AÇOREANA, 2007, Supl. 5: 59-73

THE MARINE FOSSILS FROM : AN HISTORICAL OVERVIEW

Patrícia 1, Andreas Kroh 4, António M. de Frias Martins 1, 2, 5 & Sérgio P. Ávila 1, 2, 3

1 MPB, Marine PalaeoBiogeography Working Group of the University of , Departamento de Biologia, Universidade dos Açores, 9501-855 , Azores, ; e-mail: [email protected] 2 Departamento de Biologia, Universidade dos Açores, Rua Mãe de Deus, 9501-855 Ponta Delgada, Azores, PORTUGAL 3 Centro do IMAR da Universidade dos Açores, 9901-862 , Azores, PORTUGAL; e- mail: [email protected] 4 Natural Museum Vienna, Department of & Palaeontology, Burgring 7, 1010 Vienna, AUSTRIA e- mail: [email protected] 5 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos - Pólo Açores, Departamento de Biologia, Universidade dos Açores, Rua da Mãe de Deus, Apartado 1422, 9501-855 Ponta Delgada, Azores, PORTUGAL; e-mail: [email protected]

ABSTRACT In the Azores, Santa Maria is the only island with a sedimentary cover in a nine island volcanic system. This southeast most island of the archipelago has a rich fossil record, dated to the Late -Early Pliocene and to the Pleistocene. Palaeontological investigations on the island started in the late nine- teen century. Nevertheless, through the next 150 years, the number of publica- tions and thus the attention given to the fossil record of Santa Maria shows con- siderable fluctuation over time. From the beginning, the Miocene-Pliocene outcrops, more numerous in the island, focused the attention of the scholars. Recently, however, the interest has shifted to the Pleistocene outcrops. Data obtained from these outcrops has been used extensively in the inter- pretation and discussion of the Azorean geological genesis and evolution, par- ticularly of Santa Maria, the oldest island of the archipelago. However, its role in the interpretation and discussion of the origin and subsequent evolution of insular marine communities has been meagre. The checklists produced for Santa Maria’s fossiliferous outcrops, account for a large spectrum of represented animal groups. Workers have focused their attention on the molluscs, being the group best represented in the fossil record. Data on other animal groups is still scarce, particularly in case of the Pleistocene outcrops. The fossiliferous beds of Santa Maria are far from being an exhausted matter and further research in the Island should be considered.

INTRODUCTION Archipelago, in the North-eastern Atlantic. Like the remaining eight anta Maria Island (Lat. 37º 23’ N; islands, it is of volcanic origin. SLong. 24º 45’ W) is located in the However, except for the small south-eastern most end of the Azores of (about 30 km to the NE 60 AÇOREANA 2007, Supl. 5: 59-73 of Santa Maria), Santa Maria is the With the turn of the 20th century only island where fossiliferous out- the palaeontological interest in Santa crops were detected. Maria Island decreased to almost In the 16th century, Gaspar total oblivion, with the exceptions of Frutuoso (1983) described a quarry at Friedlander (1929), and the reviews Figueiral, where the extracted cal- of Agostinho (1937a, 1937b). These careous sandstones had “seafood works focussing on the geological shells glued on it”. He also stated that origin of Santa Maria and its volcanic Santa Maria “limestone” was unique structures, but yielding no new in the archipelago, but of lesser qual- palaeontological data. ity for construction purposes, than The interest on the fossil outcrops imported limestone from the of Santa Maria rose again in the sec- Portuguese . Though ond half of the 20th century, expressed named as such by local people, the by a series of studies published in “limestones” of Santa Maria are rapid succession: Berthois (1950, mostly well lithified, lithic sand- 1951, 1953a, 1953b, 1953c), Ferreira stones and calcarenites. Pure calcare- (1952, 1955), Krejci-Graf et al. (1958) ous units, in contrast, are rare and and reproductions of the 19th century poorly developed, being restricted to studies by Teixeira (1950). This tin beds of bioclastic rud- and wacke- enhanced attention persisted through stones in most of the outcrops. the next decades, with the prolific ’s iterations, illus- publication of palaeontological trate the importance of these sedi- reports. During this time, Zbysze- mentary units for the inhabitants of wski et al. (1961) produced a geologi- these volcanic islands otherwise bare cal map with several explanatory of suitable building stone. notes, where the fossil contents of the The first scientific reports on the island outcrops were again discussed geology of this island date back to the (Ferreira, 1961a; Zbyszewski & 19th century, with the studies by Ferreira, 1961, 1962a, 1962b). After Bronn (1860), Hartung (1860) and these productive decades, the scien- Morelet (1860). The former two tific reports on the fossils of Santa authors produced the first thorough Maria Island became scarce again. geological studies on the island, with Some exceptions are the revisions on extensive descriptions of its geomor- the sedimentary rocks of the phology. These investigations, to- Macaronesian islands by Mitchell- gether with the ones of Reiss (1862), Thomé (1974, 1976, 1981), and the Hartung (1864) and Mayer (1864), papers of García-Talavera (1990) and provided the framework for the Callapez & Soares (2000) on the palaeontological work in Santa Maria Pleistocene outcrops of Prainha and during the 20th century. By the end of Lagoinhas. the 19th century, Cotter (1892) pub- In 2002, Santa Maria’s outcrops lished an update to the previously were revisited by a scientific expedi- published faunal lists. tion organized by elements of the MADEIRA ET AL: HISTORICAL OVERVIEW 61

Marine PalaeoBiogeographic Wor- with the highest point of the island at king Group (MPB) and of the Pico Alto (590 m) (Zbyszewski et al., Department of Biology of the 1961). The coast of Santa Maria is . The main characterized by steep cliffs, from 30 objectives were aimed at understand- m (Ponta do Marvão) to 342 m (Rocha ing the palaeoecology and palaeobio- Alta), in places interrupted by small geography of the Pleistocene and bays with sandy beaches (Praia, Miocene-Pliocene outcrops, as well as , São Lourenço and Anjos) towards the legal protection of the (Serralheiro, 2003). geological legacy of Santa Maria Hartung (1860) and Reiss (1862) Island. As a result, a checklist of the extensively described the geological Pleistocene molluscs of Lagoinhas landscape of Santa Maria, and located and Prainha was produced (Ávila et most of the fossil outcrops that we al., 2002), as well as a technical report know today. Almost one hundred for the protection of the outcrops of years later, Zbyszewski et al. (1961) Pedreira do Campo and Figueiral published a geological map with sev- (Cachão et al., 2003). The recently eral explanatory notes, where the fos- formed MPB intends to continue on sil contents of the island outcrops further research activities, e.g. the 2nd were again discussed. A new, more International Congress “Atlantic detailed geological map was pro- Islands ” (AINIC) that will be duced by Serralheiro et al. (1987) held at Ponta Delgada Azores, in focusing on the volcanic structures of September 2008, and the internation- Santa Maria. Recently, an update to al workshops “Palaeontology in this map was published (Serralheiro, Atlantic Islands” (this volume), all 2003). aiming at a better understanding of In his review, Mitchell-Thomé Santa Maria’s palaeontology and (1976) postulated the age of the stratigraphy. Azorean archipelago as Late Miocene or Early Pliocene. Abdel-Monem et al. The Geological History of Santa (1975) published radiometric (K-Ar) Maria Island age estimates for the volcanic rocks in Santa Maria is a relatively small the eastern Azores group. They dated island, with 97 km2 and a maximum the basaltic series exposed below the length of 16.8 km (França et al., 2003). fossiliferous sediments (“coquina The island can be divided in two dis- zone”) of Santa Maria to 6-8 Ma B.P. tinct physiographic regions: the west- A maximum estimate of 8.12 Ma was ern part, relatively flat with erosional obtained for the bellow the air- surfaces at different altitudes, and a port runway about 70 m above the more irregular eastern part (Abdel- present sea level. Although, the exact Monem et al. 1975). These regions are age of Santa Maria Island is still a separated by a central mountain matter of debate (e.g., Serralheiro & chain extending NNW-SSE between Madeira, 1990, Serralheiro, 2003), the Lagoinhas and Glória, coinciding data presented by these authors show 62 AÇOREANA 2007, Supl. 5: 59-73 that it is the oldest island in the sion of the eroded materials. Azorean archipelago, confirming the Afterwards, volcanic activity restart- assumptions made by Agostinho ed with increasing intensity and dis- (1937b), who considered that the dif- tribution, and contrary to the previ- ferential degree of deterioration of ous event, it was of subaerial nature, the volcanic rocks was possibly testi- leading ultimately to the formation of fying different periods of volcanic the Anjos Complex during the activity. Zbyszewski et al. (1961) Tortonian stage (11.61-7.25 Ma). The assumed three general phases in the Anjos Complex is responsible for history of the island formation: increasing Santa Maria about 3 km to pre-“Vindobonian”, “Vindobonian” the north, modelling the island to a (which includes the deposition of the somewhat square shape, with round Santa Maria sedimentary units) and corners, very different from today. post-“Vindobonian” (the “Vindobo- Serralheiro (2003) adds that the nian” stage is an outdated strati- Lagoinhas islets are a testimony to graphical term, corresponding to the the island growth, subsequently and Middle Miocene today). The discus- progressively destroyed by sea ero- sion on origin and evolution of the sion. Santa Maria continued throughout The eruptive period that lead to the late 20th century (e.g., Madeira, the development of Anjos Complex, 1986; Storetvedt et al., 1989; was followed by a relatively calm Serralheiro & Madeira, 1990). In the phase, characterised by the erosion following paragraphs the history of and weathering of the old volcanic the geological evolution of Santa landscape. Afterwards, the island Maria Island is given, based on the experienced a transgression, during review of Serralheiro (2003). the transition from the Messinian The Cabrestantes Formation in the (Late Miocene) to the Pliocene (~5 west of Santa Maria Island was con- Ma), when sea level reached at least sidered by Serralheiro (2003) to be the 180 m of altitude (Serralheiro, 2003). oldest formation in the island. On the south side the submarine Standing today at an altitude of no eruptions were intense, contrasting more than 33 m, this structure was with the highly erosive environment formed by initial submarine volcanic in the north, which placed the shore- activity. After the initial period of line back to the area near the volcanic activity, the sea level experi- Lagoinhas islets. The Touril Complex enced a regression, dropping below was formed during this long period, today’s level. This period was charac- including its thick sedimentary beds. terized by the erosion and weather- Volcanic activity restarted with ing of the exposed island. However, great intensity during the formation Serralheiro (2003) states that with of Facho-Pico Alto Complex (Lower today’s data it is not possible to Pliocene). This period was character- understand the magnitude or dura- ized by the explosive submarine vol- tion of the regression and the exten- canic activity, which raised the vol- MADEIRA ET AL: HISTORICAL OVERVIEW 63 canic cones of the Pico do Facho. Miocene, respectively. One hundred Volcanic activity continued through- years later, Zbyszewski & Ferreira out this period, spreading its materi- (1962b), placed them in the als from the Airport in the west of “Vinbodonian” (an outdated term Santa Maria, to the southeast in Baixa for the Middle Miocene). Based on do Sul and to the north in Ponta do the foraminiferal data of Colom Norte, increasing the size of the (1958 in Krejci-Graf et al., 1958) and island to its present dimensions. new radiometric dates, Abdel- Later volcanic activity shifted to the Monem et al. (1975) rejected these east-southeast part of the island and age assignments and proposed a changed to the subaerial type due to younger, Late Miocene to Early a marine regression. The eruptions Pliocene age (around 5.3 Ma). Aside were then interrupted by a relatively from the sub-recent sedimentary calm phase of erosion. deposits, the geological map of During the late Pliocene, volcanic Serralheiro et al. (1987) shows two activity restarted giving rise to the lithostratigraphic units composed of Complex, with great explo- sedimentary rocks of two ages: Late sive manifestations spreading Miocene and Pliocene. The former throughout the island. After this are mainly distributed on the west eruptive phase, another transgression part of the island, and the latter on occurred by the end of the Pliocene. the central and east areas, at alti- The sea level reached the altitude of tudes that can reach about 400 m 200 m, forming the old beach (e.g., in areas near Pico Alto). In deposits and extensive wave cut plat- 2003, Serralheiro reviews the data forms, especially visible in the west given by the earlier workers, includ- part of the island up to 120 m. Since ing the ones mentioned above by the Quaternary, Santa Maria has risen Zbyszewski & Ferreira (1962b) and relatively to sea level progressively, by Colom (1958 in Krejci-Graf et al., but not continuously, until the 1958), and agrees with the latter, today’s position (Serralheiro & assigning the Miocene deposits to Madeira, 1990). Presently, Santa the Messinian (Late Miocene) or pos- Maria shows no signs of recent vol- sibly early Pliocene. canic activity or any kind of sec- In general terms, the sedimentary ondary volcanism (Agostinho, deposits in Santa Maria consist of 1937b). horizontal layers intercalated in vol- canic material, and are represented The sedimentary stratigraphy of by limestones, breccias, sandstones, Santa Maria conglomerates and subaerial Mayer (1864) assigned the fossils deposits (Agostinho, 1937b; Ferreira, from the Miocene outcrops of Santa 1955; Mitchell-Thomé, 1976). Maria Island to the “Mayencian” Agostinho (1937b) claimed that the and “Helvetian” stages, today corre- maximum altitude reached by the sponding to the Middle and Lower outcrops was no higher than 130 m 64 AÇOREANA 2007, Supl. 5: 59-73

FIGURE 1. Map with the fossiliferous outcrops of Santa Maria Island (adapted from Ferreira, 1955, 1961a). above the sea level at Pico do Facho. The outcrops of Santa Maria Island Such was contradicted by Mitchell- In the west part of Santa Maria Thomé (1976), who based on the Island, the geological map of observation of the geological map by Serralheiro et al. (1987) shows the Zbyszewski et al. (1961), stated that presence of Upper Miocene and sedimentary rocks occurred as high Quaternary deposits on the area of as 400 m in Pico Alto. Berthois the Airport and Santana. Though, (1953c) in his extensive description these Quaternary beaches at about 90 of the lithology of the calcareous m of altitude can be easily correlated rocks of Santa Maria, believed that with the Airport deposits referred to the present elevations of the lime- in the bibliography (e.g., Berthois, stone depositions are partially origi- 1950; Zbyszewski & Ferreira, 1962a), nal. the same cannot be stated for the In the following paragraphs we Miocene limestone outcrops. Ferreira provide a description of the fossilif- (1961b) mentioned an old quarry of erous outcrops (see Fig. 1). The nam- relatively small extension in the area ing of the outcrops is based on the called Antigas Crés, which by its map by Serralheiro et al. (1987) and, position appears to be correlated to with few exceptions, is consistent the Acácias (or Assumada, as was with the historical works. called by the author) and Meio Moio MADEIRA ET AL: HISTORICAL OVERVIEW 65 outcrops, located to the North of the faunal diversity of Ponta dos Matos is island, near Baía da Cré. relatively low (Mayer, 1864; Ferreira, Baía da Cré, in the north of Santa 1955), yet that outcrop yielded taxa Maria Island, to the east of Baía dos not encountered anywhere else (e.g. Anjos, was noticed by the first inves- the brachiopod Terebratulina retusa tigators for its rich fossil content, par- [=T. caputserpentis of former ticularly the outcrops located in the authors]). From the first two locali- east part of the bay - Boca da Cré and ties, Zbyszewski et al. (1961) Pinheiros (Hartung, 1860; Reiss, 1862; described 3 layers of fossiliferous Mayer, 1864). Aside from those two brownish limestone, no thicker than 2 sites, additional outcrops were m, intercalated with basaltic lavas. studied the by Zbyszewski & Ferreira Following the coastline to the (1962a, b): Meio Moio, Acácias, south, a number of Miocene deposits Pedreira dos Frades, Monte Gordo, are documented: São Lourenço (on Ponta dos Frades and Casa da Cré or the southeast end of the bay), Rocha Escarpa da Cré. In general, these sed- Negra and Ponta da Rocha (see imentary deposits do not reach high- Ferreira, 1961a). The last outcrop was er that 60 m above present sea level referred also as by the name of (Ferreira, 1961a), with maximum of Pontinha or Ponta do Papagaio ~120 m at Casa da Cré (our data), and (Hartung, 1860, 1864; Krejci-Graf et about 100 m at Monte Gordo or Meio al., 1958; Zbyszewski et al., 1961). Moio (Teixeira, 1950). The sedimentary rocks in the area The Miocene sedimentary deposi- of Ponta das Salinas, known tradi- tions continue through the north tionally by the name of Feteirinha or coast of Santa Maria, to the east of Feteirinhas (Harthung, 1860; Reiss, Baía da Cré. In Baía do Raposo, the 1862; Mayer, 1864), lie about 30 m documented outcrops are located in above present sea level (Agostinho, the small valley formed by the 1937b) and are exposed in disused Ribeira do Engenho, and in the north- quarries described by Ferreira east extreme of the bay, Ponta do (1961a). Additional Miocene outcrops Pesqueiro Alto, referred also as on the east coast of Santa Maria are Tamuscal or Tamugal (Reiss, 1862; and (Zbyszewski & Ferreira, 1961a). The geology of this Ferreira, 1962a), which lie roughly area is discussed in greater detail by about 20 m above the sea level. Zbyszewski & Ferreira (1962a). Ferreira (1961a) in his survey of eco- In the north extreme of the island, nomical interesting limestone three Miocene localities have been deposits addressed these two as documented: Ponta do Norte, Ponta unsuitable. The deposits of Cedros de Badeus and Ponta dos Matos. were too small and Altares outcrop These deposits are described to some was at that time an almost exhausted extent by Reiss (1862), Ferreira quarry of extremely difficult access. (1961a), Zbyszewski et al. (1961) and To the south, sedimentary units only Zbyszewski & Ferreira (1962a). The appear again in the locality of Maia, 66 AÇOREANA 2007, Supl. 5: 59-73 mainly to the south-eastern end of the to the north outcrop, as done by island, at Ponta do Castelo. This area, Hartung (1860) and subsequent particularly to the west, in the direc- investigators. Ferreira (1961a) posi- tion of Ponta da Malbusca, is charac- tioned the Malbusca outcrop close to terised by thin beds of fossiliferous the sea level. However, Zbyszewski marine sandstone (and subordinate et al. (1961) and Zbyszewski & limestone) intercalated between vol- Ferreira (1962a) described a 180 m canic layers (basalt flows, pillow high cliff containing two sedimentary lavae and volcanoclastics) (Serra- deposits of 3.5, respectively 12 m lheiro et al., 1987). The Ponta do thickness, framed by volcanic lavas. Castelo section was described by According to their data, these layers Zbyszewski et al. (1961) and lie roughly 60 m above the present Zbyszewski & Ferreira (1962a). In the sea level, which agrees with the alti- area near Baixa do Sul, between tude given by Agostinho (1937b) for Ponta do Castelo and Rocha Alta, this outcrop and our own field obser- Serralheiro (2003) mentioned for the vations. first time a fossil outcrop about 3 m The area south of Pico do Facho, thickness which he called “Pedra- east to Vila do , is rich in sedi- que-Pica”. mentary strata (Serralheiro et al., Along the south coast of Santa 1987). In this area the Miocene out- Maria, near locality in an crops reach their highest elevations area of steep cliffs, there is an old (between 120 and 130 m above pre- limestone quarry (Ferreira, 1961a), sent sea level; Agostinho, 1937b; earlier referred to as Forno da Cré Ferreira, 1962). Nowadays, two out- (Hartung, 1860; Reiss, 1862; Mayer, crops are recognized, Pedreira do 1864; Cotter, 1892, 1953). Other desig- Campo and Figueiral, all included in nations were used too: Agostinho an area classified as a Natural (1937b) called it “Cré”, Ferreira Monument (Cachão et al., 2003). (1961a) used the name “Furna da Figueiral is well known since the first Cré” and Zbyszewski & Ferreira naturalists (Hartung, 1860; Reiss, (1962a, 1962b) called it “Boca da Cré”, 1862; Mayer, 1864), for it is the place a designation used as well for the of one of the oldest “limestone” north locality of Baía da Cré. quarries in Santa Maria and has been Zbyszewski & Ferreira (1962a, 1962b) mentioned commonly (Ferreira, argued that the use of “Bocca do Cré” 1952, 1955, 1961a; Zbyszewski & by Hartung (1860) to designate the Ferreira, 1962a; Zbyszewski et al., north outcrop was not correct, for the 1961). Pedreira do Campo (also local people use this name for the known as Pedreira do Facho) is situ- south locality near Piedade. In this ated near Figueiral, to the south of work we prefer to use the name given Pico do Facho. Although discovered in the map of Serralheiro et al. (1987), by García-Talavera (pers. com.) in Ponta da Malbusca, to prevent confu- 1997, it attracted the attention of sci- sions. We restrict the use Boca da Cré entists and the local community only MADEIRA ET AL: HISTORICAL OVERVIEW 67 after the first international expedi- outcrops: Praia and Prainha. The tion “Palaeontology in Atlantic sedimentary succession of Prainha Islands”, in 2002. A technical report was studied thoroughly by the was produced for the local govern- authors listed above. It is character- ment, in order to protect this outcrop ized by a basal conglomerate with and Figueiral (see Cachão et al., algal-limestone concretions, fol- 2003), and to stop stone extraction lowed by 0.8 to 3.0 m of fossiliferous from the Pedreira do Campo area. sands (interpreted as beaches In the area of Prainha, at an alti- deposits) rich in micro-molluscs tude between 100 to110 m above the (Ávila et al., 2002; Ávila, 2005). present sea level, Berthois (1953c) Zbyszewski & Ferreira (1961) and described Miocene sediments. The García-Talavera (1990) assigned the fossils contained are poorly pre- sediments of Prainha to the served, due to leaching and recrys- Tyrrhenian stage. tallization (aragonitic molluscs being The area near the Ilhéu da preserved in form of moulds only). Lagoinhas, to the west end of Baía In this work we call this outcrop by do Tagarete, in the north of Santa Macela, based on the name given to Maria Island, is relatively rich in the hills that surround the west part Pleistocene deposits. However, the of this bay, thus differentiating it first report on these depositions was from the Pleistocene deposits of made only in the turn of the millen- Prainha. nium by Callapez & Soares (2000). Although the Miocene outcrops These authors correlated this out- were given more attention since the crop, known as Lagoinhas, to the beginning of the palaeontogical Pleistocene outcrop of Prainha. work on Santa Maria, the Pleistocene Similar to the latter, the outcrop is deposits of Praia Formosa and located about 7 m above the present Prainha attracted even more atten- sea level and is characterized by a tion (Mayer, 1864; Berthois, 1950, basal conglomerate, coralline red 1951, 1953c; Ferreira, 1961a; algae crusts (algal biostromes of up Zbyszewski & Ferreira, 1961, 1962a; to 0.2 m thickness), followed by 0.7 Zbyszewski et al., 1961; García- m bioclastic cross-bedded sands rich Talavera, 1990; Ávila et al., 2001; in fossil molluscs (interpreted as Amen, 2002; Ávila et al., 2002; Amen beach deposits; Ávila et al., 2002). et al., 2005; Ávila, 2005). Located to Based on the malacofauna, Callapez the southeast of Pico do Facho, the & Soares (2000) assigned the exposed Pleistocene deposits lie 2 to Lagoinhas outcrop the Tyrrhenian 3 m above present sea level (García- stage, corroborating the dates for Talavera, 1990), and extend from Prainha. Both outcrops were later Prainha until the far west end of studied by Ávila and co-workers Praia Formosa (Zbyszewski & (2001, 2002, and 2005), giving further Ferreira, 1961). In the literature two details on their litho-stratigraphy names are used to designate these and fossil content. 68 AÇOREANA 2007, Supl. 5: 59-73

The fossils of Santa Maria Island: tively by Zbyszewski & Almeida molluscs and other groups (1950) and Ferreira (1961b), are the In the faunal lists produced only classical works on animal through the years, the Molluscs are groups other than the molluscs. The the best represented group from the number of works published on the outcrops of Santa Maria. Zbyszewski fossil microfauna is likewise rather & Ferreira (1962b) in their work on limited: Berthois (1950), Collom (in the Miocene fossils reported a total of Krejci-Graf et al., 1958) and Ferreira 188 animal species for the island: 32 (1960). Calcareous red algae, particu- foraminifers, 2 anthozoans, 4 echi- larly well developed in the noids, 2 annelids, 5 bryozoans, 1 bra- Pleistocene outcrops of Prainha and chiopod, 66 bivalves, 64 gastropods, 2 Lagoinhas, were only recently a tar- cirripeds, 1 decapod, 8 fishes and the get group for the studies by Amen undetermined remains of cetacean (2002) and Amen et al. (2005). In sum, vertebras and ribs. the fossil record of Santa Maria Berthois (1950) documented the Islands is without doubt an issue far presence of microfossils at Prainha from being exhausted. Additional (e.g. green algae, coccoliths and attention should be given to groups foraminifers), as well as the remains of organisms other than molluscs in of the macrofauna (e.g. fish scales, future studies. fragments of echinoid spines and tests, holothurian ossicles, corals debris, bryozoans, brachiopods, gas- The past Azorean communities and tropods and red algae). Aside from their biogeography the molluscs, Callapez & Soares Despited numerous faunal lists (2000) reported bryozoan, cirriped being available after almost 150 years and echinoid fragments from of investigations the palaeoecological Lagoinhas. The last addition to the and palaeobiogeographical implica- Pleistocene malacofauna of Santa tions of these faunas remained poorly Maria Island was given by Ávila et al. studied. Questions concerning the (2002), summing 89 records, includ- evolution of the Azorean fossil fauna ing 75 gastropods and 14 bivalves. and its relation to extant communities The particularly good fossil record are largely un-answered. of molluscs can be explained by a Biogeographical studies on the combination of initial abundance, archipelago, largely ignored palaeon- taphonomic biases and selective tological data (e.g., Boury-Esnault & investigations (most palaeontological Lopes, 1985; Prud’homme van Reine, studies on Santa Maria focused on 1988; Cornelius, 1992; Lopes et al., molluscs). 1993; Wirtz & Martins, 1993; Tittley & For the macrofauna existing in the Neto, 1995; Santos et al., 1995; Santos sedimentary outcrops of Santa Maria, et al., 1997; Ávila, 2000; Tittley & the reviews on the Portuguese Neto, 2006). Only in the most recent Miocene fishes and echinoids, respec- studies, Pleistocene mollusc faunas MADEIRA ET AL: HISTORICAL OVERVIEW 69 were included in an attempt to CONCLUSIONS understand Santa Maria’s palaeobio- geographical relationships (García- Palaeontological studies relating Talavera, 1990; Callapez & Soares, to Santa Maria Island depended pri- 2000; Ávila et al., 2001; Ávila et al., marily on few expeditions made in 2002; Ávila, 2005). the last two centuries and on valuable donations made by private collectors. Several palaeontological works were The Portuguese fossil collections produced during this time, especially Nowadays, there are five institu- about the fossil molluscs, but little is tions on the Portuguese territory known on other animal groups pre- with palaeontological collections sent in the outcrops. This is also true harbouring specimens from Santa for microfossils and nannofossils, Maria fossiliferous outcrops: the despite their biostratigraphic value. Museu Geologico (Geological During the 20th century, several Museum) (INETI) in , the workers targeted the numerous Museu de Zoologia - Museu de Upper Miocene-Lower Pliocene out- História Natural da Faculdade de crops in their studies. This tendency Ciências e Tecnologia da Univer- changed by the end of the millenni- sidade de Coimbra (MZ/MHN- um, with the production of numerous FCTUC) in Coimbra, the “Museu papers on the Pleistocene strata, dos Montanheiros” in Angra do many of which aimed at the under- Heroísmo (, Azores), standing of the biogeographical rela- the Museu Carlos Machado/ tionships of the archipelago through- História Natural (MCM(HN) in out its history. Thus, although this Ponta Delgada (São Miguel Island, recent attention on the Pleistocene Azores) and the Department of outcrops is still far from being Biology of the University of the exhausted, a renewed look must be Azores (DBUA-F) in Ponta Delgada given to the Miocene-Pliocene out- (São Miguel Island, Azores). In the crops. Mitchel-Thomé (1976) classi- Geological Museum (Lisbon) we fied the palaeontological situation in could locate the specimens studied Santa Maria Island, as “(…) a promis- by Cotter and the material collected ing field”, and it seems that it still by Zbyszewski and co-workers. The holds. houses the material from Lagoinhas collected by Soares. In the University of the ACKNOWLEDGEMENTS Azores it is deposited the most com- plete collection of Santa Maria fos- We are grateful to the organiza- sils, as a result of several expedi- tion of the 1st International Congress tions since 1998, organized by the “Atlantic Island Neogene” for finan- members of the Marine PalaeoBio- cial support. We thank Direcção geography Working Group (MPB). Regional da Ciência e Tecnologia 70 AÇOREANA 2007, Supl. 5: 59-73

(Regional Government of the Azores). Revista Española de Azores), Direcção Regional do Am- Micropaleontologia, 37(1): 63-70. biente e do Mar (Regional Go- ÁVILA, S.P., 2000. Shallow-water vernment of the Azores), CCPA/UA marine molluscs of the azores: (Centro de Conservação e Protecção Biogeographical relationships. do Ambiente / Universidade dos Arquipélago, Life and Marine Açores), Departamento de Biologia Sciences, Universidade dos Açores, da Universidade dos Açores, Câmara Supplement 2(Part A): 99-131. Municipal de (Santa Ponta Delgada. Maria Island), Clube Naval de Santa ÁVILA, S.P., 2005. Processos e Padrões Maria, “Nerus” and Geo-Fun” for de Dispersão e Colonização nos field support during our expeditions Rissoidae (: Gastropoda) dos to Santa Maria Island. Açores. PhD Thesis, Univerisdade S.P. Ávila was supported by grant dos Açores, x+329 pp., Ponta SFRH/BPD/22913/2005 (FCT - Fun- Delgada. dação para a Ciência e Tecnologia) ÁVILA, S.P., J.M.N. AZEVEDO & from the Portuguese government. R.G. AMEN, 2001. Quaternary marine molluscs of Santa Maria, Azores, Portugal: check-list and BIBLIOGRAPHY Paleobiogeographic remarks. First Island Ecossistems: a conservation AGOSTINHO, J., 1937a. Tectónica, and molecular approach: 22. Sismicidade e vulcanismo das ÁVILA, S.P., R. AMEN, J.M.N. ilhas dos Açores. Açoreana, 1: AZEVEDO, M. CACHÃO & F. 85-98. GARCÍA-TALAVERA, 2002. AGOSTINHO, J., 1937b. Sobre a Checklist of the Pleistocene tectónica da ilha de Santa Maria. Marine Molluscs of Praínha and Açoreana, 1: 281-285. Lagoínhas (Santa Maria Island, ABDEL-MONEM, A. A., L. A. Azores). Açoreana, 9(4): 343-370. FERNANDEZ & G. M. BOONE, BERTHOIS, L., 1950. Sur la présence 1975. K-Ar ages from the eastern d’une microfaune dans le calcaire Azores group (Santa Maria, São de Santa Maria (Açores). Açoreana, Miguel and the Formigas Islands). 4: 277-285. Lithos, 8: 247-254. BERTHOIS, L., 1951. Sur la présence de AMEN, R. G., 2002. Estudo da jazida basses terrasses marines dans fóssil da Prainha, Santa Maria, l’Archipel des Açores (Portugal). Açores. Tese de licenciatura, Comptes Rendus du 76ème Congrès des Universidade dos Açores, 51pp., Sociétés Savantes à Rennes: 101-106. Ponta Delgada. BERTHOIS, L., 1953a. Sur la reparti- AMEN, R. G., A. I. NETO & J. M. N. tion granulométrique des élé- AZEVEDO, 2005. Coralline-algal ments calcaires dans les sables des framework in the Quaternary of Açores (Portugal). Açoreana, 5(1): Prainha (Santa Maria island, 54-63. MADEIRA ET AL: HISTORICAL OVERVIEW 71

BERTHOIS, L., 1953b. Terrasses COTTER, J. C. B., 1892. Notícia de marines d’altitude +5 à +8 m dans alguns fósseis terciários da ilha de l’Archipel des Açores. Açoreana, 5: Santa Maria no Arquipélago dos 64-70. Açores. Comunicações da Comissão BERTHOIS, L., 1953c. Contribuition à de Trabalhos Geológicos de Portugal, l’étude lithologique de l’Archipel 2: 255-287. des Açores. Comunicações dos COTTER, J. C. B., 1953. Notícia de Serviços Geológicos de Portugal, alguns fósseis terciários da ilha de Portugal, 34: 5-64. Santa Maria. Açoreana, 5: 71-75. BOURY-ESNAULT., N & M. T. FERREIRA, M., 1960. Microforaminí- LOPES, 1985. Les Desmoponges feros do Quaternário de Santa littorales de l’archipel des Açores. Maria (Açores). Comunicações Annales dee lÍnstitut Océanographi- XXV Congresso Luso-Espanhol para que, 6l(2): 149-225. o progresso das Ciências, 3 pp., BRONN, H.G., 1860. Die fossilen Madrid. Reste von Santa Maria, der süd- FERREIRA, O. V., 1952. Os Pecti- lichsten der Azorischen Inseln. In: nídeos do Miocénico da ilha de Hartung, G. (ed.) 1860. Die Azoren Santa Maria (Açores). Revista da in ihrer äusseren Erscheinung und Faculdade de Ciência de Lisboa, 2ª nach ihrer geognostischen Natur, Série, C, 2(2): 243-258. 350 pp. Verlag von Wilhem FERREIRA, O. V., 1955. A fauna Engelmann, Leipzig. Miocénica da ilha de Santa Maria. CACHÃO, M., J. MADEIRA, C. Comunicações dos Serviços Geoló- MARQUES DA SILVA, J. M. N. gicos de Portugal, 36: 9-44. AZEVEDO, A. P. CRUZ, C. FERREIRA, O. V., 1961a. Aflora- GARCIA, F. SOUSA, J. MELO, M. mentos de calcário miocénico da AGUIAR, P. SILVA, R. MARTINS Ilha de Santa Maria (Açores). & S. P. ÁVILA, 2003. Pedreira do Comunicações dos Serviços Geoló- Campo (Santa Maria, Açores): gicos de Portugal, 45: 467-478. monumento natural. Ciências da FERREIRA, O. V., 1961b. Equinídeos Terra (UNL), nº V: 120-123. do Miocénico de Portugal Con- CALLAPEZ, P. & A. F. SOARES, tinental e Ilhas Adjacentes. Comu- 2000. Late Quaternary warm nicações dos Serviços Geológicos de marine mollusks from Santa Portugal, 45: 529-564. Maria (Azores): paleoecology and FRANÇA, Z., J. V. CRUZ, J. C. paleobiogeographic considera- NUNES & V. H. FORJAZ, 2003. tions. Ciências da Terra (UNL), 14: Geologia dos Açores: uma 313-322. perspectiva actual. Açoreana, 10(1): CORNELIUS, P. F., 1992. The Azores 11-140. hydroid fauna and its origin, with FRIEDLANDER, I., 1929. Die Azoren. discussion of rafting and medusa Zeitschrift für Vulkanologie, 12(2-3): suppression. Arquipélago, Life and 77-107. Earth Sciences, 10: 75-99. FRUTUOSO, G., 1983. Livro Terceiro 72 AÇOREANA 2007, Supl. 5: 59-73

das Saudades da Terra. Instituto MITCHELL-THOMÉ, R.C., 1976. Cultural de Ponta Delgada, Geology of the middle Atlantic Islands, clxxviii+300pp., Ponta Delgada. 351pp. Gebrüder Borntraeger, GARCÍA-TALAVERA, F., 1990. Berlin. Fauna tropical en el Neotirrenien- MITCHELL-THOMÉ, R.C., 1981. se de Santa Maria (I. Azores). Structural features of the sedimen- Lavori S. I. M., 23: 439-443. tary rocks of . Ar- HARTUNG, G., 1860. Die Azoren in quipélago, 2: 121-151. ihrer äusseren Erscheinung und nach MORELET, A., 1860. Notice sur ihrer geognostischen Natur, 350pp. l’Histoire Naturelle des Açores suivie Verlag von Wilhelm Engelman. d’une description des Mollusques ter- HARTUNG, G., 1864. Geologische Besch- restres de cet Archipel, 216pp., 5pls. reibung der Inseln Madeira und Porto J.-B. Baillière et Fils, Paris. Santo, 299 pp.Engelman Verlag. PRUD’HOMME VAN REINE, W. F., KREJCI-GRAF, K., J. FRECHEN, W. 1988. Phytogeography of seaweeds WETZEL & G. COLOM, 1958. of the Azores. Helgolander Meere- Gesteine und Fossilien von den suntersuchung, 42: 165-185. Azoren. Senckenbergiana Lethaea, REISS, W., 1862. Mitteiling über die 39: 303-351. tertiären Schichten von Santa LOPES, M. F. R., J. C. MARQUES & Maria, der südlichsten der Azoren, D. BELLAN-SANTINI, 1993. The und ihre organischen Einschlüsse. benthic amphipod fauna of the Neues Jarhrbuch für Mineralogie, Azores (Portugal): an up-to-date Geognosie, Geologie und Petrefacten- annotated list of species, and some kunde: 1-22. biogeographic considerations. SANTOS, R. S., S. HAWKINS, L. R. Crustaceana, 65: 204-217. MONTEIRO, M. ALVES & E. J. MADEIRA, J. E. O., 1986. Geologia ISIDRO, 1995. Marine research, re- Estrutural e Enquadramento Geo- sources and conservation in the tectónico da Ilha de Santa Maria Azores. Aquatic Conservation: Marine (Açores), 107 pp. MSc. Thesis, and Freshwater Ecosystems, 5: 311-354. Departamento de Geologia, SANTOS, R. S., F. M. PORTEIRO & J. Faculdade de Ciências da Uni- P. BARREIROS, 1997. Marine versidade de Lisboa, Lisboa. Fishes of the Azores: An annotated MAYER, K., 1864. Die Tertiär-Fauna checklist and bibliography. Arqui- der Azoren und Madeiren. Systema- pélago. Life and Marine Sciences. tisches Verzeichniss der fossilen Reste Supplement 1: xxviii + 244 pp. von Madeira, Porto Santo und Santa SERRALHEIRO, A., 2003. A Geologia Maria nebst Beschreibung der neuen da Ilha de Santa Maria, Açores. Arten, vi+107 pp., Zürich. Açoreana, 10(1): 141-192. MITCHELL-THOMÉ, R.C., 1974. The SERRALHEIRO, A., C. M. ALVES, Sedimentary Rocks of Macaro- V.H. FORJAZ & B. RODRIGUES, nesia. International Journal of Earth 1987. Carta Vulcanologica dos Sciences, 63: 1179-1216. Açores, Ilha de Santa Maria MADEIRA ET AL: HISTORICAL OVERVIEW 73

esc.1/15.000, 2 folhas. Centro Irish Biogeographical Society, 9: Vulcanologia INIC, Ponta Delgada. 40-54. SERRALHEIRO, A., & J. MADEIRA, WIRTZ, P. & H. R. MARTINS, 1993. 1990. Stratigraphy and geochrono- Notes on some rare and little logy of Santa Maria Island known marine invertebrates from (Azores). Livro de Homenagem ao the Azores, with a discussion of the Prof. Carlos Romariz, pp. 357-376. zoogeography of the region. Departamento Geologia da Facul- Arquipélago. Life and Marine dade de Ciências da Universidade Sciences, 11A: 55-63. de Lisboa, Lisboa. ZBYSZEWSKI, G. & F. M. ALMEIDA, STORETVEDT, K. M., A. 1950. Os peixes miocénicos por- SERRALHEIRO, M. MOREIRA & tugueses. Comunicações dos Serviços M. C. ABRANCHES, 1989. Mag- Geológicos de Portugal, 31: 309-412. netic structure and evolution of ZBYSZEWSKI, G., O. V. FERREIRA & the island of Santa Maria, Azores. C. F. T. ASSUNÇÃO, 1961. Carta Physics of the Earth and Planetary Geológica de Portugal na escale 1: Interiors, 58: 228-238. 50,000. Noticia explicativa da folha de TEIXEIRA, C., 1950. Notas sobre a Ilha de Santa Maria (Açores). geologia das ilhas Atlânticas. Serviços Geológicos de Portugal, Anais da Faculdade de Ciências da 28 pp., Lisboa. Universidade do Porto, 33: 209-215. ZBYSZEWSKI, G. & O. V. FERREIRA, TITTLEY, I. & A. I. NETO, 1995. The 1961. La faune marine des basses marine algal flora of the Azores plages quaternaires de Praia et de and its biogeographical affinities. Prainha dans l’île de Santa Maria Boletim do Museu Municipal do (Açores). Comunicações dos Serviços , Suplemento 4: 747-766. Geológicos de Portugal, 45: 467-478. TITTLEY, I. & A. I. NETO, 2006. The ZBYSZEWSKI, G. & O. V. FERREIRA, marine algal flora of the Azores: 1962a. Étude géologique de l’île de island isolation or Atlantic step- Santa Maria (Açores). Comunicações ping-stones? In: HAYDEN, T.J., dos Serviços Geológicos de Portugal, D.A. MURRAY & J. P. 46: 209-245. O’CONNOR (Eds). Proceedings of Zbyszewski, G. & O. V. Ferreira, the 5th international symposium 1962b. La faune miocène de l’île de on the fauna and flora of Atlantic Santa Maria (Açores). Comunicações Islands. Dublin, 24-27 August dos Serviços Geológicos de Portugal, 2004. Occasional Publication of the 46: 247-289.