Ichthyofauna of the Selvagens Islands. Do small coastal areas show high species richness in the northeastern Atlantic?

Abstract

The Selvagens Islands are located in the northeastern Atlantic between the Canary Islands and Madeira Island. As a result of their small dimensions, remote location and harsh sea conditions only a few studies have been conducted to describe their marine species diversity. We were able to identify 29 new coastal fish species, an increase of 33% in the ichthyofauna described for these islands (n = 88). There is a prevalence of species with tropical affinities and only 2.3% (n = 2) are endemic to Macaronesia. Considered a stepping-stone colonization model from the nearest continental shore, as proposed by other authors for this region, the Selvagens Islands represent 34.1% of the ichthyofauna described for the much larger Canary Islands (nspecies = 258, submerged area nSelvagensIs. = 2.3%) and 47.3% of the ichthyofauna described for the more distantly located Madeira Island (nspecies = 186, submerged area nSelvagensIs. = 17.9%). Interestingly 6.8% (n = 6) of the species failed to bridge the gap between the Selvagens Islands and Madeira Island. Data collected so far showed no trend toward an increasing number of species with high dispersal capability. The Selvagens Islands are an example of a high coastal species diversity occurring even in very small areas of the northeastern Atlantic Ocean.

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Ichthyofauna of the Selvagens Islands. Do small
coastal areas show high species richness in the
northeastern Atlantic?
Frederico Almadaa, David Abecasisb, David Villegas-Ríosc, Sofia Henriquesd, Miguel P.
Paisd, Marisa Batistad, Bárbara Horta e Costaab, Joana Martinsa, Inês Tojeirae, Nuno
Vasco Rodriguesf, Ricardo Araújog, Miguel Soutoe, Hany Alonsoa, Jesús M. Falcónh, Filipe
Henriquesg, Paulo Catrya, Henrique Cabrald, Manuel Biscoitog & Vitor C. Almadaa
a Unidade de Investigação em Eco-Etologia, ISPA – Instituto Universitário, Lisboa,
Portugal
b Centro de Ciências do Mar (CCMAR, CIMAR – Associate Laboratory), Universidade do
Algarve, Faro, Portugal
c Instituto de Investigaciones Marinas (CSIC), Vigo, Spain
d Faculdade de Ciências, Centro de Oceanografia, Universidade de Lisboa, Lisboa,
Portugal
e Task Group for the Extension of the Continental Shelf, EMEPC, Paço de Arcos, Portugal
f GIRM – Marine Resources Research Group, Polytecnic Institute of Leiria, Peniche,
Portugal
g Museu de História Natural do Funchal, Funchal, Portugal
h Departamento de Biología Animal (Ciencias Marinas), Facultad de Biología, Grupo de
Investigación BIOECOMAC, Universidad de La Laguna, La Laguna, Spain
Published online: 16 Jun 2014.
To cite this article: Frederico Almada, David Abecasis, David Villegas-Ríos, Sofia Henriques, Miguel P. Pais, Marisa Batista,
Bárbara Horta e Costa, Joana Martins, Inês Tojeira, Nuno Vasco Rodrigues, Ricardo Araújo, Miguel Souto, Hany Alonso,
Jesús M. Falcón, Filipe Henriques, Paulo Catry, Henrique Cabral, Manuel Biscoito & Vitor C. Almada (2014): Ichthyofauna
of the Selvagens Islands. Do small coastal areas show high species richness in the northeastern Atlantic?, Marine Biology
Research, DOI: 10.1080/17451000.2014.889306
To link to this article: http://dx.doi.org/10.1080/17451000.2014.889306
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ORIGINAL ARTICLE
Ichthyofauna of the Selvagens Islands. Do small coastal areas show high
species richness in the northeastern Atlantic?
FREDERICO ALMADA
1
*, DAVID ABECASIS
2
, DAVID VILLEGAS-RÍOS
3
,
SOFIA HENRIQUES
4
, MIGUEL P. PAIS
4
, MARISA BATISTA
4
,
BÁRBARA HORTA e COSTA
1,2
, JOANA MARTINS
1
, INÊS TOJEIRA
5
,
NUNO VASCO RODRIGUES
6
, RICARDO ARAÚJO
7
, MIGUEL SOUTO
5
, HANY ALONSO
1
,
JESÚS M. FALCÓN
8
, FILIPE HENRIQUES
7
, PAULO CATRY
1
, HENRIQUE CABRAL
4
,
MANUEL BISCOITO
7
& VITOR C. ALMADA
1+
1
Unidade de Investigação em Eco-Etologia, ISPA –Instituto Universitário, Lisboa, Portugal,
2
Centro de Ciências do Mar
(CCMAR, CIMAR –Associate Laboratory), Universidade do Algarve, Faro, Portugal,
3
Instituto de Investigaciones Marinas
(CSIC), Vigo, Spain,
4
Faculdade de Ciências, Centro de Oceanografia, Universidade de Lisboa, Lisboa, Portugal,
5
Task
Group for the Extension of the Continental Shelf, EMEPC, Paço de Arcos, Portugal,
6
GIRM –Marine Resources Research
Group, Polytecnic Institute of Leiria, Peniche, Portugal,
7
Museu de História Natural do Funchal, Funchal, Portugal, and
8
Departamento de Biología Animal (Ciencias Marinas), Facultad de Biología, Grupo de Investigación BIOECOMAC,
Universidad de La Laguna, La Laguna, Spain
Abstract
The Selvagens Islands are located in the northeastern Atlantic between the Canary Islands and Madeira Island. As a result
of their small size, remote location and harsh sea conditions only a few studies have been conducted to describe their marine
species diversity. We were able to identify 29 new coastal fish species, an increase of 33% in the ichthyofauna described for
these islands (n= 88). There is a prevalence of species with tropical affinities and only 2.3% (n= 2) are endemic to
Macaronesia. Considered a stepping-stone colonization vector from the nearest continental shore, as proposed by other
authors for this region, the Selvagens Islands host 34.1% of the ichthyofauna described for the much larger Canary Islands
(n
species
= 258, submerged area n
SelvagensIs.
= 2.3%) and 47.3% of the ichthyofauna described for the more distantly located
Madeira Island (n
species
= 186, submerged area n
SelvagensIs.
= 17.9%). Interestingly, 6.8% (n= 6) of the species failed to
bridge the gap between the Selvagens Islands and Madeira Island. Data collected so far showed no trend toward an
increasing number of species with high dispersal capability. The Selvagens Islands are an example of a high coastal species
diversity occurring even in very small areas of the northeastern Atlantic Ocean.
Key words: Biogeography, dispersal capability, ichthyofauna, Macaronesia, Selvagens Islands
Introduction
The warm temperate northeastern Atlantic Ocean is
punctuated by a number of archipelagos and sea-
mounts. This region known as Macaronesia includes
the Canary Islands, located near the north African
coast and, in a northwestward offshore direction, the
archipelagos of Madeira, Selvagens and the Azores
(Lloris et al. 1991; Brito et al. 2007). According to
several authors the ichthyogeography of Macaronesia
has mainly been influenced by the tropical west
African coast, the Mediterranean Sea and to a lesser
extent the western European continental shores
(Almada et al. 2001; Briggs & Bowen 2012).
Recently, Almada et al. (2013) have shown the
*Correspondence: Frederico Almada, Eco-Ethology Investigation Unit, ISPA –Instituto Universitário de Ciências Psicológicas, Sociais e da
Vida, Rua Jardim do Tabaco 34, 1149-041, Lisboa, Portugal. E-mail: falmada@ispa.pt
+
deceased
Published in collaboration with the Institute of Marine Research, Norway
Marine Biology Research, 2014
http://dx.doi.org/10.1080/17451000.2014.889306
(Accepted 24 January 2014; Published online xx xxx xxxx)
© 2014 Taylor & Francis
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distinctiveness of the Macaronesia ichthyofauna
compared with the continental African and Euro-
pean regions at the same latitude and its close
affinity with west African tropical communities.
These affinities were already noted by Santos et al.
(1995), who suggested that the colonization of
Macaronesia by warm-water fish followed a route
from west African continental shores along progres-
sively distant clusters of islands in a stepping-stone
process. But how is it possible to support this
hypothesis if the Canary Current, the main current
in this region, flows in the opposite direction south-
ward along the southwestern European and north-
western African coasts? Stramma (1984) showed
that the transport of eggs and larvae to the north
was possible through gyres and eddies that fre-
quently flow northward, forming a complex two-
way system of connection among these archipelagos.
Additionally this northward colonization trend was
also detected by molecular studies that analysed the
phylogeography of Macaronesian warm-water spe-
cies (e.g. Domingues et al. 2006).
The Selvagens Islands (Lat. 30°08′45″N, Long.
15°51′51″W; Lat. 30°01′59″N, Long. 16°01′00″W)
are located south of Madeira Island and north of the
Canary Islands (Figure 1). They form an isolated
group of small islands and reefs that may have played
(or may play) an important role for some fish
species, allowing them to bridge the gap between
those larger archipelagos.
Although the Selvagens Islands have been a
marine protected area for more than 40 years, their
remoteness and the rough oceanographic conditions
usually found in this region of the Atlantic Ocean are
probably the reasons why only one paper has
addressed their ichthyofaunal richness. Falcón et al.
(2000) described the ichthyofauna around these
islands, reporting a total of 60 coastal fish species.
More surveys are still needed to provide a better
description of the species composition at the Selvagens
Islands. This is of major importance to understand-
ing the forcing agents underlying the colonization
patterns presented above.
The ichthyogeography of the northeastern Atlantic
may have been shaped significantly by climatic
events. Macaronesia is included in the Lusitania
Province which encompasses the warm and temper-
ate waters of the northeastern Atlantic, that extend
from the southern British Isles in the north to
southern Morocco in the south and eastwards
through the Mediterranean (Briggs 1995; Spalding
et al. 2007; Briggs & Bowen 2012). This province is
Figure 1. Map showing the location of the Selvagens Islands in the northeastern Atlantic between Madeira and the Canary Islands.
Bathymetric data of these small remote islands are also presented.
2F. Almada et al.
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one of the poorest regions on the planet in terms of
fish species diversity, a fact probably related to the
climatic deterioration that has been evolving since
the Miocene (Briggs 1995) and that became espe-
cially severe with the Pleistocene glaciations (CLI-
MAP 1981; Hayes et al. 2005). During the glacial
maxima much of the habitat for warm-water species
ceased to be available on the western European
shores. In the last glacial peak the polar front
advanced to the south, possibly reaching the west
Iberian coast (Dias et al. 1997).
While the continental shores of west Africa and
Europe experienced a severe cooling (Briggs 1995),
Macaronesia maintained milder conditions together
with some warmer water pockets that persisted in the
Mediterranean (Thiede 1978; Briggs 1995; Almada
et al. 2001). Indeed, the Macaronesian archipelagos
were affected differently by the glaciations. While the
Azores, in the north, experienced a moderate drop in
sea surface temperature (2–3°) in the last glacial
maximum (LGM), Madeira seems to have been
almost unaffected while in the Canaries, the islands
nearer the African shore were more affected than
those located offshore (Crowley 1981).
During the LGM the average sea level was approxi-
mately 130 m lower than at the present time (Dias
et al. 1997). These eustatic changes led to an increase
in the coastal area available for most inshore fish both
in the present northeastern Atlantic Islands and on
several seamounts in this region that are currently
submerged (e.g. Gorringe Bank). This fact may have
favoured stepping-stone processes along the Macar-
onesian islands by inshore fish fauna during the
regression of the polar front in the last 11,000 years.
This hypothesis is also supported by the low number
of endemic species in the Macaronesian archipelago
(Briggs 1966; Santos et al. 1997).
In this paper we provide an updated checklist and
abundance data of fish species from the Selvagens
Islands. We discuss the biogeographic affinities of
the ichthyofauna of the Selvagens Islands reported so
far and also analyse the role of some ecologically
relevant features traditionally related to the dispersal
ability of fish species in an attempt to further
understand the factors shaping the ichthyofaunal
richness of these islands.
Materials and methods
The Selvagens Islands are small, uninhabited islands
of volcanic origin dating from the Oligocene (24.2 to
29.5 Ma) (Mata et al. 2010). The largest islands,
Selvagem Grande (2.5 km
2
) and Selvagem Pequena
(0.16 km
2
), are approximately 18 km from each other
and are surrounded by small islets. The largest island,
Selvagem Grande, is mostly flat and surrounded by
abrupt cliffs, about 80 m high, with scarce intertidal
platforms. The smaller island, Selvagem Pequena,
rises just a few metres above sea level, with large
areas of intertidal rocky platforms that lead into a
shallow subtidal area connecting small rocks and
islets. The shallow seascape is mostly rocky. The
calculation of available submerged areas between 0
and 60 m depth and distances between groups of
islands was obtained using the ArcGIS Desktop 9.3
(ESRI) software and the method described by Vin-
centy (1975). The coordinate system used in this
work was the WGS84 Gebco (08-Grid, version
20100927) bathymetry data with 30 arc-second
(IOC et al. 2003) and CAOP (version 2011) coastline
data.
The EMEPC/M@rbis/Selvagens expedition in
June–July 2010 involved more than 80 investigators
and had the larger objective of sampling the marine
communities of this poorly known archipelago
including algae, invertebrates, fish and marine birds.
No additional sampling was made in the remaining
seasons due to the remoteness of these islands and the
rough sea conditions which can almost completely
submerge some islands of this archipelago and make
sampling in these coastal areas a difficult or danger-
ous task. Ichthyofauna censuses included non-
destructive sampling methods like underwater scuba
diving visual census, photography and video surveys,
snorkelling surveys, intertidal censuses during low
tide and additional sampling methods described
below. Photos and videos of the species observed
during the campaign are available from EMEPC
(2010). Specimens of some species were euthanized
with MS-222 and preserved in ethanol (marked with
an (S) in Table I) for later molecular studies. Species
names follow Eschmeyer (2014) and specific literat-
ure on the taxonomy of some fish families (Blennii-
dae: Almada et al. (2005a,2005b), Muss et al.
(2001); Diodontidae: Leis (2006); Gadidae: Roa-
Varón & Ortí (2009); Gobiesocidae: Almada et al.
(2008), Henriques et al. (2002); Labridae: Almada
et al. (2002), Hanel et al. (2002)) was also considered.
Intertidal specimens were captured with hand nets
and released at the same site. Large pools were also
visually sampled by snorkelling. Subtidal sampling
involved groups of two to four divers who performed
six snorkelling and 39 scuba diving censuses (includ-
ing five nocturnal dives) around Selvagem Grande,
and 18 snorkelling and 35 scuba diving censuses
(including three nocturnal dives) around Selvagem
Pequena. A strip transect technique was adopted
(Brock 1954) and transects were established along
isobaths and allocated to pre-established depth strata
between 0 and 30 m. A fixed transect area of 50 × 4 m
was sampled following the recommendations of
Fowler (1987). Under-estimation of cryptobenthic
Ichthyofauna of the Selvagens Islands 3
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Table I. Species list for the Selvagens Islands. N, new records for Selvagens Islands (species highlighted in bold); S, specimens collected and preserved. Abundance levels: 1, single observation (one
individual); 2, rare (2–10); 3, common (11–100); 4, abundant (>100); (a), species sampled by coastal angling, (b), species identified in fresh stomach contents of Calonectris diomedea; C, species
reported for the Canary Islands but absent from Madeira Island; asterisks mark species previously reported by Falcón et al. (2002), whose valid name has changed due to recent taxonomic updates.
Biogeographic affinities: TAS, tropical west African species; WTS, warm-temperate species; TES, temperate species; EAS, eurythermic eastern Atlantic species; AAS, amphiatlantic species; WAS,
west Atlantic species; COS, Cosmopolitan species and END, endemic Macaronesian species; MED, distribution range includes the Mediterranean. Habitats: B, benthic, D, demersal, BP,
benthopelagic, P, pelagic. Adult mobility: s, sedentary, m, mobile, vm, very mobile. Reproductive functional group: v, viviparous; w, ovoviviparous; oviparous with pelagic eggs (op) and with
adhesive eggs attached to substrata (ov); og, species in which one or the other parent guards externally; os, species that shed their eggs and protect them in a part of their body; ob, species that
produce eggs which settle on the substratum (see Materials and methods section for more information).
Species Common name
Abundance level
(no. of
independent
observations)
Falcón
et al.
(2000)
EMEPC/
M@rbis/
Selvagens
(2010)
Main
biogeographic
affinities
Water
column
habitat
Adult
mobility
Reproductive
functional
group
Recent taxonomic
updates and
references
Subclass Elasmobranchii
CARCHARHINIDAE
Prionace glauca (Linnaeus, 1758) Blue shark 1 (1) N COS P vm v
DASYATIDAE
Dasyatis pastinaca (Linnaeus, 1758) Common stingray 2 (3) √√EAS/MED B m w
Taeniura grabata (Geoffroy Saint-Hilaire, 1817) Round stingray 1 (1) N COS B m w
MYLIOBATIDAE
Myliobatis aquila (Linnaeus, 1758) Common
eagle ray
2 (2) √√EAS/MED B m w
Pteromylaeus bovinus (Geoffroy Saint-Hilaire, 1817) Bull ray 1 (1) N COS B m w
TORPEDINIDAE
Torpedo marmorata Risso, 1810 Spotted torpedo 1 (1) N EAS/MED B s w
Class Actinopterygii
ANTENNARIIDAE
Antennatus nummifer (Cuvier, 1817) Spotfin frogfish 1 (1) N COS B s ov
APOGONIDAE
Apogon imberbis (Linnaeus, 1758) Cardinal fish 3 (43) √√TAS/MED D s os
ATHERINIDAE
Atherina presbyter Cuvier, 1829 Sand smelt 4 (5) √√ TES P vm op
AULOSTOMIDAE
Aulostomus strigosus Wheeler, 1955 Atlantic cornetfish 3 (17) N TAS D m op
BALISTIDAE
Balistes capriscus Gmelin, 1789 (S) Grey triggerfish 3 (23) √√AAS/MED BP vm og
Canthidermis sufflamen (Mitchill, 1815) Ocean triggerfish 3 (19) √√ WAS BP vm og
BELONIDAE
Belone belone (Linnaeus, 1761) Garfish 1 (1) N TES/MED P vm ov
BLENNIIDAE
Coryphoblennius galerita (Linnaeus, 1758) (S) Montagu’s blenny 3 (14) √√TES/MED B s og
Lipophrys pholis (Linnaeus, 1758) (S) Shanny 2 (2) √√TES/MED B s og Almada et al. (2005a)
Lipophrys trigloides (Valenciennes, 1836)* (S) N/A 1 (1) √√WTS/MED B s og Almada et al. (2005a)
Ophioblennius atlanticus (Valenciennes, 1836) Redlip blenny 3 (69) √√ AAS B s og Muss et al. (2001)
Parablennius incognitus (Bath, 1968) N/A 0 √WTS/MED B s og
Parablennius parvicornis (Valenciennes, 1836) (S) Rock-pool blenny 4 (57) √√ TAS B s og Almada et al. (2005b)
Scartella cristata (Linnaeus, 1758) (C) Molly miller 2 (2) √√ COS B s og
BOTHIDAE
Bothus podas (Delaroche, 1809) Wide-eyed
flounder
1 (1) √√TAS/MED B s op
4F. Almada et al.
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Table I. (Continued)
Species Common name
Abundance level
(no. of
independent
observations)
Falcón
et al.
(2000)
EMEPC/
M@rbis/
Selvagens
(2010)
Main
biogeographic
affinities
Water
column
habitat
Adult
mobility
Reproductive
functional
group
Recent taxonomic
updates and
references
CARANGIDAE
Caranx crysos (Mitchill, 1815) Blue runner 3 (19) √√AAS/MED BP vm op
Naucrates ductor (Linnaeus, 1758) Pilot fish (b) N COS P vm op
Pseudocaranx dentex (Bloch & Schneider, 1801) White trevally 4 (24) √√ COS BP vm op
Seriola dumerili (Risso, 1810) Greater amberjack 3 (14) √√ COS BP vm op
Seriola fasciata (Bloch, 1793) Lesser amberjack 2 (2) √√AAS/MED BP vm op
Seriola rivoliana Valenciennes, 1833 Longfin yellowtail 4 (43) √√ COS BP vm op
Trachinotus ovatus (Linnaeus, 1758) Pompano, Derbio 3 (13) √√EAS/MED P vm op
CLUPEIDAE
Sardinella maderensis (Lowe, 1838) Madeira
sardinella
4 (1) N TAS/MED P vm op
CONGRIDAE
Conger conger (Linnaeus, 1758) Conger eel 2 (5) N EAS/MED B s op
DIODONTIDAE
Chilomycterus reticulatus (Linnaeus, 1758)* Spotfin burrfish 2 (6) √√ COS D m op Leis (2006)
Diodon eydouxii Brisout de Barneville, 1846 Pelagic
porcupinefish
1 (1) N COS BP vm op Leis (2006)
EXOCOETIDAE
Cheilopogon exsiliens (Linnaeus, 1771) (C) Bandwing
flyingfish
(a) N WAS P vm op
Cheilopogon pinnatibarbatus (Bennett, 1831) Bennett’s
flyingfish
(a) N COS P vm op
Exocoetus volitans Linnaeus, 1758 Tropical two-wing
flyingfish
(a) N COS P vm op
GADIDAE
Gaidropsarus guttatus (Collett, 1890) Macaronesian
rockling
1 (1) N WTS B s op Roa-Varón &
Ortí (2009)
GOBIESOCIDAE
Apletodon dentatus (Facciolà, 1887) (C) Sea-urchin
clingfish
1 (1) N TES/MED B s og
Apletodon pellegrini (Chabanaud, 1925) Chubby clingfish 2 (2) N TAS B s og
Lepadogaster candolii Risso, 1810* Connemarra
clingfish
2 (6) √√TES/MED B s og But see Almada
et al. (2008)
Lepadogaster lepadogaster (Bonnaterre, 1788)* (S) Shore clingfish 3 (11) √√WTS/MED B s og Henriques
et al. (2002)
GOBIIDAE
Gnatholepis thompsoni Jordan, 1904 Goldspot goby 2 (2) N AAS B s og
Gobius niger Linnaeus, 1758 (C) Black goby 2 (2) N EAS/MED B s og
Gobius paganellus Linnaeus, 1758 Rock goby 2 (8) √√EAS/MED B s og
Gobius xanthocephalus Heymer & Zander, 1992 (C) Yellow-
headed goby
1 (1) N WTS/MED B s og
Ichthyofauna of the Selvagens Islands 5
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Table I. (Continued)
Species Common name
Abundance level
(no. of
independent
observations)
Falcón
et al.
(2000)
EMEPC/
M@rbis/
Selvagens
(2010)
Main
biogeographic
affinities
Water
column
habitat
Adult
mobility
Reproductive
functional
group
Recent taxonomic
updates and
references
Mauligobius maderensis (Valenciennes, 1837) (S) Madeira
rock goby
4 (72) √√ END B s og
Thorogobius ephippiatus (Lowe, 1839) Leopard-
spotted goby
0√TES/MED B s og
HAEMULIDAE
Parapristipoma octolineatum (Valenciennes, 1833) African striped
grunt
2 (2) √√TAS/MED D m op
KYPHOSIDAE
Kyphosus sectatrix (Linnaeus, 1758)* Bermuda
sea chub
4 (21) √√AAS/MED BP vm op Eschmeyer (2014)
LABRIDAE
Bodianus scrofa (Valenciennes, 1839) Barred hogfish 3 (61) √√ TAS D m op
Coris julis (Linnaeus, 1758) Mediterranean
rainbow wrasse
3 (16) √√TES/MED D m op
Symphodus trutta (Lowe, 1834)* Atlantic wrasse 3 (16) √√ END D m og Almada et al. (2002),
Hanel et al. (2002)
Thalassoma pavo (Linnaeus, 1758) Ornate wrasse 4 (534) √√TAS/MED D m op
Xyrichtys novacula (Linnaeus, 1758) Pearly razorfish 2 (4) √√AAS/MED D m op
LABRISOMIDAE
Labrisomus nuchipinnis (Quoy & Gaimard, 1824) Hairy blenny 2 (9) N AAS B s og
MUGILIDAE
Chelon labrosus (Risso, 1827) Thicklip grey
mullet
4 (26) √√EAS/MED BP m op
Liza aurata (Risso, 1810) (S) Golden grey
mullet
4 (46) N EAS/MED BP m op
Mugil cephalus Linnaeus, 1758 (C) Flathead grey
mullet
0√COS BP m op
MULLIDAE
Mullus surmuletus Linnaeus, 1758 Striped red mullet 2 (2) N TES/MED B m op
MURAENIDAE
Enchelycore anatina (Lowe, 1838) Fangtooth moray 3 (11) √√ WAS B s op
Gymnothorax unicolor (Delaroche, 1809) Brown moray 3 (33) √√TAS/MED B s op
Muraena augusti (Kaup, 1856) (S) Black moray 3 (69) √√ TAS B s op
Muraena helena Linnaeus, 1758 Mediterranean
moray
1 (1) N EAS/MED B s op
PHYCIDAE
Phycis phycis (Linnaeus, 1766) Forkbeard 2 (5) √√EAS/MED D m op
POMACENTRIDAE
Abudefduf luridus (Cuvier, 1830) (S) Canary damsel 4 (343) √√ TAS D m og
Chromis limbata (Valenciennes, 1833) Azores chromis 4 (117) √√ AAS BP m ov
PRIACANTHIDAE
6F. Almada et al.
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Table I. (Continued)
Species Common name
Abundance level
(no. of
independent
observations)
Falcón
et al.
(2000)
EMEPC/
M@rbis/
Selvagens
(2010)
Main
biogeographic
affinities
Water
column
habitat
Adult
mobility
Reproductive
functional
group
Recent taxonomic
updates and
references
Heteropriacanthus cruentatus (Lacepède, 1801) Glasseye 3 (22) √√ COS D m op
SCARIDAE
Sparisoma cretense (Linnaeus, 1758) Mediterranean
parrotfish
4 (243) √√TAS/MED D m op
SCOMBRIDAE
Katsuwonus pelamis (Linnaeus, 1758) Skipjack tuna (a) N COS P vm op
Sarda sarda (Bloch, 1793) Atlantic bonito 2 (4) N AAS/MED P vm op
Scomber colias Gmelin, 1789 Atlantic chub
mackerel
(b) N AAS/MED P vm op
SCORPAENIDAE
Scorpaena maderensis Valenciennes, 1833 (S) Madeira rockfish 4 (254) √√TAS/MED B s ov
Scorpaena notata Rafinesque, 1810 Small red
scorpionfish
2 (2) N TAS/MED B s ov
SERRANIDAE
Anthias anthias (Linnaeus, 1758) Swallowtail
seaperch
2 (1) N TAS/MED D m op
Epinephelus marginatus (Lowe, 1834) Dusky grouper 3 (26) √√ COS D m op
Mycteroperca fusca (Lowe, 1838) Island grouper 3 (14) √√ TAS D m op
Serranus atricauda Günther, 1874 (S) Blacktail comber 4 (135) √√TAS/MED D m op
SPARIDAE
Boops boops (Linnaeus, 1758) Bogue 4 (34) √√EAS/MED P vm op
Diplodus cervinus (Lowe, 1838) Zebra seabream 3 (13) √√TAS/MED D m op
Diplodus sargus (Linnaeus, 1758) White seabream 3 (16) √√TAS/MED D m op
Diplodus vulgaris (Geoffroy Saint-Hilaire, 1817) Common two-
banded seabream
3 (8) √√TAS/MED D m op
Oblada melanura (Linnaeus, 1758) Saddled seabream 4 (22) √√
EAS/MED BP vm op
Sarpa salpa (Linnaeus, 1758) Salema 4 (27) √√EAS/MED D m op
SPHYRAENIDAE
Sphyraena viridensis Cuvier, 1829 Yellowmouth
barracuda
3 (7) √√TAS/MED P vm op
SYNODONTIDAE
Synodus saurus (Linnaeus, 1758) Atlantic lizardfish 3 (11) √√AAS/MED B s op
Synodus synodus (Linnaeus, 1758) Diamond
lizardfish
3 (28) √√ WAS B s op
TETRAODONTIDAE
Canthigaster capistrata (Lowe, 1839) Macaronesian
sharpnose puffer
4 (131) √√TAS/MED D m og
Sphoeroides marmoratus (Lowe, 1838) Guinean puffer 3 (18) √√TAS/MED D m og
TRIPTERYGIIDAE
Tripterygion delaisi Cadenat & Blache, 1970 Black-faced
blenny
3 (47) √√WTS/MED B s og
Ichthyofauna of the Selvagens Islands 7
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reef fish species (see Willis 2001) was minimized
with the implementation of the method proposed
by Beldade & Gonçalves (2007) following each
transect: an interference visual census technique
was applied to an additional 25 × 1 m crypto-
benthic-focused transect. Additionally, all new
species were recorded with no distance-to-observer
limit.
Abundances in the Selvagens Islands were esti-
mated following the scale suggested by Gonçalves
et al. (2002): 1, Single observation (one individual);
2, Rare (2–10 individuals); 3, Common (11–100
individuals); 4, Abundant (>100 individuals). The
number of independent observations of each species
was also recorded in order to discriminate between
isolated observations of a few individuals and large
schools of a given species.
Traditional trawl surveys were not used due to the
rough bottom structure but the sampling of fast-
swimming pelagic species was partially bridged by
obtaining fish from a small number of licensed
fishermen working within the marine reserve. Addi-
tionally, from 2008 to 2011 very detailed studies of
the diet and foraging ecology of Cory’s shearwater
Calonectris diomedea (Scopoli, 1769) have taken place
at the colony on Selvagem Grande. While much of
the prey captured by these birds comes from far
away, some fish species consistently appeared in
regurgitated samples in large numbers and in a very
fresh state. This, combined with information on the
movements of birds obtained by GPS tracking,
allowed us to be confident that these species must
be present in the immediate vicinity of the Selvagens
Islands.
The relative contributions of each biogeographic
region to the Selvagens Islands were defined accord-
ing to species distribution data provided by Froese &
Pauly (2012) and Almada et al. (2013). The biogeo-
graphic units follow the classification provided by
Henriques et al. (2007): tropical west African species
(TAS); warm-temperate species (WTS); cold-tem-
perate species (CTS); temperate species (TES).
Species with a widespread geographic range from
tropical waters to the Bay of Biscay or boreal waters
were classified as eurythermic eastern Atlantic spe-
cies (EAS). Species that occur on both sides of the
Atlantic and are well represented along American
and African/European continental coasts were classi-
fied as amphiatlantic species (AAS). Species reported
in the Selvagens Islands that are absent along
African/European continental coasts but are widely
distributed along the East American coast were
considered of west Atlantic origin (WAS). Species
whose distribution encompasses the Atlantic and
other oceanic basins were classified as cosmopolitan
species (COS). Endemic species (END) were those
whose distribution is restricted to Macaronesia (sensu
Almada et al. 2001; Wirtz 2011), being absent along
the continental coasts of Europe and Africa. When-
ever the distribution range of one species includes
the Mediterranean (MED) that information is also
provided.
Species-specific attributes that may be related to
the ability to disperse, such as the reproductive
functional group and the mobility of adult fish,
followed the categories already defined by other
authors (Elliott et al. 2007; Claudet et al. 2010)
and are referred to in Table I. Larval dispersion and
pelagic larval duration was not included because
there is no information available for many species
presented in this study. Fish were also classified
according to the degree of dependence on the
substrate as: strict benthic species (almost always in
close contact with the substrate, e.g. blennies);
demersal species (highly dependent on the substrate
but are not strictly benthic, e.g. groupers); bentho-
pelagic species (active swimmers that are almost
always in the water column but occasionally use the
substrate to feed, e.g. jacks, or for spawning, e.g.
triggerfish); or pelagic species (long-distance swim-
mers always found in the water column, e.g. tuna).
Results and discussion
The 29 additional species found for the Selvagens
Islands (see Table I) in the current study represent
an increase of 33% compared with what had
previously been reported by Falcón et al. (2000).
Some specimens were only identified to the genus
level because they are fast-swimming pelagic fish that
were observed for a very short period: Sphyrna sp.,
Decapterus sp. and Trachurus sp. Although we report
here their presence in the Selvagens Islands we did
not include them in subsequent analysis.
Differences between fish assemblages in Selvagem
Grande and Selvagem Pequena islands were
assessed calculating the Jaccard index of similarity.
A value of 72% indicates that the fish assemblages
from both islands are very similar. This index was
not higher due to the number of rare species with
one single report from one of the islands (see
Table I). In fact, only 20.5% (n= 18) of the species
were considered abundant, which reflects the relative
contribution to the fish community of rare and
common taxa, with 47.7% (n= 42) and 31.8%
(n= 28), respectively. This similarity is further
emphasized by the fact that no differences were
found between the number of times each species
was reported from each island (Wilcoxon test, n= 88,
Z= 0.99, P= 0.32). The Spearman rank correlation
between abundances at the two islands was 0.73 (n=
88, P< 0.05). Taken together, these results suggest
8F. Almada et al.
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that the variation of abundances of species follows a
similar pattern in the two islands and that none of
them displays higher levels of abundance. In view of
these findings and due to the very close proximity
of those islands, they will be treated as a single entity.
The species reported for the Selvagens Islands are
a subgroup (34.1%) of the coastal ichthyofauna de-
scribed for the much larger Canary Islands (n
species
=
258), meaning that all species reported for the
Selvagens Islands are also present in the Canaries.
It is also remarkable that the species richness of the
Selvagens Islands represents almost half (47.3%) of
the ichthyofauna described for the much larger
Madeira Island (n
species
= 186). This last figure
retains its interest even if we take into consideration
that 6.8% (n= 6) of the Selvagens Islands species
(Apletodon dentatus (Facciolà, 1887); Cheilopogon
exsiliens (Linnaeus, 1771); Gobius niger Linnaeus,
1758; Gobius xanthocephalus Heymer & Zander,
1992; Mugil cephalus Linnaeus, 1758 and Scartella
cristata (Linnaeus, 1758)) are not reported for
Madeira and Porto Santo Islands.
The biogeographic relationships of the Selvagens
Islands with the surrounding regions must be con-
sidered provisional because, although involving a
significant and diversified sampling effort, this cam-
paign was limited to one season as reported above.
Additionally, the increase of 33.0% in the cumulat-
ive number of fish species identified between the
survey of Falcón et al. (2000) and the present one
probably implies that the species richness of these
islands may be greater than the one currently
recorded.
Taking into consideration these limitations there
are some patterns that emerge from the data cur-
rently available. In a decreasing order of importance
the ichthyofauna of the Selvagens Islands is com-
posed of eastern Atlantic tropical species (27.3%,
n= 24), temperate species (10.2%, n= 9) and warm-
temperate species (6.8%, n= 6). The remaining
taxa: widespread eurythermic eastern Atlantic spe-
cies (15.9%, n= 14), cosmopolitan (19.3%, n= 17)
and amphiatlantic species (13.6%, n= 12) provide
no biogeographic information. Two Macaronesian
endemisms were also detected: Mauligobius
maderensis (Valenciennes, 1837) and Symphodus
trutta (Lowe, 1834). It is also interesting to note
that 3.4% (n= 3) of the fish species reported for the
Selvagens Islands are widespread along the western
Atlantic coast, being absent from the European and
African continental coasts. In fact, Canthidermis
sufflamen (Mitchill, 1815), Enchelycore anatina
(Lowe, 1838) and Synodus synodus (Linnaeus,
1758) have their centres of distribution along the
eastern American coast.
In an attempt to find a pattern that could explain
the presence of some species at the Selvagens Islands
we analysed the water column habitat, adult mobility
and reproductive functional group of each species.
The degree of dependence on the substrate revealed
that most species are benthic (42.0%, n= 37) or
demersal (26.1%, n= 23), and only a small propor-
tion are benthopelagic (15.9%, n= 14) or pelagic
(15.9%, n= 14) fish. Furthermore, sedentary
(37.5%, n= 33) and mobile species (35.2%, n=
31) clearly outnumber very mobile species (27.3%,
n= 24). The type of eggs produced by each species
revealed that the majority spawn pelagic eggs (58%,
n= 51), followed by oviparous species with adhesive
eggs with or without parental care of the eggs
(34.1%, n= 30). The remaining reproductive func-
tional groups represent a minority (7.9%, n=7)of
the total number of species.
Compared with other archipelagos the species
richness reported for the Selvagens Islands is
remarkable when one considers that the submerged
area with depths less than 60 m (Table II) is much
smaller than that available in larger Macaronesian
archipelagos: 43 times smaller than the Canaries, 11
times smaller than the Azores and six times smaller
than Madeira (differences in species richness are
addressed below). This conclusion is further sup-
ported by the fact that the species list presented here
probably represents an under-estimation of the
species richness at these islands.
A similar pattern, even more extreme than the
present one, was previously reported by Abecasis
et al. (2009) for the Gorringe Bank with an esti-
mated submerged area above 60 m of approximately
0.8 km
2
(71 times smaller than that of the Selvagens
Table II. Pairwise distances (km) calculated for island groups and Gorringe Bank in the Macaronesian region and estimated area (km
2
)
between 0 and 60 m depth for each location.
Azores Gorringe Bank Madeira, Porto Santo Selvagens Canaries
Azores Is. –1174.2 838.4 1126.9 1112.6
Gorringe Bank –572.0 807.6 805.9
Madeira, Porto Santo Is. –251.3 417.6
Selvagens Is. –153.1
Submerged area 0–60 m (km
2
) 584.8 0.8 308.6 55.2 2392.2
Ichthyofauna of the Selvagens Islands 9
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Islands). The summit of Gorringe is at 30–40 m
depth (Alteriis et al. 2003), which means that this
seamount has no habitat for shallow subtidal fishes.
Despite the very small area and reduced number of
habitat types, the estimated species richness of
Gorringe was 42, about half of that for the Selvagens
Islands. These results strengthen the conclusion that
in Macaronesia islands or seamounts of small or very
small size may harbour a substantial fraction of the
species richness of this region.
It is likely that both the Selvagens Islands and the
Gorringe Bank are within the dispersal range of
many fish species that can be exported from other
locations, in particular those that are at a shorter dis-
tance. The closest land masses to the Selvagens
Islands are the Canary Islands (153 km) and the
closest land mass to the Gorringe Bank is that of
mainland Portugal (228 km). It remains unclear,
however, whether the Selvagens Islands and the
Gorringe seamounts sustain their own populations
or if they are supplied from other sources and act
as sinks sensu Pulliam (1988). In 2010 we found 57
of the 60 species described by Falcón et al (2000),
but if colonization is frequent the finding of the
same species in a given area in different years does
not demonstrate that populations are self-recruiting.
Also, we failed to detect three species: Mugil cephalus
(Mugilidae), Parablennius incognitus (Bath, 1968)
(Blenniidae) and Thorogobius ephippiatus (Lowe,
1839) (Gobiidae). Either these species were present
in low numbers and we have failed to detect them
during the campaign (because the fish assemblages
are dominated by a small number of species and
species richness is highly influenced by rare or very
rare species), or there are changes in the community
structure of the Selvagens Islands from year to year.
Additional surveys of the Selvagens Islands ichthyo-
fauna and fine-scale molecular studies may eventu-
ally answer this question in the future.
When comparing data from the Selvagens Islands
ichthyofauna with similar data from the Canary
Islands (Brito et al. 2002), Madeira (Wirtz et al.
2008) and the Azores (Porteiro et al. 2010), the high
affinity between the first two regions becomes clear.
Indeed, all species listed for the Selvagens Islands
are also reported for the Canary Islands, while the
corresponding numbers for Madeira are smaller.
This is to be expected when we consider that the
Canaries are the nearest neighbour to the Selvagens
Islands (153 km) while Madeira and the Azores are
at a distance of 251 km and 1127 km, respectively.
Also, the Canary Islands are the largest and the
oldest of the Macaronesian archipelagos and have
the highest species richness, yielding 258 coastal fish
species, while Madeira has 186 species and the
Azores 148 species (Almada et al. 2013).
The high species richness of the Canary Islands is
also likely to favour this archipelago as the dominant
source of fish propagules (e.g. eggs and larvae)
reaching the Selvagens Islands. Although the Canar-
ies Current (north to south) and northwest winds are
predominant, other current patterns that may trans-
port these propagules from the Canary Islands
northward to the Selvagens Islands are also common
in this region (Stramma 1984; Morton et al. 1998).
These results are congruent with the colonization of
the Macaronesian Islands through a stepping-stone
process, as proposed by Santos et al. (1995) and
Almada et al. (2001). Overall, the high affinity between
the ichthyofauna of the Macaronesian Islands and
the west African coast parallels the biogeographic
patterns found with other taxonomic groups (see
Morton & Britton 2000; Tuya & Haroun 2009).
Excluding cosmopolitan species, some species
reported for the Selvagens Islands are widely dis-
tributed along the western Atlantic coast but are not
reported for the eastern Atlantic continental coast.
These species only reach the archipelagos of the
eastern Atlantic, including Cape Verde, São Tomé
and Principe. Although they represent a small
percentage of the total number of fish species from
these islands, we must bear in mind that this
transatlantic route is more than 10 times (closest
American continental coast: 3669 km) the distance
between the west African coast and the Selvagens
Islands (353 km). This presumptive eastward colon-
ization process could be favoured by the North
Equatorial Countercurrent (see Fonseca et al. 2004).
When considering the potential capability of each
species to colonize remote regions over long
stretches of oceanic waters, no trend toward highly
mobile pelagic fish can be found. In fact, most
species reported for the Selvagens Islands are ben-
thic or demersal and sedentary fish. Even consider-
ing that a few benthopelagic species may be found
offshore associated with floating objects, e.g. Balistes
capriscus Gmelin, 1789 and Canthidermis sufflamen
(Mitchill, 1813), only a small number of species are
able to cross the oceanic barrier between the west
African coast or the Canary Islands and the Selva-
gens Islands.
On the other hand, when considering reproductive
biology, most species reported for the Selvagens
Islands spawn pelagic eggs or are viviparous and
ovoviviparous, mobile or very mobile species
(63.6%, n= 56). These fish may cross oceanic
barriers, since there are no apparent barriers to the
transport of eggs or breeding fish by marine cur-
rents. The high number of species that spawn pelagic
eggs could represent a particularity of the fish
communities of small and isolated islands.
10 F. Almada et al.
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The complexity of fish distribution patterns in
Macaronesia is exemplified by the blenniids (the
adults being sedentary and unable to cross deep
water), with different species occurring in particular
combinations of island groups but being absent in
others which, judging by distance alone, seem to be
accessible to eggs and larvae coming from the
nearest island group. For instance, Salaria pavo
(Risso, 1810) and Parablennius pilicornis (Cuvier,
1829), which occur in the Canary Islands and the
shores of southwest Europe and northwest Africa, do
not reach the Selvagens Islands. On the contrary,
Scartella cristata (Linnaeus, 1758) shows a distribu-
tion that includes the Mediterranean, northwest
Africa, both shores of the South Atlantic, the Canary
Islands and the Selvagens Islands but not Madeira or
the Azores. Parablennius ruber (Valenciennes, 1836),
a very common blenniid at the Azores, occurs in
western Europe, Madeira and Gorringe, but does
not reach the Selvagens Islands and the Canaries.
This diversity of distribution patterns in a single
family suggests that interspecific differences in life
history patterns, larval behaviour, ecology or even
stochastic events may constrain each species to cope
differently with transport to and survival at these
islands (see Cowen 2002; Cowen & Sponaugle
2009). In a paper comparing reef fish communities
across the Atlantic Ocean, Floeter et al. (2008)
concluded that examples of recent dispersal out-
number those of historical events, although both
factors have a strong influence on Atlantic fish
diversity and biogeographic patterns.
In the scenario outlined above it is not surprising
that no endemic fishes are known for the Selvagens
Islands or the much larger Canary Islands. Macar-
onesian endemic species reported for other islands
(e.g. Scorpaena azorica Eschmeyer, 1969 for the
Azores) are considered doubtful reports because
they are based on the description of a single
specimen (Santos et al. 1997).
In the future, additional surveys are essential to
improve our knowledge of the ichthyofauna of the
Selvagens Islands in order to better understand fish
dispersion in this geographic area. Additional data
on the pelagic larval development and behaviour,
together with a comprehensive phylogeographic
study of the ichthyofauna from the Macaronesian
islands, will improve our understanding of fish
distribution patterns and processes in the north-
eastern Atlantic and Mediterranean.
Funding
The EMEPC/M@rbis/Selvagens 2010 oceanographic
expedition was funded by EMEPC, Task Group for the
Extension of the Continental Shelf under the Portuguese
Ministry of National Defence, Eco-Ethology Research
Unit (331/94) pluriannual financing programme (Funda-
ção para a Ciência e Tecnologia, partially FEDER
funded). The research of FA (SFRH/BPD/63170/2009),
DA (SFRH/BD/46286/2008), SH (SFRH/BD/47034/2008),
MP (SFRH/BD/46639/2008), MB (SFRH/BD/64395/
2009), BHC (SFRH/BD/41262/2007), HA (BD/47055/
2008) and PC (PTDC/MAR/71927/2006) was supported
by grants from the Portuguese Science and Technology
Foundation (FCT) and DVR was partially financed by a
FPU scholarship from the Spanish Minister of Science and
Technology. Thanks are also due to the Parque Natural da
Madeira and their wardens who provided crucial logistic
support during the expedition.
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