Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: Paulo A.V. Borges (pborges@uac.pt)
Academic editor: Pavel Stoev
Received: 27 Oct 2016 | Accepted: 09 Dec 2016 | Published: 22 Dec 2016
© 2016 Paulo Borges, Clara Gaspar, Luís Carlos Crespo, François Rigal, Pedro Cardoso, Fernando Pereira, Carla Rego, Isabel Amorim, Catarina Melo, Carlos Aguiar, Genage André, Enésima Mendonça, Sérvio Ribeiro, Joaquín Hortal, Ana Santos, Luís Barcelos, Henrik Enghoff, Volker Mahnert, Margarida Pita, Jordi Ribes, Arturo Baz, António Sousa, Virgílio Vieira, Jörg Wunderlich, Aristeidis Parmakelis, Robert Whittaker, José Alberto Quartau, Artur Serrano, Kostas Triantis
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Borges P, Gaspar C, Crespo L, Rigal F, Cardoso P, Pereira F, Rego C, Amorim I, Melo C, Aguiar C, André G, Mendonça E, Ribeiro S, Hortal J, Santos A, Barcelos L, Enghoff H, Mahnert V, Pita M, Ribes J, Baz A, Sousa A, Vieira V, Wunderlich J, Parmakelis A, Whittaker R, Quartau J, Serrano A, Triantis K (2016) New records and detailed distribution and abundance of selected arthropod species collected between 1999 and 2011 in Azorean native forests. Biodiversity Data Journal 4: e10948. https://doi.org/10.3897/BDJ.4.e10948
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In this contribution we present detailed distribution and abundance data for arthropod species identified during the BALA – Biodiversity of Arthropods from the Laurisilva of the Azores (1999-2004) and BALA2 projects (2010-2011) from 18 native forest fragments in seven of the nine Azorean islands (all excluding Graciosa and Corvo islands, which have no native forest left).
Of the total 286 species identified, 81% were captured between 1999 and 2000, a period during which only 39% of all the samples were collected. On average, arthropod richness for each island increased by 10% during the time frame of these projects. The classes Arachnida, Chilopoda and Diplopoda represent the most remarkable cases of new island records, with more than 30% of the records being novelties. This study stresses the need to expand the approaches applied in these projects to other habitats in the Azores, and more importantly to other less surveyed taxonomic groups (e.g. Diptera and Hymenoptera). These steps are fundamental for getting a more accurate assessment of biodiversity in the archipelago.
Azores, terrestrial arthropods, BALA project, laurissilva forest, Linnean, Wallacean and Prestonian shortfalls.
In 1999 a group of researchers from the University of the Azores and the University of Lisbon started a long-term (1999-2004) standardized sampling program to inventory the arthropod biodiversity in native forest remnants of the Azores - the BALA I project – Biodiversity of Arthropods from the Laurisilva of the Azores (
Eight years of standardized survey of the native forest in seven of the nine Azorean islands resulted in a major improvement on the knowledge of the Azorean arthropod fauna, in particular concerning Araneae, Opiliones, Pseudoscorpionida, Diplopoda, Chilopoda and Insecta (excluding Collembola, Diptera and Hymenoptera). As a consequence, several new endemic taxa were described for the archipelago (e.g.
During this period, two complete checklists of Azorean arthropod fauna were produced (
Area of study: The Azores
The remote Azores archipelago extends for 615 km in the North Atlantic Ocean (37-40 °N, 25-31 °W), 1584 km to the east (southern Europe) and 2150 km to the west (northern America) of the nearest mainland. It comprises nine main islands and some small islets, all of volcanic origin, and is located at the triple junction of the Eurasian, African and American tectonic plates. The nine islands are divided into three groups: the western group (Corvo and Flores isls.), the central group (Faial, Pico, Graciosa, São Jorge and Terceira isls.), and the eastern group (São Miguel and Santa Maria isls) (Fig.
Main characteristics of the Azorean islands (bold) and native forest fragments sampled from 1999 to 2011, including area (hectares), highest point (altitude in metres), distance to the nearest island/fragment (isolation in kilometres) and the oldest geological age of emerged substrate (million years BP) (adapted from
Island |
Fragment |
Code |
Area (ha) |
Altitude (m) |
Isolation (km) |
Age (my) |
Flores |
FLO |
14102 |
911 |
236.43 |
2.16 |
|
Morro Alto e Pico da Sé |
MO |
1331 |
911 |
6.02 |
2.16 |
|
Caldeiras Funda e Rasa |
FR |
240 |
773 |
6.02 |
2.16 |
|
Faial |
FAI |
17306 |
1043 |
34.26 |
0.73 |
|
Caldeira do Faial |
CA |
190 |
934 |
4.67 |
0.73 |
|
Cabeço do Fogo |
CG |
36 |
597 |
4.67 |
0.60 |
|
Pico |
PIC |
44498 |
2350 |
32.42 |
0.30 |
|
Mistério da Prainha |
MP |
689 |
881 |
2.92 |
0.26 |
|
Caveiro |
CA |
184 |
1077 |
4.61 |
0.27 |
|
Lagoa do Caiado |
LC |
79 |
945 |
2.92 |
0.28 |
|
São Jorge |
SJG |
24365 |
1053 |
32.42 |
0.55 |
|
Topo |
TO |
220 |
946 |
15.13 |
0.55 |
|
Pico Pinheiro |
PP |
73 |
717 |
15.13 |
0.55 |
|
Terceira |
TER |
40030 |
1021 |
71.67 |
3.52 |
|
S. Bárbara e M. Negros |
SB |
1347 |
1021 |
7.20 |
1.24 |
|
Biscoito da Ferraria |
BF |
557 |
809 |
3.03 |
0.10 |
|
Guilherme Moniz |
GM |
223 |
487 |
2.70 |
0.41 |
|
Terra Brava |
TB |
180 |
726 |
2.70 |
0.10 |
|
Pico do Galhardo |
PG |
38 |
655 |
2.79 |
0.10 |
|
São Miguel |
SMG |
74456 |
1105 |
97.53 |
4.01 |
|
Pico da Vara |
PV |
306 |
1105 |
3.42 |
3.20 |
|
Graminhais |
GR |
15 |
930 |
4.02 |
3.20 |
|
Atalhada |
AT |
10 |
500 |
3.42 |
4.01 |
Sampling protocol
Eighteen native forest fragments distributed across seven of the nine islands were sampled (Table
Canopy sampling was conducted during the trapping period, when the vegetation was dry. A 5 m wide square was established every 15 m (total of 10 squares per transect). Two woody plant specimens of the most abundant species (up to three species when available) were sampled in each square. For each selected plant, a branch was chosen at random and a beating tray placed beneath. The tray consisted of a 1 m wide and 60 cm deep cloth inverted pyramid, with a plastic bag at the vertex. Five beatings were made using a stick for each plant individual sampled.
The arthropod taxa considered in this study were selected based on the availability of expert taxonomists and ability to readily separate them by morphological criteria. All Araneae, Opiliones, Pseudoscorpionida, Diplopoda, Chilopoda and Insecta (excluding Collembola, Diptera and Hymenoptera) were assigned to morphospecies through comparison with a reference collection. Various taxonomists (PAVB, ARMS, LC, PC, HE, FI, VM, MTP, JR, AB, ABS, RzS, VV, JW, JAQ, and see also Acknowledgments) checked the assignment to morphospecies, performed species identifications and supplied additional ecological information. The taxonomic nomenclature follows the most recent list of Azorean arthropods (
All specimens are deposited in the Entomological Collection Dalberto Teixeira Pombo at the University of the Azores (Portugal), under the curation of Paulo A. V. Borges (pborges@uac.pt).
In this contribution we list the 286 species for which we obtained an identification. The new records for each island are marked with *. For this list two families of Coleoptera were not considered since they will be presented elsewhere, Staphylinidae (Borges et al. in prep.) and Zopheridae (
All specimens were assigned a SITE CODE composed of several letters and numbers that read as follows (see Suppl. material
i) the first three letters refer to island name (FLO – Flores; FAI – Faial; PIC – Pico; SJG – São Jorge; GRA – Graciosa; TER – Terceira; SMG – São Miguel; SMR – Santa Maria);
ii) the following two letters refer to fragment name (Flores: FR - Caldeiras Funda e Rasa, MA - Morro Alto e Pico da Sé; Faial: CF – Caldeira do Faial, CG – Cabeço do Fogo; Pico: CA – Caveiro, LC – Lagoa do Caiado, MP – Mistério da Prainha; São Jorge: PP – Pico Pinheiro, TO – Topo; Terceira: BF – Biscoito da Ferraria, GM – Caldeira do Guilherme Moniz, PG – Pico do Galhardo, SB –Serra de Santa Bárbara, TB – Terra Brava; São Miguel: AT – Atalhada, GR – Graminhais, PV – Pico da Vara; Santa Maria: PA – Pico Alto);
iii) the following three characters refer to the sampling transect; and
iv) the next letter refers to the sampling technique: P - pitfall, B - canopy beating; for pitfall samples (P) TU – Turquin and ET – ethylene glycol; for canopy samples (B) the next two letters refer to the plant sampled: CA = Calluna vulgaris, CL = Clethra arborea, ER = Erica azorica, FR = Frangula azorica, IL = Ilex perado azorica, JU = Juniperus brevifolia, LA = Laurus azorica, MC = Morella faya, MS = Myrsine africana, PI = Picconia azorica, PT = Pittosporum undulatum, VA = Vaccinium cylindraceum.
For the geographical location of transects within reserves (UTM coordinates) see Suppl. material
Accumulation curves were obtained using the software “Species Diversity and Richness” V.4.
COR; FLO
Azorean Arthropod biodiversity - towards a more complete knowledge
The ultimate goal of biodiversity assessments is documenting all species inhabiting a region. However, this has often proven impossible to achieve given the unfeasibility of collecting every single species that exists in a study area. This study focuses on the terrestrial arthropod diversity of the Azores and encompasses most orders of the phylum Arthropoda. A pool of a total of 1215 species and subspecies was surveyed, representing 53% of the whole arthropod fauna known from the Azores (
Species richness for the Azores archipelago and each island. Total currently known species, the number of species surveyed during this study and those that represent new records are presented.
Known species in the Azores |
Pool of surveyed taxa |
New records |
New records (%) |
|
AZORES |
2316 |
1215 |
26 |
2.13 |
FLO |
797 |
461 |
55 |
11.93 |
FAI |
945 |
537 |
51 |
9.49 |
PIC |
808 |
463 |
46 |
9.93 |
SJG |
620 |
359 |
76 |
21.17 |
TER |
1224 |
731 |
52 |
7.11 |
SMG |
1592 |
861 |
28 |
3.25 |
SMR |
799 |
573 |
38 |
6.63 |
Total species and subspecies records for the Azores, new species and subspecies records during this study and increment for the most speciose classes and orders. Values for all islands are added, so richness may be up to 7 times higher than the archipelago's richness (as 7 islands were surveyed). (*)The Coleoptera families Staphylinidae and Zopheridae were not considered (see text).
Total records |
New records |
New Records (%) |
|
Class Arachnida |
362 |
124 |
34.25 |
Order Pseudoscorpiones |
19 |
5 |
26.32 |
Order Opiliones |
12 |
11 |
91.67 |
Order Araneae |
331 |
108 |
32.63 |
Class Diplopoda |
67 |
24 |
35.82 |
Order Polydesmida |
18 |
8 |
44.44 |
Order Polyxenida |
0 |
0 |
0.00 |
Order Julida |
44 |
12 |
27.77 |
Order Chordeumatida |
5 |
4 |
80.00 |
Class Chilopoda |
21 |
9 |
42.86 |
Order Scutigeromorpha |
0 |
0 |
0.00 |
Order Lithobiomorpha |
7 |
0 |
0.00 |
Order Scolopendromorpha |
4 |
2 |
50.00 |
Order Geophilomorpha |
10 |
7 |
70.00 |
Class Insecta |
1012 |
189 |
18,68 |
Order Microcoryphia |
13 |
4 |
30.77 |
Order Zygentoma |
0 |
0 |
0.00 |
Order Ephemeroptera |
6 |
0 |
0.00 |
Order Odonata |
0 |
0 |
0.00 |
Order Blattaria |
7 |
3 |
42.86 |
Order Orthoptera |
10 |
0 |
0.00 |
Order Phasmatodea |
0 |
0 |
0.00 |
Order Dermaptera |
14 |
0 |
0.00 |
Order Psocoptera |
75 |
40 |
53.33 |
Order Thysanoptera |
76 |
6 |
7.89 |
Order Hemiptera |
290 |
82 |
28.28 |
Order Neuroptera |
7 |
3 |
42.86 |
Order Coleoptera (*) |
361 |
36 |
9.97 |
Order Trichoptera |
6 |
4 |
66.67 |
Order Lepidoptera |
147 |
11 |
7.84 |
The number of species identified for each of the 18 native forest fragments surveyed is shown in Fig.
BALA2 samples only added 4% of species to the previous BALA survey (Fig.
Species accumulation curve for the 286 species of arthropods collected in 152 pitfall and beating samples between 1999 and 2011. The solid line corresponds to the chronological sample sequence and the dotted line is a randomized curve (1000 runs). Samples to the left of the vertical line were collected in BALA1 and to the right in BALA2.
A total of 163744 individuals were identified as belonging to the 286 species (see Suppl. material
The increase in the number of islands from where each species is known and the distribution increase for many species within each island shows the importance of regional standardized surveys, which provided a major improvement in the knowledge of the distribution of arthropod species in the native forests of the Azores.
The fact that most diversity was captured during the first two years of the project reflects the importance of sampling a wide geographic range covering all the islands and the maximum number of sites. Increasing the number of samples per fragment (sampling performed in 2004) or replicating the sampling at a different time (29 sites in 2010 to 2011; BALA2 project) had a lesser impact in increasing our knowledge about biodiversity (Fig.
The future agenda for surveying and monitoring Azorean arthropod biodiversity includes:
a) expanding the standardized survey of Azorean arthropods to other habitat types, mostly man-modified, an already on-going task for some of the islands (see e.g.
b) selecting study areas along a comprehensive environmental gradient where an optimal sampling strategy will be applied in order to sample the entire arthropod communities (All Taxa Biodiversity Inventory - ATBI). ATBIs are intensive sampling efforts to identify and record all living species that exist within a given area and simultaneously create a common and standardized biodiversity database (
c) finishing the identification of many morphospecies. Good progress has been made with Staphylinidade (Borges et al. in prep.), but other taxa need further effort to reach proper identification;
d) increase sampling and update the current list of Azorean Hymenoptera and Diptera, which is clearly incomplete (
e) contributing to the validation and updating of the pan-European checklists programs, including Fauna Europaea (
This study advances the knowledge on the unique arthropod biodiversity of the Azores, but at the same time highlights the need for further surveys. We strongly believe that the BALA project will stimulate further research and conservation actions towards the preservation of Azorean biodiversity. Furthermore, we hope that all the taxa yet to be identified will entice taxonomist to join us in the endeavour of cataloguing all terrestrial arthropods of the most remote of the Macaronesian archipelagos, the Azores. The ongoing longterm research projects in Azores and the recent creation of the E-Repository ISLANDLAB will create new opportunities for biodiversity studies in Azores.
We are grateful to all researchers that collaborated in the field and lab work: Álvaro Vitorino, Anabela Arraiol, Annabella Borges, Ana Rodrigues, Francisco Dinis, Emanuel Barcelos, Hugo Mas, João Amaral, João Moniz, Lara Dinis, Paula Gonçalves, Sandra Jarroca and Luís Vieira. The Forest Services provided logistic support on each island. Acknowledgments are due to the taxonomists Fernando Ilharco, Ole Karsholt, Wolfgang Rücker and Richard zur Strassen who assisted with species identification. This work was funded by the Azorean Direcção Regional dos Recursos Florestais (Azorean Government / project 17.01-080203) and FCT- project PTDC/BIA-BEC/100182/2008 – “Predicting extinctions on islands: a multi-scale assessment”. Additional data was obtained from project ATLANTISMAR (DRCT–M2.1.2 /I/027/2011). CR was supported by FCT grant SFRH/BPD/91357/2012. AMCS was supported by a Marie Curie Intra-European Fellowship (IEF 331623 ‘COMMSTRUCT’) and by a Juan de la Cierva Fellowship (IJCI-2014-19502) funded by the Spanish ‘Ministerio de Economía y Competitividad’. Open Access was funded by national funds through FCT - Fundação para a Ciência e a Tecnologia in the frame of the project UID/BIA/00329/2013 (2015-2017).
PAVB, ARMS, JQ and KAT conceived the ideas; PAVB, CG, LC, FR, PC, FP, CR, IRA, CM, CA, GA, SPR, JH, AMCS, ABS, JW, JAQ, ARMS and KAT obtained samples; PAVB, CG, EM and LB prepared the databases; PAVB, LC, PC, HE, FI, VM, MTP, JR, AB, ABS, RzS, VV, JW, JAQ and ARMS performed taxonomic work and identified species; PAVB led the writing with substantial input from the other authors.
Detailed data on the occurrences and abundances of the studied species. Data on species abundance in each individual sample (pitfall trap or canopy beating) for the 152 transects in eighteen protected areas and seven Azorean islands.
METADATA from Appendix 1 – Detailed data on the distribution and abundance of the studied species
UTM coordinates (regions 25S for Flores and 26S for all other islands), altitude (meters) and supporting project of the studied transects in the Azores. Transect code according to island, reserve and transect number (see text)
The complete list of new records per island.
Detailed abundance for each species in each of the 18 protected areas