão, V. F. Fonseca, B. Duarte, R. Rosa (2022) Effects of hypoxia on coral photobiology and oxidative stress. Biology 11, 1068. DOI: 10.3390/biology11071068

36) S. Ramírez-Calero, J. R. Paula, E. Otjacques, R. Rosa, T. Ravasi, C. Schunter (2022) Neuro-molecular characterization of fish cleaning interactions. Scientific Reports 12, 8468. DOI: 10.1038/s41598-022-12363-6 

35) J. R. Paula, T. Repolho, A.S. Grutter, R.Rosa (2022) Access to cleaning services alters fish physiology under parasite infection and ocean acidification. Frontiers in Physiology 13, 859556. DOI: 10.3389/fphys.2022.859556 Rising stars in aquatic physiology 2022

34) B. Duarte, T. Repolho, J.R. Paula, I. Caçador, A. Matos, R. Rosa (2022) Ocean acidification alleviates dwarf eelgrass (Zostera noltii) lipid landscape remodeling under warming stress. Biology, 11, 780. DOI: 10.3390/biology11050780

33) M. Franzitta, T. Repolho, J. R. Paula, I. Caçador, A.R. Matos, R. Rosa, B. Duarte (2021) Dwarf eelgrass (Zostera noltii) fatty acid remodelling induced by climate change. Estuarine, Coastal and Shelf Science, 261, 107546. DOI: 10.1016/j.ecss.2021.107546

32)  F.O. Borges, C. Santos, J. R. Paula, E. Mateos-Naranjo, S. Redondo-Gómez, J.B. Adams, I. Caçador, V.F. Fonseca, P. Reis-Santos, B. Duarte, R. Rosa (2021) Invasion and extirpation potential of native and invasive Spartina species under climate change. Frontiers in Marine Science 8:696333. DOI:10.3389/fmars.2021.696333

31) J. R. Paula, D. Sun, V. Pissarra, P. Narvaez, A. Grutter, R. Rosa, P. Sikkel (2021) The role of corals on the abundance of a fish ectoparasite in the Great Barrier Reef Coral Reefs, 40, 535–542. DOI:10.1007/s00338-021-02051-8. 

30)  J. R. Paula (2020) Lockdowns due to COVID-19 threaten PhD students’ and early-career researchers’ careers. Nature Ecology and Evolution 4, 999. DOI: 10.1038/s41559-020-1231-5. 

29) E. Otjacques, T.  Repolho, J. R. Paula, S. Simão, M. Baptista, R. Rosa (2020) Cuttlefish buoyancy control in response to food availability and ocean acidification. Biology 9, 147. DOI:10.3390/biology9070147. 

28) M. Corriere, M. Baptista, J.R. Paula, T. Repolho, R. Rosa, P. Costa, L. Soliño (2020) Impaired fish swimming performance following dietary exposure to the marine phycotoxin okadaic acid. Toxicon 179: 53-59. DOI:10.1016/j.toxicon.2020.02.022. 

27) J.R. Paula, E. Otjacques, C. Hildebrandt, A.S. Grutter, R. Rosa (2020) Ocean acidification does not affect fish ectoparasite survival. Oceans 1: 27-33. DOI: 10.3390/oceans1010003 

26) J. R. Paula, M. Baptista, F. Carvalho, T. Repolho, R. Bshary, R. Rosa (2019) The past, present and future of cleaner fish cognitive performance as a function of CO2 levels. Biology Letters, 15, 20190618. DOI: 10.1098/rsbl.2019.0618 Cover of Biology Letters

25) J. R. Paula, T. Repolho, M. Pegado, P-O Thörnqvist, R. Bispo, S. Winberg, P. Munday, R. Rosa (2019) Neurobiological and behavioural responses of cleaning mutualisms to ocean warming and acidification. Scientific Reports 9, 12728. DOI:10.1038/s41598-019-49086-0

24) A. Rodrigues, F. O. Borges, V. Pissarra, A. Maulvault, J. R. Paula, R. Bispo, R. Rosa (2019) First indication of deleterious impacts in white-seabream larvae (Diplodus sargus) survival and behaviour following acute venlafaxine exposure. Ecotoxicology 28, 612-618. DOI: 10.1007/s10646-019-02057-7 

23) M. Baptista, O. Azevedo, C. Figueiredo, J. R. Paula, M. Santos, N. Queiroz, R. Rosa, J. Raimundo (2019) Body size and season influence elemental composition of tissues in ocean sunfish Mola mola juveniles. Chemosphere 223, 714-722. DOI: 10.1016/j.chemosphere.2019.02.061 

22)  F. Borges, C. Santos, E. Sampaio, C. Figueiredo, J. R. Paula, C. Antunes, R. Rosa, T. F. Grilo (2019) Ocean warming and acidification may challenge the riverward migration of glass eels. Biology Letters 15, 20180627. DOI: 10.1098/rsbl.2018.0627 

21) A. Maulvault, C. Camacho, V. Barbosa, R. Alves, P. Anacleto, S. Cunha, J. Fernandes, P. Pousão-Ferreira, J. R. Paula,R. Rosa, M. Diniz, A. Marques (2019) Bioaccumulation and ecotoxicological responses of juvenile white seabream (Diplodus sargus) exposed to triclosan, warming and acidification. Environmental Pollution 245, 427-442. DOI: 10.1016/j.envpol.2018.11.020 

20) M. Baptista, A. Couto, J. R. Paula, J. Raimundo, N. Queiroz, R. Rosa (2018) Seasonal variations in the abundance and body size distribution of the ocean sunfish Mola mola in coastal waters off southern Portugal. J. Marine Biological Association UK 99, 1197-1203. DOI: 10.1017/S002531541800111X

19) G. Dionísio, F. Faleiro, R. Bispo, A. Lopes, S. Cruz, J. R. Paula, T. Repolho, R. Calado, R. Rosa (2018) Distinct bleaching resilience of photosynthetic plastid-bearing mollusks under thermal stress and high CO2 conditions. Frontiers in Physiology 9:1675. DOI: 10.3389/fphys.2018.01675

18) L. Mincarelli, J. R. Paula, P. Pousão-Ferreira, R. Rosa, P. Costa (2018) Effects of acute waterborne exposure to harmful algal toxin domoic acid on foraging and swimming behaviours of fish early stages. Toxicon 156, 66-71. DOI: 10.1016/j.toxicon.2018.11.297 

17) A. Maulvault, L. Santos, J. R. Paula, C. Camacho, V. Pissarra, F. Fogaça, V. Barbosa, R. Alves, P. Pousão Ferreira, D. Barceló, S. Rodriguez-Mozaz, A. Marques, M. Diniz, R. Rosa (2018) Differential behavioural responses to venlafaxine exposure route, warming and acidification in juvenile fish (Argyrosomus regius). Science of The Total Environment 634:1136-1147. DOI: 10.1016/j.scitotenv.2018.04.015. 

16)  E. Sampaio, A. Lopes, S. Francisco, J. R. Paula, M. Pimentel, A. Maulvault, T. Repolho, T. Grilo, P. Pousão-Ferreira, A. Marques, R. Rosa (2017) Ocean acidification dampens physiological stress response to warming and contamination in a commercially-important fish (Argyrosomus regius). Science of The Total Environment 618:388-398. DOI: 10.1016/j.scitotenv.2017.11.059 

15) R. Rosa, J. Kelly, V. Lopes, J. R. Paula, J. Gonçalves, R. Calado, M. Norman, J. P. Barreiros (2017) Deep-sea seven-arm octopus hijacks jellyfish in shallow waters. Marine Biodiversity 49, 495-499. DOI: 10.1007/s12526-017-0767-3 

14) T. Repolho, B. Duarte, G. Dionísio, J. R. Paula, A. R. Lopes, I. C. Rosa, T. F. Grilo, I. Caçador, R. Calado, R. Rosa (2017) Seagrass ecophysiological performance under ocean warming and acidification. Scientific Reports 7:41443. DOI: 10.1038/srep41443 

13) E. Sampaio, A. L. Maulvault, V. Lopes, J. R. Paula, V. Barbosa, R. Alves, P. Pousão-Ferreira, T. Repolho, A. Marques, R. Rosa (2016) Habitat selection disruption and lateralization impairment of cryptic flatfish in a warm, acid, and contaminated ocean. Marine Biol. 163, 217. DOI: 10.1007/s00227-016-2994-8 

12) A. R. Lopes, M. Baptista, I. C. Rosa, G. Dionísio, J. Gomes-Pereira, J. R. Paula, C. Figueiredo, N. Bandarra, R. Calado, R. Rosa (2016) “Gone with the wind”: Fatty acid biomarkers and chemotaxonomy of stranded pleustonic hydrozoans (Velella velella and Physalia physalis). Biochemical Systematics and Ecology 66: 297-306. DOI: 10.1016/j.bse.2016.03.016 

11) R. Rosa, J. R. Paula, E. Sampaio, M. Pimentel, A. Lopes, M. Baptista, M. Guerreiro, C. Santos, D. Campos, V. Almeida-Val, R. Calado, M. Diniz, T. Repolho (2016) Neuro-oxidative damage and aerobic potential loss of sharks under elevated CO2 and warming. Marine Biol 163, 119. DOI: 10.1007/s00227-016-2898-7 

10) M. Soares, J. R. Paula, R. Bshary (2016) Serotonin blockade delays learning performance in a cooperative fish. Animal Cognition 19, 1027-1030. DOI: 10.1007/s10071-016-0988-z 

9)    J. Messias, J. R. Paula, A.S. Grutter, R. Bshary, M.C. Soares (2016) Dopamine disruption increases negotiation for cooperative interactions in a fish. Scientific Reports 6:20817. DOI: 10.1038/srep20817 

8)    V. Lopes, F. Faleiro, M. Baptista, M. Pimentel, J. R. Paula, A. Couto, N. Bandarra, P. Anacleto, A. Marques, R. Rosa (2015) Amino and fatty acid dynamics of octopus (Octopus vulgaris) early life stages under ocean warming. Journal of Thermal Biology 55: 30-38. DOI: 10.1016/j.jtherbio.2015.11.006 

7)    M. Payne, M. Barange, W. Cheung, B. MacKenzie, H. Batchelder, X. Cormon, T. Eddy, J. Fernandes, A. Hollowed, M. Jones, J. Link, P. Neubauer, I. Ortiz, A. Queiros, J. R. Paula (2015) Uncertainties in projecting climate change impacts in marine ecosystems. ICES Journal of Marine Science 73: 1272-1282. DOI:10.1093/icesjms/fsv231 

6)    E. Kulczykowska, S. Cardoso, M. Gozdowska, G. André, J. R. Paula, M. Ślebioda, R.F. Oliveira, M.C. Soares (2015) Brain levels of nonapeptides in four labrid fish species with different levels of mutualistic behavior. General and Comparative Endocrinology 222,99-105. DOI: 10.1016/j.ygcen.2015.06.005 

5)    S. Cardoso, A. Grutter, J. R. Paula, G. André, J. Messias, M. Gozdowska, E. Kulczykowska, M. Soares (2015) Forebrain neuropeptide regulation of pair association and behavior in cooperating cleaner fish. Physiology & Behavior145, 1-7. DOI: 10.1016/j.physbeh.2015.03.024 

4)    J. R. Paula, J.P. Messias, A.S. Grutter, R. Bshary, M. C. Soares (2015) The role of serotonin in the modulation of cooperative behavior. Behavioral Ecology 26,1005-1012. DOI: 10.1093/beheco/arv039 T. Cited: 42. Cover of Behavioral Ecology.

3)    F. Faleiro, M. Baptista, C. Santos, M. Aurélio, M. Pimentel, M. Pegado, J. R. Paula, R. Calado, T. Repolho, R. Rosa (2015) Seahorses under a changing ocean: The impact of warming and acidification on the behaviour and physiology of a poor-swimming bony-armoured fish. Conservation Physiology 3, cov009. DOI: 10.1093/conphys/cov009

2)    M. Soares, G.André, J. R. Paula (2015) Preliminary notes on brain weight variation across labrid species with different levels of cooperative behaviour. Cur. Zoo. 61, 274-280. DOI:10.1093/czoolo/61.2.274 

1)    R. Rosa, M. Baptista, V. Lopes, M. Pegado, J. R. Paula, K. Trübenbach, M. Leal, R. Calado, T. Repolho (2014) Early-life exposure to climate change impairs tropical shark survival. Proceedings of the Royal Society B 281,1793-1793. DOI: 10.1098/rspb.2014.1738