Zootaxa 2680: 33–44 (2010)
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ISSN 1175-5326 (print edition)
Article
Copyright © 2010 · Magnolia Press
ZOOTAXA
ISSN 1175-5334 (online edition)
Zullobalanus santamariaensis sp. nov., a new late Miocene barnacle species of the
family Archaeobalanidae (Cirripedia: Thoracica), from the Azores
KAI WINKELMANN1,2,4, JOHN S. BUCKERIDGE3, ANA CRISTINA COSTA5, MARIA ANA MANSO
DIONÍSIO5, ANDRÉ MEDEIROS4 , MÁRIO CACHÃO6 & SÉRGIO P. ÁVILA4,5
1
Faculty of Science, University of Bristol, UK
Natural History Museum London, UK. E-mail: [email protected]
3
Earth & Oceanic Systems Research Group, RMIT University, Melbourne, VIC 3001, Australia.
E-mail: [email protected]
4
MPB, Marine PalaeoBiogeography Working Group of the University of Azores, Departamento de Biologia, Universidade dos Açores,
9501-855 Ponta Delgada, Azores, Portugal. E-mail: [email protected]; [email protected]
5
CIBIO- Pólo Açores, Departamento de Biologia, Universidade dos Açores, Rua Mãe de Deus, 9501-855 Ponta Delgada, Azores,
Portugal. E-mail: [email protected]; [email protected]
6
Centro de Geologia e Departamento de Geologia da Faculdade de Ciências. Universidade de Lisboa. Rua da Escola Politécnica, 58,
1250-102 Lisboa, Portugal. E-mail: [email protected]
2
Abstract
A new species of Archaeobalanidae barnacle is described from the late Miocene and early Pliocene of Santa Maria Island
(Azores). Samples were collected primarily from outcrops at Malbusca and “Pedra-que-Pica”. Zullobalanus
santamariaensis sp. nov. is endemic to the Azores archipelago and until the discovery of this material, the genus was
known only from the Southern Hemisphere. For such a long-range distribution we assume a combination of
planktotrophic larval development, sea-surface oceanic currents and transport by cetaceans. This study endorses the
elevation of the subgenus Zullobalanus to generic level.
Key words: Balanomorpha, Archaeobalanidae, Zullobalanus, Azores, Miocene
Introduction
The Azores archipelago is located in the central North Atlantic Ocean and consists of nine volcanic islands.
Of these, Santa Maria (lat. 37º 23’ N; long. 24º 45’ W), which has rocks dated as having been formed 8.12
million years ago (Abdel-Monem et al. 1975; Feraud et al. 1980), is the only island with a marine fossil
record.
Although palaeontological research in the Azores began in the second half of the 19th century, scientific
reports on the subject are rare until the turn of the 20th century (Callapez & Soares 2000; García-Talavera
1990; Madeira et al. 2007). In 2000, researchers from the Department of Biology of the University of the
Azores visited the fossil outcrops of Santa Maria Island and found well preserved marine fossil assemblages
of late-Miocene/upper-Pliocene and Pleistocene age (Ávila et al. 2002; Ávila et al. 2007; Ávila et al. 2008a;
Ávila et al. 2008b; Ávila et al. 2009a; Ávila et al. 2009b; Ávila et al. 2010; Kirby et al. 2007; Janssen et al.
2008; Kroh et al. 2008). Since 2002, the Marine PalaeoBiogeography Working Group of the University of the
Azores has organized “Palaeontology in Atlantic Islands”, a series of international workshops held on Santa
Maria Island to study the geology and geomorphology of the island, the petrology and geochemistry of
magmatic rocks, the volcanostratigraphy, the palaeontology, the palaeoecology and the palaeobiogeography
of invertebrates within these strata (i.e. bryozoans, foraminifera, molluscs, echinoderms and crustaceans –
although the latter did not include fossil barnacles). Over the last 150 years at least 12 expeditions to the
Azores have described the recent barnacle fauna (Southward 1998) but apart from brief mention of Balanus
Accepted by D. Jones: 26 Oct. 2010; published: 18 Nov. 2010
33
laevis Bruguière, 1789 and further Balanus species as fossils (Zbyszewski & Ferreira 1962; Callapez &
Soares 2000) there is no detailed description of fossil cirripedes from the island.
This paper reviews the material collected during the 2006–2009 “Palaeontology in Atlantic Islands”
expeditions and describes the first new fossil cirripede species recorded from the Azores: an archaeobalanid
from the Miocene of Santa Maria Island.
Material and methods
Location and geological setting. Santa Maria Island is the most southeastern island of the Azores
archipelago; it has an area of 97 km2, with a maximum length of 16.8 km (França et al. 2003). The oldest part
of the island, the Cabrestantes Formation, was formed by submarine volcanic activity and can be found on the
west side of the island. After a marine regression, erosion resulted in the peneplanation of this area.
Subsequent intensive subaerial volcanic activity formed the Anjos Complex, which enlarged the island in the
north and gave it a square shape, very different from today. About 5 Ma ago a marine transgression occurred
and submarine lavas (the Touril Complex) were erupted on the south side of the island. The most recent
volcanic activity took place during the lower Pliocene (the Facho-Pico Alto Complex) when explosive
volcanic activity occurred and formed the island’s current shape as well as its highest point, Pico Alto, at 590
m. Today old beach deposits and extensive wave cut platforms are visible in the western part of the island up
to heights of 120 m above sea level. Pillow-lavas (submarine lavas) are exposed along the shore of Santa
Maria Island, from the intertidal (although they are also known subtidally) up to 100 m (e.g., at Malbusca, in
the south), testifying for a progressive uplift of the island since the end of the Pliocene. A more detailed
description of the evolution of Santa Maria Island is given in Madeira (1986), Serralheiro & Madeira (1990)
and Serralheiro (2003).
Figure 1 illustrates the fossiliferous outcrops of Santa Maria Island visited and fossil barnacle remains
collected during the 5th (2008) and 6th (2009) workshops “Palaeontology in Atlantic Islands”.
FIGURE 1. Fossiliferous outcrops of Santa Maria sampled by MPB team (© Secção de Geografia, Universidade dos
Açores).
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WINKELMANN ET AL.
Sampling. Barnacles were found either as whole specimens attached to a substrate and/or as single
fragments. At Malbusca and at Cré, sediment samples were taken and sieved using mesh sizes of 2 mm, 1 mm
and 0.5 mm. After this process, a number of well-preserved barnacle opercula were obtained, which are of
significant value in determining taxonomy. SEM images were taken with a JEOL JSM-5410 Scanning
Microscope (SEM) and acquired with SemAfore at the University of the Azores in July 2009. Apart from
paratype material, see below, the holotype and additional material collected was deposited at DBUA-F (the
fossil collection of the Department of Biology, University of the Azores).
Table 1 summarizes the information on the outcrops (e.g.: altitude, GPS coordinates and the geologic
complex). This table also gives the altitude where the fossils were found and the code attributed for the fossil
collection of the University of the Azores (DBUA-F).
TABLE 1. List of all the samples of fossil barnacles (Zullobalanus santamariaensis sp. nov.) collected in Santa Maria
Island. For all information regarding the outcrops, dates and authors of collecting the samples, please consult http://
www.mpb.uac.pt/PDFs/collections/DBUA-F.pdf.
Collection site
Altitude of the
outcrops (m)
Collection
altitude (m)
GPS coordinates
Volcanostratigraphic
Complex (after Madeira,
1986)
DBUA-F Collection
number
Ponta do Castelo
0–6
2–3
25º 00‘ 59.443319‘‘W
36º 55‘ 43.823011‘‘ N
Facho-Pico Alto Complex
660
“Pedra-que-Pica“ 0–2
1–2
25º 01‘ 29.730591 W
36º 55‘ 48.323454‘‘ N
Facho-Pico Alto Complex
96, 106, 108, 185, 186,
187, 200, 202, 210, 237,
317, 318, 389, 498, 499,
509, 526, 527, 544, 661,
662
Malbusca
30–50, 90
30
25° 4'7.04"W
36°55'45.53"N
Touril Complex
164, 170, 179, 188, 191,
192, 193, 201, 251, 252,
292, 295, 298, 299, 473,
475, 478, 479, 494, 528,
530, 531, 536, 621, 659
Figueiral
80–110
80
25º 07‘ 43.014356 W
36º 56‘ 52.694857 N
Touril Complex
198
Cré
90–130
100
25º 08‘ 10.700918‘‘ W
37º 00‘ 04.040618‘‘ N
Touril Complex
663, 664
Ponta dos Frades
20–100
50
25º 08‘ 46.675793‘‘ W
37º 00‘ 35.218918‘‘ N
Touril Complex
658
Calcareous nannofossils biostratigraphy. Facies of the fossiliferous outcrops near the coast at Malbusca
and Cré were deposited under conditions of high bottom hydrodynamics and as such are generally too coarse
to yield abundant and diversified assemblages of calcareous nannofossils. Because of this, taphonomic
material was not collected as vertical samples but laterally, along the most favorable beds, avoiding the basal
or topmost horizons. Each sample was prepared following the rippled smear slide procedure described in
Cachão & Moita (2000) and the nannofossil assemblages screened for biostratigraphic markers at x1,250
magnification under a petrographic optical microscope. For taxonomic references see Bown (1998) and
Perch-Nielsen (1989).
Systematics
Subclass Cirripedia Burmeister, 1834
Superorder Thoracica Darwin, 1854
Order Sessilia Lamarck, 1818
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Superfamily Balanoidea Leach, 1817
Family Archaeobalanidae Newman and Ross, 1976
Genus Zullobalanus Buckeridge, 1989
Solidobalanus (Withersella) Buckeridge, 1983: 354
Solidobalanus (Zullobalanus) Buckeridge, 1989: 704
Zullobalanus. — Newman, 1996: 502
Diagnosis. Archaeobalanids with six solid compartments, radii with denticulate sutural edges, basis solid,
calcareous; tergal spur narrow, removed from basi-tergal angle by its own width or more; scutum with
moderately weak to wanting adductor ridge and shallow, pyriform pit for lateral depressor muscles.
Age and distribution. ?Lower Oligocene, Middle Oligocene to Pliocene, Australia and New Zealand;
Lower Miocene to Upper Pliocene, Azores.
Type. Solidobalanus (Withersella) cudmorei Buckeridge, 1983: 91. Lower Pliocene, South Australia.
Remarks. Until the discovery of this material, Zullobalanus had been collected only from the Southern
Hemisphere. The occurrence in the North Atlantic indicates that this genus had a wider distribution than
anticipated.
Zullobalanus santamariaensis Buckeridge & Winkelmann sp. nov.
(Figures 2–3)
Diagnosis. Zullobalanus with shell lacking external apico-basal ribbing; tergum only weakly reflexed
apically, but possessing extensive, well-formed crests for depressor muscle attachment; scutum with
moderately produced articular ridge and moderate to weak adductor muscle scar.
Distribution and age. Azores; Late Miocene – Early Pliocene.
Type material. Holotype containing 2 sample compositions: DBUA-F 536.1, complete shell (3.23 mm in
height and 3.39 mm in basal diameter), carina (5.44 mm in height), carinolatus, (5.36 mm in height), latus
(1.84 mm in height), rostrum (2.59 mm in height), scutum (2.50 mm in articular margin), tergum (3.44 mm in
articular margin); DBUA-F 536.2, complete shell (6.31 mm in height and 6.63 mm in basal diameter), carina
(4.89 mm in height), carinolatus, (4.79 mm in height), latus (3.36 mm in height), rostrum (2.30 mm in height),
scutum (2.43 mm in articular margin), tergum (3.61 mm in articular margin); Santa Maria, Malbusca outcrop
(25°4'7.04"W, 36°55'45.53"N [30 m asl]); S.P. Ávila & K. Winkelmann, 22–29 June 2008. Paratypes all with
same locality, collectors and date as the holotype: Natural History Museum Berlin: MB.A 1733, MB.A 1734;
Natural History Museum Vienna: NHMW 2010/0088/0001, NHMW 2010/0088/0002; Royal Belgian Institute
of Natural Sciences Brussels: IRSNB 7255a–g, IRSNB 7256a–g; Natural History Museum London: NHM IC
549 – NHM IC 562; Muséum National d’Histoire Naturelle Paris: MNHN A32053, MNHN A32054.
Description. Shell conic, height approximating basal diameter; exterior rough, longitudinal ribbing weak
to absent, lateral growth lines weak; orifice rhomboidal (Fig. 2A, B); interior with strong ribbing, with minor
bifurcation of some ribs near basis; rib cross-section showing simple arborescent interlaminate structures (Fig.
2B, C); sheath smooth, well developed, pendant, occupying up to 50% of height of inner surface; basis firmly
interlocking with parietal ribs; radii moderately narrow, with oblique summits, growth lines parallel to basis
crossed by fine growth striae parallel to alar abutment (Fig. 2B); alae with marginal extensions, summits
almost horizontal (Fig. 2B).
Scutum (Fig. 3A, B) very weakly reflexed apically; exterior with strong transverse growth ribs crossed by
weak apico-basal striae; interior with articular ridge moderately produced, slightly pendant basally; adductor
ridge weak, broadly rounded; adductor muscle scar moderate to weak, pit for lateral depressor muscles well
formed lacking any obvious muscle attachment crests. Tergum (Fig. 3C, D) moderately elongate, spur wellformed, removed from basi-scutal angle by own width and extending beyond same angle by approximately its
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own width, furrow well-formed distally, externally with well spaced growth lines, in some areas (especially
basally), fine apico-basal striae present; interior with longitudinal furrow on articular margin wide, open,
shallow, smooth except for weak growth striae; articular ridge well-formed, concave towards articular margin;
crests for depressor muscles well developed basally, extending as a zone of raised papillae nearer the apex;
articular ridge gently curved to extend slightly beyond a broadly concave basal margin; apex with
approximately 3–4 strong transverse growth lines in adults.
FIGURE 2. Zullobalanus santamariaensis sp. nov. All specimens from lower-Miocene / early-Pliocene of Malbusca
outcrop (Santa Maria Island, Azores, Portugal); samples DBUA-F (see. Table 1). I: Zullobalanus santamariaensis sp.
nov. lateral and apical views. Letters at apical view; C = carina, CL = carinolatus, L = latus, R = rostrum. II: C - R; left
interior, right exterior view. III: wall from below. Scale: I = 10 mm, II = 1 mm, III = 200 µm.
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FIGURE 3. Zullobalanus santamariaensis sp. nov. All specimens from lower-Miocene / early-Pliocene of Malbusca
outcrop (Santa Maria Island, Azores, Portugal), from samples DBUA-F (see Table 1); drawings based on SEM pictures.
A: moveable scutum exterior (enlarged area showing detail from the lower part of the exterior of the scutum,
accentuating fine apico-basal striae), B: moveable scutum interior, C: moveable tergum exterior, D: moveable tergum
interior.
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Occurence at Santa Maria Island. All the Late-Miocene and Early-Pliocene outcrops studied in Santa
Maria contained the endemic barnacle Zullobalanus santamariaensis sp. nov. Most specimens were found at
Malbusca and at “Pedra-que-Pica” outcrops. The beds at “Pedra-que-Pica” contained some well preserved but
rare barnacles with opercula cemented to diverse marine molluscs (e.g. Gigantopecten latissimus (Brocchi,
1814) and Lopha plicatuloides (Mayer, 1864)); the beds at Malbusca contained a number of loose opercula in
a sandy matrix. At Cré outcrop, rare single walls as well as opercula were present. In the more lithified marine
sediments of Figueiral and Ponta do Castelo, barnacle fragments were only observed on eroded surfaces. At
Ponta dos Frades, a few barnacle fragments were found together with well preserved micro-molluscs on
eroded bottom surfaces. Nevertheless, we detected fragments of Zullobalanus santamariaensis sp. nov. in
every sampled Miocene/Pliocene outcrop.
Remarks and affinities. There is an extensive archaeobalanid fauna known from western Europe
(Carriol, 2008) and the eastern seaboard of North America (Zullo 1984; Zullo & Kite 1985; Zullo & Perreault
1989; Zullo & Portell 1991). Of these, species of Actinobalanus are distinguished from this taxon by their
porous bases; Hesperibalanus, the terga of which lack the well-formed nodose papillae on the internal surface
and the broad spur; Solidobalanus by the externally smooth compartments; and Lophobalanus by their very
narrow radii.
Zullobalanus santamariaensis sp. nov. differs from all other Zullobalanus species by a lack of clearly
formed external ribbing on the shell. Although there are some weak “ribs” these are somewhat obscured by
the generally rough surface of the shell. The weakly reflexed scutal apex is close to that seen in the Australian
species Zullobalanus australiae victoriae (Buckeridge, 1983) and in most respects it is closest to this species.
In addition to the lack of external ribbing, it differs from Z. australiae victoriae by the presence of numerous
well-formed crests for depressor muscles on the tergum.
The scutal articular ridge is often basally pendant in larger specimens of Zullobalanus species
(Buckeridge 1983), and although this has not been clearly seen in the material available, we should be mindful
that all larger scuta are isolated and since shell disarticulation, have been abraded, as such wearing away
delicate extensions.
Etymology. Geographic: Island of Santa Maria (noun in apposition).
Discussion
Stratigraphic range of Zullobalanus santamariaensis sp. nov. The abundance of suitable Miocene/Pliocene
and Pleistocene (MIS5e) 1 outcrops of benthic habitats around the island of Santa Maria permits a
determination of the stratigraphic range (Fig. 4) of Zullobalanus santamariaensis sp. nov. The fossil
assemblage of Malbusca, located on the south of the island, and in which Z. santamariaensis sp. nov. occurs,
was dated as Messinian (nannofossil biozone NN11; ~6.1–6.4 Ma), based on the presence of calcareous
nannofossils, in particular on the presence of the species Reticulofenestra rotaria (Theodoridis, 1984), while
“Pedra-que-Pica” on the south eastern corner of Santa Maria has been dated by Strontium isotope
chronostratigraphy (87Sr/86Sr) as 5.51±0.21 Ma (Kirby et al. 2007). The Cré outcrop was dated as Zanclean
(5.33–3.60 Ma), based on holoplanktonic mollusc ranges (Janssen et al. 2008). Thus, the record of Z.
santamariaensis sp. nov. is restricted to lithologies from the latest Miocene to early Pliocene (Zbyszewski &
Ferreira 1961; García-Talavera 1990; Callapez & Soares 2000; Ávila et al. 2002). It is unknown from more
recent deposits, even though suitable facies are known, e.g. the Pleistocene Lagoinhas and Prainha outcrops,
both dated by Ávila et al. (2008a) as from the MIS 5e.
Biogeography and distribution. In spite of the occurrence of fossil barnacles in several Miocene and
Pliocene outcrops along the European and American Atlantic coasts (Pilsbry 1930; Ross 1964; Zullo 1984;
Zullo & Kite 1985; Zullo & Perreault 1989; Zullo & Portell 1991; Carriol 2005; 2008), no taxa systematically
close to Zullobalanus santamariaensis sp. nov. are known so far, leading us to conclude that it was endemic to
1.
MIS5e —Marine Isotopic Substage 5e, that is, 130–117 ka
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the Azores archipelago. Endemism in the fossil record of Santa Maria Island is also know in gastropods from
the Miocene (eg. Cardium hartungi (Bronn in Hartung, 1860)).
Most of the species associated with Z. santamariaensis sp. nov. (e.g., the sea-urchin Clypeaster altus
(Lamarck, 1816), the bivalves Gigantopecten latissimus (Brocchi, 1814), Manupecten pesfelis (Linné, 1758)
and Hinnites crispus (Brochi, 1814)) are examples of thermophilic species, which have disappeared from the
Azores after the mid-Pliocene and Pleistocene cooling events (Raffi & Monegatti 1993; Landau et al. 2007;
Landau et al. 2009). Most of these species lived in shallow waters, thus we infer that the endemic barnacle Z.
santamariaensis sp. nov. also was a shallow water taxon, and this is not incompatible with Zullobalanus
species from Australasia (Buckeridge 1983). Zullobalanus has its earliest record in the early Oligocene of
New Zealand. Shortly after this, it is recorded from nearby Tasmania, Australia, and then from the Miocene of
Victoria (Buckeridge 1983).
FIGURE 4. Malbusca outcrop biostratigraphic interval, based on a checklist of species of calcareous nannofossils.
The distribution of other balanomorph barnacles may provide clues as to the disjunct distribution of
Zullobalanus. During the middle Cenozoic ocean currents are known to have dispersed elminiine barnacles
from the Australasian region to the southern Atlantic (Buckeridge & Newman 2010). The tetraclitid
Tesseropora also has a disjunct distribution – although primarily an Indo-Pacific genus a single species,
Tesseropora atlantica (Newman & Ross, 1977), is known from the Azores (Newman & Ross 1977). The
genus Tesseropora survived on Atlantic coast of France during the Miocene with T. dumortieri (Fisher, 1866)
and the Pliocene with T. sulcata (Carriol, 1993) (see Carriol 1993; 2008). The dispersal systems that operated
for Tesseropora are less clear than with the Elminiinae, for although Tesseropora is found in strata dated late
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WINKELMANN ET AL.
Oligocene-early Miocene in the southwest Pacific (Buckeridge 1983), the genus also occurs at about the same
time (or even slightly earlier) in Italy (De Alessandri, 1895). Tesseropora atlantica is, however, a nonplanktotrophic species and this would drastically minimize any likelihood of long-range dispersal as larvae.
Nonetheless, it has a wide-range distribution in the Atlantic, being recorded from Saint Paul’s Rocks in the
equatorial Atlantic (Edwards & Lubbock 1983a; Edwards & Lubbock 1983b), Bermuda (Newman & Ross
1977), Madeira archipelago (Wirtz et al. 2006) and the Azores (Newman & Ross 1977; Southward 1998;
Costa & Jones 2000). Recently, Cardigos et al. (2006) re-evaluated the introduction status of T. atlantica in
the Azores and have classified it as cryptogenic, stating that it was probably introduced in the islands by boat
hulls. However, and in spite of Tesseropora atlantica possessing a very short dispersal larval phase, it is
present with well established populations in all Azorean islands and in several spots, some of them quite far
from harbours. Moreover, populations are not dense and although frequent, are not very conspicuous, thus we
believe its presence in the Azores was not human mediated.
Zullobalanus santamariaensis sp. nov. may well have had a pelagic larvae and thus could have been
dispersed by sea currents from the Pacific to northern Atlantic (see Southward 1987); this record certainly
expands our knowledge of the potential for long-range dispersal of sessile barnacles. A combination of
probable planktotrophic larval development, sea-surface oceanic currents and transport of adult specimens on
cetaceans provides a plausible explanation for the arrival of the ancestral of Z. santamariaensis sp. nov. at
Santa Maria in the late Miocene. In support of this hypothesis are fossil whale remains reported in Estevens &
Ávila (2007) from Santa Maria Island (Cré outcrop) and identified as Balaenopteridae indet., Mysticeti indet.
and Mesoplodon sp. Many whales, especially balaenopterids such as the humpback Megaptera novaeangliae
(Borowski, 1781), are characterised by barnacle infestations. Whilst these barnacles are obligate commensals
on whales rather than parasites, they are probably irritating and along with ectoparasites are lost when whale
skin is shed. Bianucci et al. (2006) report remarkably rich deposits of coronulids, along with occasional
balanids, in what are interpreted as whale-breeding areas in Plio-Pleistocene Ecuador. Although no coronulid
remains have been recovered from the Santa Maria lithologies, it is not unreasonable to suspect that the
Azores was close to north-south cetacean migration routes during the late Miocene.
Most whale infesting barnacles are either coronulids, (e.g. Coronula diadema (Linnaeus, 1767)) or
lepadiforms (e.g. Conchoderma virgatum (Spengler, 1790)). Balanoid barnacles have been found attached to
Coronula species (as opposed to the cetacean’s skin), although this association is relatively rare. Two
specimens of Balanus trigonus (Darwin, 1854), attached to C. diadema, are figured in Newman & Ross
(1971: Pl. 3) and Bianucci et al. (2006: Fig. 1) illustrate Coronula with balanid barnacles attached, but it is
likely that the latter, along with associated bryozoans, colonised the barnacles after the whale’s skin had been
shed. Olsen et al. (2009) observed that Balaenoptera borealis (Lesson, 1828) apparently hitches a ride on the
ocean current conveyor belts for a considerable part of its migration following the path of major currents.
Thus, it is possible that the ancestral Z. santamariaensis sp. nov. could have reached the Azores passively
attached to whale barnacles that were subsequently dislodged near Santa Maria Island, ultimately establishing
a viable population.
Fossil barnacles of other Macaronesian islands, especially the Middle Miocene outcrops of Porto Santo
(Madeira archipelago) need to be further investigated in order to test for the endemic status of the Azorean
barnacle Zullobalanus santamariaensis sp. nov.
Acknowledgements
We thank the organization of the “Palaeontology in Atlantic Islands” for the logistics at Santa Maria Island
during the international workshops. We are grateful to Sara Silva, Pedro Monteiro and Patrícia Madeira (all
MPB team, University of the Azores) for support in fieldwork. We thank Direcção Regional da Ciência e
Tecnologia (Açores) and Câmara Municipal de Vila do Porto for financial support, and Clube Naval de Santa
Maria for providing sea transportation to reach the outcrops of Malbusca and “Pedra-que-Pica”. Jorge
Medeiros (CIRN/University of the Azores) is acknowledged for the SEM photos. We also thank William
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Newman for useful comments, including confirmation that the two unnamed balanids attached to the figured
Coronula diadema (in Newman & Ross, 1971) are Balanus trigonus. We thank the anonymous reviewers for
their helpful comments that improved this work. We thank also FCT project PTDC/MAR/102800/2008
(calcareous nannofossils).
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