Long Term Ecological Monitoring Projects

BALA - Biodiversity of Arthropods the Laurissilva of Azores

Experimental Design:

Permanent PLOT:

Team Leaders

  • Paulo A. V. Borges


  • Ana M. C. Santos
  • Carla José Azevedo Rego
  • François Rigal
  • Isabel R. Amorim
  • Joaquín Hortal
  • Konstantinos Triantis
  • Mário Rui Canela Boieiro
  • Paulo A. V. Borges
  • Pedro Cardoso
  • Sofia Terzopoulou

Since 1999, our knowledge of arthropods in native forests of the Azores has improved greatly. Under the BALA project (Biodiversity of Arthropods of Laurisilva of the Azores), an extensive standardised sampling protocol was employed in most of the native forest cover of the Archipelago. Additionally, in 2003 and 2004, more intensive sampling was carried out in several fragments, resulting in nearly a doubling of the number of samples collected.

A total of 6,770 samples from 100 sites distributed amongst 18 fragments of seven islands have been collected, resulting in almost 140,000 specimens having been caught. Overall, 452 arthropod species were recorded.

Almost ten years later (2010-2012) the same basic sampling protocol was applied. More precisely, fifteen native forest fragments distributed across seven of the nine islands were sampled in this study. Altogether, they represent most of the native forest cover of the Azores. During the summer of 2010, transects of 150 m long and 5 m wide were established again in 30 ofthe 100 original BALA sites.

Specifically we want to examine:  

  • Whether species richness and abundance distribution have changed after 10 years for both indigenous and non-indigenous species assemblages?
  • If changes are detected, whether they could be related to human-mediated factors such as area of the fragment or forest disturbance?
  • Whether species-specific temporal changes could be predicted by macroecological and functional attributes of the species?



Direcção Regional dos Recursos Florestais (Project 17.01-080203) (1999-2004)

FCT- PTDC/BIA-BEC/100182/2008 – “Predicting extinctions on islands: a multi-scale assessment” (2010-2013)

FCT - PTDC/BIA-BEC/104571/2008 – “What can the Macaronesian islands teach us about speciation? A case study of Tarphius beetles and Hipparchia butterflies” (2010-2013)

FCT- PTDC/BIA-BIC/119255/2010 – “Biodiversity on oceanic islands: towards a unified theory” (2012-2015)

DRCT- M2.1.2/I/027/2011 - Mapping coastal and marine biodiversity of the Azores - ATLANTIS-MAR(2012-2014)

DRCT- (M3.1.7/F/002/2011) Impact of Land-Use changes in the arthropod fauna of the Azores (2012-2015).



1)      Joaquin Munoz Hortal (2005-2007); FCT – BPD/20809/2004) with the project "Assessing the potential of environmental- and assemblage-based approaches to conservation biogeography" 

2)      Pedro Cardoso (2007-2008); DRCT – Medida 1.1.2/FRCT – with the project: “Agriculture, habitat fragmentation, indicator species and conservation of endemic fauna and flora in the Azores – the 2010 Target”

3)      Isabel Amorim do Rosário (2007-2010) FCT – SFRH/BPD/29578/2006 with the project: Dispersal and speciation in the archipelagoes of Macaronesia: a multi-gene analysis of Tarphius beetles and Hipparchia butterflies”.

4)      Clara Gaspar (2009-2011); DRCT – Medida 1.1.2/FRCT – with the project: “Agriculture, habitat fragmentation, indicator species and conservation of endemic fauna and flora in the Azores – the 2010 Target”

5)      Carla Rego (2010-2012); FCT - SFRH/BPD/66934/2009, with the project: “Predicting extinction risk in island endemics: integrating ecology and conservation genetics in a multi-scale approach” 

6)      Konstantinos Triantis (2008-2012) FCT – SFRH/BPD/44306/2008 with the project:Native and invasive species on islands: the functionality perspective” 

7)      Pedro Cardoso (2009-2012); FCT- SFRH/BPD/40688/2007, with the project: “Adapting IUCN conservation categories to Invertebrates” 

8)  François Rigal (2010-2013); FCT - PTDC/BIA-BEC/100182/2008 – with the project: “Predicting extinctions on islands: a multi-scale assessment” 

9)  Margarita Florencio (2012-2015); DRCT – Medida 1.1.2/FRCT – with the project: “Agriculture, habitat fragmentation, indicator species and conservation of endemic fauna and flora in the Azores – the 2010 Target”,

10)  François Rigal (2012-2015); FCT - PTDC/BIA-BIC/119255/2010– with the project: “Biodiversity on oceanic islands: towards a unified theory”

11Carla Rego (2013-2016); FCT - SFRH/BPD/91357/2012, with the project: “Radiation of phytophagous insects on oceanic islands: analysis of host associated genetic divergence in a speciose group of endemic bugs”.

12) Mário Boieiro (2013-2016); FCT - SFRH/BPD/86215/2012, with the project: “Host specialization in insect-plant food webs from a relict forest: assessing the roles of phylogenetic, ecological and functional determinants as drivers of community assembly”

13) Isabel Amorim do Rosário (2015-2018); FCT - SFRH/BPD/102804/2014, with the project "Biodiv-WhoCares? Communicating the value of unique biodiversity: targeting non-traditional audiences in the Azores"


1) Clara Gaspar (2007) Ph.D. in Biodiversity by the University of Sheffied, U.K, with  co-supervision of Kevin J. Gaston. Thesis:Arthropod diversity and conservation planning in native forests of the Azores archipelago

2) Sofia Terzopoulou (2011-2015) Ph.D. in Ecology by the Univ. of Athens also under supervision of Prof. Kostas A. Triantis and Aris Paramakelis (Univ. Athens): “Species richness, functional, and genetic diversity of Azorean beetles: a holistic approach

3) Rui Nunes (2012-2015) Ph.D. in Landscape Interdisciplinary Management by the Univ. of Azores and Univ. Évora under supervision of Prof. Paulo Borges, Prof. Pedro Cardoso and Prof. António Onofre Soares with the project Whole Azorean Arthropod diversity: understanding the trophic relations and functional diversity at a plot scale”.

We are grateful to all researchers that collaborated in the field and laboratory during the BALA project: Álvaro Vitorino, Anabela Arraiol, Ana Rodrigues, Carlos Aguiar,Francisco Dinis, Genage André, Emanuel Barcelos, Hugo Mas,  João Amaral, Joaquín Hortal, Lara Dinis, Paula Gonçalves, Sandra Jarroca and Luís Vieira.

The Azores Forest Services provided local support in each island in the years of 1999-2004. Since 2005 most logistical support was given by the Natural Parks of each Island.

Acknowledgments are due to all the taxonomists who assisted in the identification of the morphotypes: Andrew Polaszek, António Bivar de Sousa, Artur Serrano, Arturo Baz, Fernando Ilharco, Henrik Enghoff, Jordi Ribes, José Quartau, Jörg Wunderlich,  Ole Karsholt, Richard Strassen, Volker Manhert.

Ribeiro, S.P., Borges, P.A.V., Gaspar, C., Melo, C., Serrano, A.R.M., Amaral, J., Aguiar, C., André, G. & Quartau, J.A. (2005). Canopy insect herbivores in the Azorean Laurisilva forests: key host plant species in a highly generalist insect community. Ecography, 28: 315-330. DOI: 10.1111/j.0906-7590.2005.04104.x(IF2013:4.207; Q1 Ecology)  (

Borges, P.A.V., Aguiar, C., Amaral, J., Amorim, I.R., André, G., Arraiol, A.,. Baz A., Dinis, F., Enghoff, H., Gaspar, C., Ilharco, F., Mahnert, V., Melo, C., Pereira, F., Quartau, J.A., Ribeiro, S., Ribes, J., Serrano, A.R.M., Sousa, A.B., Strassen, R.Z., Vieira, L., Vieira, V., Vitorino, A. & Wunderlich,J. (2005). Ranking protected areas in the Azores using standardized sampling of soil epigean arthropods. Biodiversity and Conservation, 14: 2029-2060.DOI: 10.1007/s10531-004-4283-y (IF2013: 2.065; Q2 Ecology) (

Borges, P.A.V., Serrano, A.R.M. & Quartau, J.A. (2000). Ranking the Azorean Natural Forest Reserves for conservation using their endemic arthropods. Journal of Insect Conservation, 4: 129-147. DOI: 10.1023/A:1009629012205 (IF2013: 1.789; Q1 Entomology) (

Borges, P.A.V., Lobo, J.M., Azevedo, E.B., Gaspar, C., Melo, C. & Nunes, L.V.(2006). Invasibility and species richness of island endemic arthropods: a general model of endemic vs. exotic species. Journal of Biogeography, 33: 169-187. DOI:10.1111/j.1365-2699.2005.01324.x (IF2013: 4.969; Q1 Ecology) (

Hortal, J., Borges, P.A.V. & Gaspar, C. (2006). Evaluating the performance of species richness estimators: Sensitivity to sample grain size. Journal of Animal Ecology, 75:274-287. DOI:10.1111/j.1365-2656.2006.01048.x (IF2013:4.726; Q1 Ecology)

Gaston, K.J., Borges, P.A.V., He, F &. Gaspar, C. (2006). Abundance, spatial variance and occupancy: arthropod species distribution in the Azores. Journal of Animal Ecology, 75: 646-656.DOI 10.1111/j.1365-2656.2006.01085.x (IF2013:4.726; Q1 Ecology) (

Cardoso, P., Borges, P.A.V.& Gaspar, C. (2007). Biotic integrity of the arthropod communities in the natural forests of Azores. Biodiversity and Conservation 16: 2883-2901.DOI: 10.1007/s10531-006-9078-x ((IF2013: 2.065; Q2 Ecology) (

Borges, P.A.V. & Wunderlich, J. (2008). Spider biodiversity patterns and their conservation in the Azorean archipelago, with description of new taxa. Systematics and Biodiversity, 6: 249-282. DOI: 10.1017/S1477200008002648. (IF2013: 2.153; Q2 Biodiversity and Conservation) (

Ribes, J & Borges, P.A.V. (2001). A new subspecies of Orthotylus junipericola Linnavuori, 1965 (Heteroptera; Miridae) from the Azores. Arquipélago Life and Marine Sciences 18A: 1-4. (

Platia, G. & Borges, P.A.V. (2002) Description of a new species of Athous and record of the female of A. azoricus Platia & Gudenzi from the Azores (Coleoptera, Elateridae). Elytron, 16: 91-95. (

Quartau, J.A. & Borges, P.A.V. (2003). A new species of the genus Aphrodes Curtis from the Azores (Hemiptera, Cicadellidae). Bocagiana, 213: 1-11. (

Gaspar, C.,Borges, P.A.V. & Gaston, K.J. (2008). Diversity and distribution of arthropods in native forests of the Azores archipelago. Arquipélago Life and marine Sciences, 25: 1-30. (

Jiménez-Valverde, A., Diniz, F. Azevedo, E.B. & Borges, P.A.V. (2009). Species distribution models do not account for abundance: the case of arthropods in Terceira Island. Annales Zoologici Fennici, 46: 451-464.(IF2013: 1.03; Q4 Ecology) (

Cardoso, P., Borges, P.A.V. & Veech, J.A. (2009). Testing the performance of beta diversity measures based on incidence data: the robustness to undersampling. Diversity and Distributions, 15: 1081-1090. Doi: 10.1111/j.1472-4642.2009.00607.x  (IF2013: 5.469; Q1 Ecology) (

Cardoso, P., Lobo, J.M., Aranda, S.C., Dinis, F., Gaspar, C. & Borges, P.A.V. (2009). A spatial scale assessment of habitat effects on arthropod communities of an oceanic island. Acta Oecologica, 35: 590-597. Doi:10.1016/j.actao.2009.05.005 (IF2013: 1.841; Q3 Ecology) (

Gaspar, C.,Gaston, K.J. & Borges, P.A.V. (2010). Arthropods as surrogates of diversity at different spatial scales. Biological Conservation, 143: 1287–1294. Doi: 10.1016/j.biocon.2010.03.007. (IF2013: 4.036; Q1 Ecology) (

Borges, P.A.V., Ugland, K.I, Dinis, F.O. & Gaspar, C. (2008). Insect and spider rarity in an oceanic island (Terceira, Azores): true rare and pseudo-rare species. In: S. Fattorini (Ed.). Insect Ecology and Conservation. pp. 47-70. Research Signpost, Kerala, India. ISBN: 978-81-308-0297-8, pp. 317. (

Triantis, K.A., Borges, P.A.V., Ladle, R.J., Hortal, J., Cardoso, P., Gaspar, C., Dinis, F., Mendonça, E., Silveira, L.M.A.,  Gabriel, R., Melo, C., Santos, A.M.C., Amorim, I.R., Ribeiro, S.P., Serrano, A.R.M., Quartau, J.A. & Whittaker, R.J. (2010). Extinction debt on oceanic islands. Ecography, 33: 285-294. DOI: 10.1111/j.1600-0587.2010.06203.x. (IF2013:4.207; Q1 Ecology) (

Meijer, S.S., Whittaker, R.J. & Borges, P.A.V. (2011). The effects of land-use change on arthropod richness and abundance on Santa Maria Island (Azores): unmanaged plantations favour endemic beetles.  Journal of Insect Conservation, 15: 505-522. DOI 10.1007/s10841-010-9330-2. (IF2013: 1.789; Q1 Entomology) (

Gaspar, C.,Gaston, K.J., Borges, P.A.V. & Cardoso, P.(2011). Selection of priority areas for arthropod conservation in the Azores archipelago. Journal of Insect Conservation, 15: 671–684. DOI: 10.1007/s10841-010-9365-4. (IF2013: 1.789; Q1 Entomology) (

Fattorini, S., Cardoso, P., Rigal, F. & Borges, P.A.V. (2012). Use of Arthropod Rarity for Area Prioritisation: Insights from the Azorean Islands. PLoS ONE 7(3):e33995. doi:10.1371/journal.pone.0033995. (IF2013: 3.534; Q1 Multidisciplinary Sciences) (

Amorim, I., Emerson, B.C, Borges, P.A.V. & Wayne, R.K. (2012). Phylogeography and molecular phylogeny of Macaronesian island Tarphius (Coleoptera: Zopheridae): why so few species in the Azores?. Journal of Biogeography, 39: 1583–1595. DOI:10.1111/j.1365-2699.2012.02721.x(IF2013: 4.969; Q1 Ecology) (

Fattorini, S. & Borges, P.A.V.(2012). Biogeographical kinetics on a island volcano (Capelinhos, Azores): fast colonization rates and dominance of arthropod exotic species. Insect Conservation and Diversity, 5: 331–345. DOI: 10.1111/j.1752-4598.2011.00169.x (IF2013: 1.937; Q1 Entomology) (

Rigal, F., Whittaker, R.J., Triantis, K.A. & Borges, P.A.V. (2013). Integration of non-indigenous species within the interspecific abundance-occupancy relationship. Acta Oecologica, 48: 69–75. DOI: 10.1016/j.actao.2013.02.003 (IF2013: 1.841; Q3 Ecology) (

Cardoso, P., Rigal, F., Fattorini, S., Terzopoulou, S. & Borges, P.A.V. (2013). Integrating Landscape Disturbance and Indicator Species in Conservation Studies. PLoS ONE, 8: e63294. DOI:10.1371/journal.pone.0063294(IF2013: 3.534; Q1 Multidisciplinary Sciences) (

Carvalho, J.C.,Cardoso, P., Borges, P.A.V., Schmera, D. & Podani, J.(2013).Measuring fractions of beta diversity and their relationships to nestedness: a theoretical and empirical comparison of novel approaches. Oikos, 122: 825–834. DOI: 10.1111/j.1600-0706.2012.20980.x. (IF2013: 3.559; Q1 Ecology) ((

Florencio, M., Cardoso, P., Lobo, J.M., Azevedo, E.B. & Borges, P.A.V. (2013). Arthropod assemblage homogenization in oceanic islands: the role of exotic and indigenous species under landscape disturbance. Diversity and Distributions, 19: 1450-1460. DOI: 10.1111/ddi.12121(IF2013: 5.469; Q1 Ecology) (

Crespo, L.C., Bosmans, R., Cardoso, P.  & Borges, P.A.V. (2013). On the endemic spider species of the genus Savigniorrhipis Wunderlich, 1992 (Araneae: Linyphiidae) in the Azores (Portugal), with description of a new species. Zootaxa, 3745: 330-342.DOI:10.11646/zootaxa.3745.3.2(IF2013: 1.06; Q2 Zoology) (

Cardoso, P., Rigal, F., Carvalho, J.C., Fortelius, M., Borges, P.A.V., Podani, J.&Schmera, D. (2014). Partitioning taxon, phylogenetic and functional beta diversity into replacement and richness difference components. Journal of Biogeography, 41: 749–761.DOI: 10.1111/jbi.12239. (IF2013: 4.969; Q1 Ecology) (

Cardoso, P., Rigal, F., Borges, P.A.V. &Carvalho, J.C. (2014). A new frontier in biodiversity inventory: a proposal for estimators of phylogenetic and functional diversity. Methods in Ecology and Evolution, 5: 452–461. DOI: 10.1111/2041-210X.12173 (IF2013: 5.322; Q1 Ecology) (

Matthews, T.J., Borges, P.A.V. & Whittaker, R.J. (2014). Multimodal species-abundance distributions: a deconstruction approach reveals the processes behind the pattern. Oikos, 123: 533–544.DOI: 10.1111/j.1600-0706.2013.00829.x(IF2013: 3.559; Q1 Ecology) (

Crespo, L.C., Bosmans, R., Cardoso, P. & Borges, P.A.V. (2014). On three endemic species of the linyphiid spider genus Canariphantes Wunderlich, 1992 (Araneae, Linyphiidae) from the Azores archipelago. Zootaxa, 3841:403–417. DOI: 10.11646/zootaxa.3841.3.5 (IF2013: 1.06; Q2 Zoology) (

Matthews, T.J., Borregaard, M.K., Ugland, K., Borges, P.A.V., Rigal, F., Cardoso, P. & Whittaker, R.J. (2014). The gambin model provides a superior fit to species abundance distributions with a single free parameter: evidence, implementation and interpretation. Ecography, 37: 1002–1011, DOI: 10.1111/ecog.00861 (IF2013:4.207; Q1 Ecology) (

Whittaker, R.,Rigal, F., Borges, P.A.V., Cardoso, P., Terzopoulou, S., Casanoves, F., Pla, L., Guilhaumon, F., Ladle, R. & Triantis, K.A. (2014). Functional biogeography of oceanic islands and the scaling of functional diversity in the Azores. PNAS - Proceedings of the National Academy of Sciences, 111: 13709–13714.  DOI: 10.1073/pnas.1218036111 (IF2013: 9.809; Q1 Multidisciplinary Sciences) (

Borges, P.A.V., Gaspar, C.S., Santos, A.M.C, Ribeiro, S.P., Cardoso, P., Triantis, K. & Amorim, I.R. (2011). Patterns of colonization and species distribution for Azorean arthropods: evolution, diversity, rarity and extinction.In: Martins, A.M.F., & M.C. Carvalho (Eds.), Celebrating Darwin: Proceedings of the Symposium "Darwin's Mistake and what we are doing to correct it", Ponta Delgada, 19-22 September, 2009. Açoreana, Supl. 7: 93-123. (

Gaspar, C., Borges, P.A.V.& Gaston, K.J. (2014). Towards systematic conservation planning in the Azores. Ecologi@, 7: 40-49. (

Gaspar, C., Cardoso, P., Borges, P.A.V. & Gaston, K.J. (2014). Efficiency of sampling methods and effort to assess arthropod diversity in Azorean native forests. Arquipélago Life and Marine Sciences, 31: 21-36. (

Florencio, M., Lobo, J.M., Cardoso, P.,Almeida-Neto, M. & Borges, P.A.V. (2015). The colonisation of exotic species does not have to trigger faunal homogenisation: lessons from the assembly patterns of arthropods on oceanic islands.PLoS ONE10(5): e0128276. DOI: 10.1371/journal.pone.0128276(IF2013: 3.534; Q1 Multidisciplinary Sciences)

Terzopoulou, S., Rigal, F., Whittaker, R.J., Borges, P.A.V. & Triantis, K.A. (2015). Drivers of extinction: the case of Azorean beetles. Biology Letters, 11: 1-4. DOI: 10.1098/rsbl.2015.0273 (IF2013: 3.425; Q1 Ecology)