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AÇOREANA 

Revista de Estudos Açoreanos

AÇOREANA 

Revista de Estudos Açoreanos 

PROPRIEDADE da 

Sociedade Afonso Chaves 

Associação de Estudos Açorianos 

Sede: Edifício do Museu “Carlos Machado” 

Apartado 258 – 9501-903 Ponta Delgada 

São Miguel, Açores, Portugal 

PRESIDENTE e EDITOR 

António M. de Frias Martins 

QUADRO EDITORIAL 

Brian Morton 

Scientific Associate 

Department of Zoology 

The Natural History Museum 

Cromwell Road 

London SW7 5BD, UK 

António Serralheiro 

Departamento de Geologia 

Universidade de Lisboa, Portugal 

Paulo A.V. Borges 

Departamento de Ciências Agrárias 

Universidade dos Açores, Portugal 

PAGINAÇÃO 

CLASSICOR, LDA. 

IMPRESSÃO E ACABAMENTO 

EGA - Empresa Gráfica Açoreana, Lda 

Edição subsidiada pela 

Direcção Regional da Ciência, 

Tecnologia e Comunicações 

do Governo Regional dos Açores 

Setembro de 2009 

TIRAGEM 

750 exemplares 

DEPÓSITO LEGAL 

113 234 / 98 

ISSN 

0874 - 0380 

Esta edição é totalmente impressa (à excepção da capa) 

em papel ecológico, sem cloro, ácidos ou branqueamentos ópticos

AÇOREANA 

Revista de Estudos Açoreanos 

SUPLEMENTO 6 SETEMBRO 2009 

THE MARINE FAUNA AND FLORA OF THE AZORES 

Proceedings of the 

Third International Workshop of Malacology 

and Marine Biology 

Vila Franca do Campo, São Miguel, Açores 

July 17-28, 2006 

Sponsored by 

Edited by 

Sociedade Afonso Chaves 

Departamento de Biologia 

Universidade dos Açores 

Ponta Delgada

C O N T E Ú D O / C O N T E N T S 

7 LIST OF PARTICIPANTS 

9 THE AZOREAN WORKSHOPS 


15 ILLUSTRATED CHECKLIST OF THE INFRALITTORAL MOLLUSCS OFF 

VILA FRANCA DO CAMPO 

António M. de Frias Martins, José Pedro Borges, Sérgio Ávila, Ana C. Costa, 

Patrícia Madeira & Brian Morton 

105 ASPECTS OF THE BIOLOGY AND FUNCTIONAL MORPHOLOGY OF 

TIMOCLEA OVATA (BIVALVIA: VENEROIDEA: VENERINAE) IN THE 

AÇORES, PORTUGAL, AND A COMPARISON WITH CHIONE ELEVATA 

(CHIONINAE) 

Brian Morton 

121 ANATOMY AND BIOLOGY OF MITRA CORNEA LAMARCK, 1811 (MOL- 

LUSCA, CAENOGASTROPODA, MITRIDAE) FROM THE AZORES 

M. G. Harasewych 

137 COMPARATIVE STUDY OF CHEMICAL DEFENCES FROM TWO 

ALLOPATRIC NORTH ATLANTIC SUBSPECIES OF HYPSELODORIS 

PICTA (MOLLUSCA: OPISTHOBRANCHIA) 

Helena Gaspar, Ana Isabel Rodrigues & Gonçalo Calado 

145 THE BIOLOGY OF THE ZONING SUBTIDAL POLYCHAETE DITRUPA 

ARIETINA (SERPULIDAE) IN THE AÇORES, PORTUGAL, WITH A 

DESCRIPTION OF THE LIFE HISTORY OF ITS TUBE 

Brian Morton & Andreia Salvador 

157 DRILLING PREDATION UPON DITRUPA ARIETINA (POLYCHAETA: 

SERPULIDAE) FROM THE MID-ATLANTIC AÇORES, PORTUGAL 

Brian Morton & E.M. Harper 

167 THE PYCNOGONIDS (ARTHROPODA: PYCNOGONIDA) OF SÃO 

MIGUEL, AZORES, WITH DESCRIPTION OF A NEW SPECIES OF 

ANOPLODACTYLUS WILSON, 1878 (PHOXICHILIDIIDAE) 

Roger N. Bamber & Ana Cristina Costa 

183 THE TANAIDACEANS (ARTHROPODA: PERACARIDA: TANAIDACEA) 

OF SÃO MIGUEL, AZORES, WITH DESCRIPTION OF TWO NEW 

SPECIES, AND A NEW RECORD FROM TENERIFE 

Roger N. Bamber & Ana Cristina Costa 

201 THE SOFT-SEDIMENT INFAUNA OFF SÃO MIGUEL, AZORES, AND A 

COMPARISON WITH OTHER AZOREAN INVERTEBRATE HABITATS 

Roger N. Bamber & Roni Robbins 

211 SHELL OCCUPANCY BY THE HERMIT CRAB CLIBANARIUS ERYTHRO- 

PUS (CRUSTACEA) ON THE SOUTH COAST OF SÃO MIGUEL, AÇORES 

Pedro Rodrigues & Roshan K. Rodrigo 

217 A CONSERVATIONAL APPROACH ON THE SEABIRD POPULATIONS 

OF ILHÉU DE VILA FRANCA DO CAMPO, AZORES, PORTUGAL 

Pedro Rodrigues, Joana Micael, Roshan K. Rodrigo & Regina T. Cunha 

227 REMEMBERING Joseph C. Britton (1942-2006)

1 

2 

3 

4 

5 

6 8 10 

12 

11 

7 

14 15 

9 13 16 

Third International Workshop on Malacology and Marine Biology. 1, António M. de Frias Martins; 

2, Brian Morton; 3, José Pedro Borges; 4, Ana Cristina Costa; 5, Roshan Rodrigo; 6, Jerry 

Harasewych; 7, Vera Malhão; 8, Sérgio Ávila; 9, Joana Xavier; 10, Roni Robbins; 11, Paola Rachello; 

12, Roger Bamber; 13, Patrícia Madeira; 14, Andreia Salvador; 15, Pedro Rodrigues; 16, Daniela 

Gabriel. 

LIST OF PARTICIPANTS 

Ana Cristina Costa - Departamento de Biologia, Universidade dos Açores, 9501-801 

Ponta Delgada, São Miguel, Açores, Portugal - accosta@uac.pt 

Andrea Cunha - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal -andrea_cunha@linus.uac.pt 

Andreia Salvador - Environment: Coastal & Marine, The Natural History Museum, 

Cromwell Road, London SW7 5BD, United Kingdom - a.salvador@nhm.ac.uk 

António M. de Frias Martins - Departamento de Biologia, Universidade dos Açores, 

9501-801 Ponta Delgada, São Miguel, Açores, Portugal - frias@uac.pt 

Brian Morton - Scientific Associate, Department of Zoology, The Natural History 

Museum, Cromwell Road, London SW7 5BD, U.K. - prof_bsmorton@hotmail.com 

Daniela Gabriel - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - danielagferreira@hotmail.com; 

dgabriel@uac.pt

Gonçalo Calado - IPM-Instituto Português de Malacologia, Zoomarine EN 125 KM 65, 

8200-864 Guia, Portugal - bagoncas@mail.telepac.pt 

Jerry Harasewych - Curator of Marine Mollusca, Smithsonian Institution, National Museum 

of Natural History, Department of Invertebrate Zoology, P.O. Box 37012 MRC 163, 

Washington DC 20013-7012, USA - HarasewychJ@si.edu 

Joana Micael - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - joanamicael@cimar.org; jomicap@hotmail.com 

Joana Xavier - Institute for Biodiversity and Ecosystem Dynamics (IBED), University of 

Amsterdam, Faculty of Science, Mauritskade 57, 1092 AD Amsterdam, Netherlands - 

xavier@science.uva.nl 

José Pedro Borges - IPM-Instituto Português de Malacologia, Zoomarine EN 125 KM 65, 

8200-864 Guia, Portugal - josepedroborges@sapo.pt 

Kathe Jensen - Reasearch Associate, Zoological Museum, Copenhagen; Sjælør Boulevard 49, 

2.tv., 2450 København SV, Denmark - KRJensen@zmuc.ku.dk 

Manuela Parente - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - mparente@uac.pt 

Maria Ana Dionísio - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - anamdionisio@gmail.com 

PaolaG. Rachello Dolmen - Institute for Biodiversity and Ecosystem Dynamics (IBED), 

University of Amsterdam, Faculty of Science, Mauritskade 57, 1092 AD Amsterdam, 

Netherlands - P.Rachello@student.uva.nl 

Patrícia Madeira - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - tamissa@hotmail.com 

Pedro Rodrigues - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - pedroreisrodrigues@yahoo.com 

Roger Bamber - Consultancy Leader, Environment: Coastal & Marine, The Natural History 

Museum, Cromwell Road, London SW7 5BD, United Kingdom - r.bamber@nhm.ac.uk 

Roni Robbins - Environment: Coastal & Marine, The Natural History Museum, Cromwell 

Road, London SW7 5BD, United Kingdom - r.robbins@nhm.ac.uk 

Roshan K. Rodrigo - No.47, Saman Mawatha, Gangula, Panadura, Sri Lanka - 

rodrigoroshan@yahoo.com 

Sérgio Ávila - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - avila@notes.uac.pt 

Vera Malhão - Departamento de Biologia, Universidade dos Açores, 9501-801 Ponta 

Delgada, São Miguel, Açores, Portugal - vmalhao@notes.uac.pt; 

veramalhao@hotmail.com

AÇOREANA, Suplemento 6, Setembro 2009: 9-13 

THE AZORES WORKSHOPS 


Sociedade Afonso Chaves, 9501-903 Ponta Delgada, São Miguel, Açores, Portugal 

e-mail: afonsochaves.sac@gmail.com 

Twenty-one years ago, as a joint initiative 

of Sociedade Afonso Chaves and 

the Department of Biology of the 

University of the Azores, and under the 

sponsorship of the local Municipality, 

about half a dozen foreign scientists gathered 

in Vila Franca do Campo and, setting 

up an improvised laboratory in the dark 

and humid facilities of the local newspaper 

“A Crença”, made history with the 

realization of the 1 st International 

Workshop of Malacology. The workshop 

participants came from the most prestigious 

scientific institutions: the 

Universities of Hong Kong, Harvard, 

Rhode Island, Liverpool, from the 

Smithsonian Institution, the Muséum 

national d’Histoire naturelle, Paris and 

the California Academy of Sciences. With 

another half dozen local scientists and 

academics, and aspiring young students, 

science was made right there, next to one 

of Vila Franca’s most significant monuments 

to culture and education of: the 

“Externato”. So worried with the (primitive) 

logistics were the organizers that 

they did not think of registering the event 

photographically for posterity. 

Nevertheless, in 1990 the proceedings of 

this first workshop came to light and 

remain today a reference document for 

the scientific study of the Ilhéu de Vila 

Franca do Campo and of the malacological 

fauna of the Azorean littoral. 

The research workshops, as regular 

events, were popularized in the scientific 

community by Brian Morton when 

Professor of Marine Ecology at the 

University of Hong Kong and Director of 

the Swire Institute of Marine Science. 

Through them, every three years, Brian 

rendered available for invited specialists 

in various scientific fields two weeks of 

research, with but one requisite in return: 

to publish in the proceedings of the event 

at least one research paper on the marine 

biology of Hong Kong. Brian Morton had 

visited the Azores in 1965, as a Chelsea 

College graduate, and since then has 

developed a special appreciation for these 

islands. It was then that I had the opportunity 

of meeting him and, from that time 

on, a relationship based on friendship 

and mutual respect developed that time 

has only made stronger. 

The 2 nd workshop, in 1991, and this 

time in the facilities of the fish market in 

Vila Franca do Campo, attracted about 40 

participants, national and foreign, with 

the corresponding scientific production 

broadened to other endeavours of marine 

biology besides malacology. The spectrum 

of participants was equally widened 

to include Australia and Hong Kong, various 

American institutions (University of 

Rhode Island, Smithsonian Institution,

10 AÇOREANA 

2009, Sup. 6: 9-13 

Harbor Branch Oceanographic Institution, 

Field Museum of Natural History, 

Chicago, Texas Christian University, Long 

Island University) and a fair representation 

from Europe (Odense University, 

Zoological Museum, Copenhagen, both 

in Denmark, Antwerp University/Institut 

Royal des Sciences naturelles de Belgique; 

University Marine Biological Station, 

Millport, UK). The workshop added the 

novelty of a panel on Marine 

Conservation, sponsored by UNESCO’s 

National Committee. The proceedings of 

the workshop were published in 1995. 

Ecology of the Açores had been undertaken 

by the Sociedade Afonso Chaves to 

replace a workshop, so that a systematic 

and all-encompassing study of the littoral 

would be conducted and reported upon 

comprehensively for the benefit, especially 

of local students. On the other hand, 

the famous expeditions of Albert the 1 st , 

Prince of Monaco, had unravelled some 

of the biological secrets of the great 

depths around the Azores. To know better 

that band between 50 and 200 metres, 

well known to the fishermen for its abundant 

fish life but practically absent from 

scientific publications due to lack of 

research directed at it, was then an obvious 

choice for a workshop project. 

This practical objective – to better 

know Azorean marine biodiversity – was 

linked to another, more theoretical goal, 

resulting from studies of the distribution 

of species through time. Knowing that 

After a long interval, marked by other 

types of events – for example, the production 

of the book Coastal Ecology of the 

Açores, in 1998 –, the Sociedade Afonso 

Chaves and the Department of Biology of 

the University of the Azores united in 

their efforts once again to promote, in the 

eternal capital of the island of São Miguel, 

from July 17 to 28 of 2006, the 3 rd 

International Workshop of Malacology 

and Marine Biology, under the auspices of 

the Municipality of Vila Franca do Campo 

and with the much-appreciated collaboration 

of the Clube Naval de Vila Franca 

do Campo. 

Although open to every aspect of science 

of interest to the participants, the 3 rd 

workshop had a specific objective: to 

research virtually unstudied grounds. In 

fact, the Azorean coastal zone has been 

profusely studied and the book Coastal 

cyclical variations in temperature resulting 

from the glaciations lead to the disappearance 

of species less adapted to such 

fluctuations, could we find species of previous 

colder ages that have sought refuge 

deeper, where temperatures are lower? 

The solution to this double quest rested 

on a dredging plan down to 200 metres. 

In fact, as it transpired, samples of the sea 

bed off Vila Franca do Campo were taken 

down to 360 metres during the workshop.

MARTINS: THE AZORES WORKSHOPS 11 

The collected material has been deposited 

in the Department of Biology reference 

collection and remains available to everyone 

who desires to conduct further 

research on it. 

Honouring the core interest of the 

workshop – Malacology – special attention 

was paid to molluscs and a listing of 

those species collected is provided. 

However, many other inhabitants of the 

sea bed came up in the dredge. 

The larger fractions of such samples were 

sorted immediately and the finer fractions 

preserved for sorting later, in the 

laboratory. 

As in previous meetings, the proceedings 

of the 3 rd workshop are published. 

The papers included in the proceedings 

result from short projects that the participants 

had taken upon themselves to 

develop, on the basis of any particular scientific 

aspect judged appropriate for the 

short time available for gathering data.

12 AÇOREANA 

2009, Sup. 6: 9-13 

Whenever possible, a photographic 

record of each living animal was 

obtained. The wealth of species collected 

is a clear sign that this workshop needs to 

be continued. 

The scientific content of the workshop 

is the most visible, desirable, outcome of 

an effort that had its roots in the statutes 

of the oldest scientific society in the 

Azores - the Sociedade Afonso Chaves – 

and which are to promote the realization 

of scientific meetings dealing with the 

natural history of the Azores. As in previous 

events, the participation of the 

Department of Biology of the University of 

the Azores provided the scientific support. 

The warm hospitality of the Mayor of Vila 

Franca do Campo, Rui Melo, and the enthusiastic 

collaboration of the Clube Naval and 

its president, Paulo Melo, paved the way to 

comfortable and efficient logistics. The skill 

of Moisés Bolarinho, the skipper of the "Vila 

Franca" and the dedication of is crew took 

us to the best sites for successful dredging. 

The indispensable financing for this 3 rd 

workshop came from the DRCT – the 

regional governmental organism responsible 

for Science and Technology. The Luso- 

American Foun-dation for Development 

(FLAD), the Foundation for Science and 

Technology (FCT) and the Municipality of 

Vila Franca do Campo were also generous 

in this area.

MARTINS: THE AZORES WORKSHOPS 13 

The choice of Vila Franca do Campo 

for the base camp of each workshop is 

somehow connected with the beautiful 

islet which is the ex-libris of this noble old 

capital of the island of São Miguel. 

Science is obviously the primary objective 

of these meetings. However, besides the 

knowledge acquired due to the research 

therein developed, there is a clear interest 

in fostering culture and education within 

the community. We are sure that the better 

knowledge about the Ilhéu de Vila 

Franca do Campo, in particular, resulting 

from the workshops has contributed to 

the strengthening of its stature as a nature 

reserve, cared for and respected by the 

thousands of users that visit it every summer. 

The prospect of an aquarium in 

Vila Franca do Campo has turned the 

attention of the local community towards 

one of its most precious resources - the 

sea. 

Finally, and, as a silent highlight of the 

workshop meetings, past, present and 

future, the generous offer of the founder 

of the workshop concept and of the Swire 

Institute of Marine Science, Professor 

Brian Morton, to help in the setting up of 

a Marine Biology Station in the facilities 

where this 3 rd workshop took place, will 

continue to bring to Vila Franca do 

Campo, a nucleus of resident researchers 

and academic guests from around the 

world who will undoubtedly act as a catalyst 

for science and education in the 

community which has so generously 

hosted these pioneering adventures. 

It is the belief of this author, his local 

colleagues and the groups of overseas scientists 

who have contributed, over the 

years, to the greater understanding of the 

unique marine life of the Azores in general 

and of the waters of Vila Franca do 

Campo in particular, that there is no better 

place where a permanent, international, 

marine research facility would not just 

flourish but would act as the archipelago’s 

flagship promoting marine conservation 

and technological advancement. 

For the benefit of all.


ILLUSTRATED CHECKLIST OF THE INFRALITTORAL MOLLUSCS 

OFF VILA FRANCA DO CAMPO 

António M. de Frias Martins 1 , José Pedro Borges 2 , Sérgio P. Ávila 3,4 , Ana C. Costa 1 , 

Patrícia Madeira 3 & Brian Morton 5 

1 

CIBIO-Pólo Açores, Department of Biology, University of the Azores, 9501-801 Ponta Delgada, São Miguel, 

Azores, Portugal. e-mail: frias@uac.pt 

2 

IPM-Instituto Português de Malacologia, Zoomarine EN 125 KM 65, 8200-864 GUIA, Portugal 

3 

Marine PalaeoBiogeography group, Department of Biology, University of the Azores, 9501-801 Ponta Delgada, 

São Miguel, Azores, Portugal 

4 

Centro do IMAR da Universidade dos Açores, 9901-862 Horta, Azores, Portugal 

5 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. 

ABSTRACT 

A list of the molluscan species dredged during the 3 rd International Workshop of 

Malacology and Marine Biology is presented. Positive identification has not been possible 

for a number of taxa. However, almost all species are illustrated so as to provide a practical 

guide to the species which may occur as beach drift and others which naturally range 

up into the intertidal. 

The distribution of the species at the various collecting stations is also presented as a 

table organized by depth, and identifying where specimens were collected alive or, in the 

case of bivalves, with both valves attached. 

RESUMO 

Apresenta-se uma listagem das espécies de moluscos dragadas durante o 3º Workshop 

Internacional de Malacologia e Biologia Marinha. Uma identificação positiva não foi 

possível para um determinado número de taxa. No entanto, quase todas as espécies estão 

ilustradas, de modo a providenciar um guia prático para as que podem ocorrer no material 

arrojado nas praias e outras que naturalmente estendem a sua distribuição até ao 

intertidal. 

A distribuição das espécies pelas várias estações de colheita é também apresentada 

numa tabela organizada por profundidade, identificando também onde os exemplares 

foram recolhidos vivos ou, no caso dos bivalves, com ambas as valvas ligadas. 

INTRODUCTION 

The main purpose behind the 3 rd 


and Marine Biology was to test the hypothesis 

that the deeper, colder, waters around 

the Azorean islands acted as a refuge during 

the warm periods that interspersed 

glaciations. It is thought that when surface 

temperatures began to rise, cold-water 

species would likely follow their temperature 

optima by descending to appropriate 

depths. Present results, preliminary as 

they are, do not support such a suggestion. 

The extensive dredging has produced a 

wealth of information about the molluscan 

biodiversity of the sublittoral off Vila 

Franca do Campo, in a depth range poorly 

studied by previous authors in the Azores. 

This herein published list of 232 molluscan 

species and the pictorial representations of 

most of them is a contribution to that 

knowledge. The illustrations, whenever 

possible, were not only of the collected 

shells but of the living animals. This will 

provide students and researchers with an 

easier identification guide to facilitate 

their studies. Also, although in many

16 AÇOREANA 

2009, Sup. 6: 15-103 

cases positive identifications were not 

possible, the provided illustrations supply 

a basis for further studies leading to 

more accurate taxonomic determinations. 

The micromolluscs are unevenly 

reported upon because the minimum 

dredge mesh was 2.5 mm and only some 

hauls were sieved to retain the residual 

1mm fraction. 

A list of the stations related to the 

workshop is provided (see Text figure 1). 

All species sorted are reported upon 

(Table 1), even though their shells could 

only have been transported from other 

habitats, including the intertidal. Such 

situations are commented upon for each 

species identified. To facilitate inferences 

related to depth distribution, the species 

collected alive or, in the case of bivalves, 

with both valves attached, are shown in 

bold in the Taxonomic List, and the sampling 

depths also indicated. Also, in 

Table 1 the stations are arranged by 

increasing depth and the state of the collected 

specimens (fragment, empty 

shell/one valve, with animal/two valves) 

is differentiated by variations in the density 

of their gray overlays, so as to give an 

overall pictorial view of the distribution 

of each taxon. 

The depth ranges of the species given 

by Poppe & Gotto (1991-1993) (P&G) and 

Macedo et al. (1999) (MM&B) are referred 

whenever possible. Also, where appropriate, 

reference is made to the species 

found alive in the nearby Ilhéu de Vila 

Franca do Campo (IVFC) (Martins, 2004). 

This is intended as a clarification to the 

distribution of these species in the 

dredged material. 

Previous works on Ilhéu de Vila 

Franca do Campo have listed the marine 

molluscs of that islet or of the shores nearby, 

and Ávila et al. (2000) have provided 

the most recent list of the shallow-water 

marine molluscs of São Miguel. For comparison 

purposes reference to these publications 

will be made under the appropriate 

taxa. Species are presented following 

their current systematic position as set 

out in CLEMAM (Check List of European 

Marine Molluscs) (http://www.somali.asso.fr/ 

clemam/index.clemam.html). 

LIST OF SAMPLING STATIONS 

Forty-six stations were sampled during 

the workshop (Text figure 1). Stations 3-4, 

9-11, 17 were dive sites and are not included 

in this report, except in the notes provided 

under the respective station code. 

Stations 47-54 were sampled after the 

workshop timeframe but followed the 

same procedure and are, therefore included 

herein. Similarly, stations 56-58, 

sampled during fieldwork undertaken for 

the Malacology class of the Department of 

Biology of the University of the Azores, 

were added to the workshop material. 

Station 55 represented from the biological 

material collected from a stone snagged 

by a fishing net. 

STATION 1 – in front of the marina – Vila 

Franca do Campo. 

Date: 17-07-2006. 

Depth: 32 fathoms (57 m). 

Co-ordinates: N 37° 42’ 12” W 25° 25’ 09”. 

Collected by: Frias Martins, Brian Morton, Jerry 

Harasewych. 

Observations: (large dredge) two tows (A and B) 

pooled. 

STATION 2 – in front of the marina - Vila 


Date: 17-07-2006. 



Collected by: Frias Martins, Brian Morton, Jerry 

Harasewych. 

Observations: (large dredge). 

STATION 3 – around Farilhão, SE of Ilhéu de 

Vila Franca do Campo. 

Date: 18-07-2006. 

Depth: 5-19 m.

MARTINS ET AL: INFRALITORAL MOLLUSCS OFF VILA FRANCA DO CAMPO 17 

TEXT FIGURE 1. Distribution of the stations. 

Co-ordinates: —- 

Collected by: Joana Xavier, Paola Rachello, José 

Pedro Borges, Gonçalo Calado. 

Observations: (SCUBA) 1 ophiurid (10 m) 

Hypselodoris picta webbi (4 spcs.), Chromodoris 

purpurea (2 spcs.), Platydoris argo

18 AÇOREANA 

2009, Sup. 6: 15-103 

(1 spc.), Flabellina (1 spc.), Hypselodoris 

midatlantica (1 spc.); Umbraculum 

umbraculum (1 spc. feeding on Halichondria 

aurantiaca). 

STATION 4 – off Cais do Tagarete, Vila Franca 

do Campo. 

Date: 18-07-2006. 

Depth: 7 m. 


Collected by: Gonçalo Calado, José Pedro 

Borges, Paola Rachello. 

Observations: (SCUBA). 

STATION 5 

Date: 19-07-2006. 

Depth: 23-24 fathoms (41-43 m) 


Collected by: Brian Morton, Sérgio Ávila, Pedro 

Rodrigues. 

Observations: (dredge) 6 minute tow at 1.5 

knotts. 

STATION 6 

Date: 19-07-2006. 

Depth: 22-23 fathoms (40-41 m). 



Rodrigues. 

Observations: (dredge) “earthworm” sand fish 

Apterichthus caecus (Linnaeus, 1758). 

STATION 7 

Date: 19-07-2006. 




Rodrigues. 

Observations: (dredge) 10 min tow at 1.5 – 2 

knotts. 

STATION 8 

Date: 19-07-2006. 




Rodrigues. 

Observations: (dredge) 6 minute tow at 1.5 – 2 

knotts. 

STATION 9 – Portinho da Ribeirinha. 

Date: 20-07-2006. 

Depth: 5-14 m. 


Collected by: Gonçalo Calado, Joana Xavier, 

Patrícia Madeira, Paola Rachello. 

Observations: (SCUBA) ophiurids. 

STATION 10 – wall and mouth of the marina – 

Vila Franca do Campo. 

Date: 20-07-2006. 

Depth: 6 m. 


Collected by: Gonçalo Calado, Patrícia Madeira. 

Observations: (SCUBA) echinoderms. 

STATION 11 – Ilhéu de Vila Franca do Campo 

(NE). 

Date: 19-07-2006. 

Depth: 16 m. 


Collected by: Gonçalo Calado, José Pedro 

Borges, Joana Xavier, Paola Rachello, 

Patrícia Madeira. 

Observations: (SCUBA). 

STATION 12 

Date: 21-07-2006. 



Collected by: Brian Morton. 

Observations: (dredge). 

STATION 13 

Date: 21-07-2006. 

Depth: 47.9–40.8 fathoms (86-73 m). 




STATION 14 

Date: 21-07-2006. 

Depth: 25-26.2 fathoms (45-47 m). 




STATION 15 

Date: 21-07-2006. 

Depth: 25,7 – 26 fathoms (46-47 m). 



Observations: (dredge).


STATION 16 

Date: 21-07-2006. 

Depth: 9.9 – 11 fathoms (18-20 m). 




STATION 17 – mouth of Ilhéu de Vila Franca 

do Campo. 

Date: 20-07-2006. 

Depth: 6 m. 


Collected by: Andrea Cunha, Daniela Gabriel. 

Observations: (SCUBA) sponges, sand urchin, 

algae. 

STATION 18 – off Ribeira das Tainhas. 

Date: 21-07-2006. 



Collected by: Frias Martins, Brian Morton, Roger 

Bamber. 

Observations: (small dredge) 5 minute tow. 


Date: 21-07-2006. 



Collected by: Roger Bamber, Frias Martins, Brian 

Morton. 

Observations: (grab). 


Date: 21-07-2006. 




Morton. 



Date: 21-07-2006. 




Morton. 



Date: 24-07-2006. 



Collected by: Frias Martins, Brian Morton. 



Date: 24-07-2006. 

Depth: 65-25 fathoms (117- 45 m). 



Observations: (large dredge) 


Date: 24-07-2006. 


Co-ordinates: 37° 42’ 04” W 25° 25’ 02”. 



STATION 25 – off Praia da Vinha da Areia. 

Date: 24-07-2006. 

Depth: 7.6 fathoms (14 m). 





Date: 21-07-2006. 




Morton. 



Date: 24-07-2006. 



Collected by: Sérgio Ávila. 

Observations: (large dredge) Holothuria. 

STATION 28– off Ribeira das Tainhas. 

Date: 24-07-2006. 






Date: 24-07-2006. 




Observations: (large dredge).

20 AÇOREANA 

2009, Sup. 6: 15-103 


Date: 24-07-2006. 





STATION 31 – off Cais do Tagarete, Vila 


Date: 25-07-2006. 



Collected by: Frias Martins. 


STATION 32 – off Cais do Tagarete, Vila 


Date: 25-07-2006. 





STATION 33 – off Rosto Branco, Água d’Alto. 

Date: 25-07-2006. 


Co-ordinates: N 37° 41” 08” W 25° 27’ 18”. 



STATION 34 

Date: 25-07-2006. 



Collected by: Roger Bamber, Joana Xavier, Paola 

Rachello. 

Observations: (grab) two samples pooled. 

STATION 35 

Date: 25-07-2006. 




Rachello. 


STATION 36 

Date: 25-07-2006. 




Rachello. 


STATION 37 

Date: 25-07-2006. 

Depth: 130-75 fathoms (234-135 m) (G); 117-65 

fathoms (210-117 m) (H). 

Co-ordinates: N 37° 42’ 13” W 25° 24’ 36” (G) 

N 37° 41’ 59” W 25° 24’ 44” to N 37° 42’ 11” 

W 25° 24’ 45” (H). 

Collected by: Joana Xavier, Paola Rachello, 

Roger Bamber. 

Observations: (large dredge) G+H were pooled. 

STATION 38 

Date: 25-07-2006. 


Co-ordinates: N 37° 41’ 41” W 25° 25’ 06” to N 

37° 41’ 17” W 25° 25’ 10”. 

Collected by: Joana Xavier, Paola Rachello, 

Roger Bamber. 


STATION 39 

Date: 25-07-2006. 




Rachello. 


STATION 40 – off Amora, Ponta Garça. 

Date: 26-07-2006. 



Collected by: Frias Martins, Jerry Harasewych. 

Observations: (small dredge). 


Date: 26-07-2006. 




Observations: (large dredge); bottom drops suddenly 

to about 600 fathoms; tow up slope. 

STATION 42 – off Praia de Água d’Alto. 

Date: 26-07-2006. 




Observations: (large dredge) rocky; octopus. 


Date: 26-07-2006.





Observations: (large dredge) rock. 


Date: 26-07-2006. 





STATION 45 – off Vinha da Areia, Vila Franca 

do Campo. 

Date: 26-07-2006. 




Observations: (small dredge) near sewage outlet. 


do Campo. 

Date: 26-07-2006. 






do Campo. 

Date: 05-09-2006. 



Collected by: Frias Martins, Patrícia Madeira, 

Henk van Goor. 



do Campo. 

Date: 05-09-2006. 







do Campo. 

Date: 05-09-2006. 







do Campo. 

Date: 05-09-2006. 







Date: 05-09-2006. 





Observations: (large dredge) sponge, corals. 


Date: 05-09-2006. 






STATION 53 – off Ponta Garça. 

Date: 05-09-2006. 







do Campo. 

Date: 05-09-2006. 






STATION 55 – off Vila Franca do Campo. 

Date: 07-09-2006. 


Co-ordinates: — 

Collected by: Moisés Bolarinho, skipper of the 

fishing boat ‘Vila Franca’.

22 AÇOREANA 

2009, Sup. 6: 15-103 

Observations: rock snagged on fishing gear; 

covered with mud, two white gorgonians, 

some sponges and living Chama. 

STATION 56 – off Vila Franca do Campo, east 

of Ilhéu. 

Date: 03-10-2006. 



Collected by: Malacology class (Department of 

Biology, University of Azores). 

Observations: (small dredge) one tow 5 min 

and another 15 min, pooled. 

STATION 57 – off Vila Franca do Campo, 

west of Ilhéu. 

Date: 03-10-2006. 





Observations: (small dredge) tow during 15 

min. 

STATION 58 – off Vila Franca do Campo, 

west of Ilhéu. 

Date: 03-10-2006. 





Observations: (grab) 4 samples, pooled. 

TAXONOMIC LIST 

Phylum MOLLUSCA 

Class POLYPLACOPHORA Gray, 1821 

Order LEPIDOPLEURIDA Thiele, 1909 

Family Leptochitonidae Dall, 1889 

Lepidochiton cimicoides (Monterosato, 1879) 

(Figure 1) 

Remarks: Specimens collected only at Station 

37; alive. Depth range: 0-50 m (P&G); 0-200 m 

(MM&B); this study, alive: 117-234 m. 

Order CHITONIDA Thiele, 1909 

Family Acanthochitonidae Simroth, 1894 

Acanthochitona fascicularis (Linnaeus, 1767) 

(Figure 2) 

Remarks: Only loose valves were found; however, 

the number and freshness of the valves 

collected suggests that it could live on nearby 

rocky habitats. Ávila et al., 2000. Depth range: 

0-50 m (P&G); 0-200 m (MM&B); this study: 

30-129 m. Alive on IVFC (Martins, 2004). 

Class GASTROPODA Cuvier, 1797 

Subclass PROSOBRANCHIA Milne Edwards, 

1848 

Order ARCHAEOGASTROPODA Thiele, 1925 

Suborder DOCOGLOSSA Troschel, 1866 

Superfamily PATELLOIDEA Rafinesque, 1815 

Family Patellidae Rafinesque, 1815 

Patella candei d’Orbigny, 1840 

Patella aspera Röding, 1798 

Remarks: Only small and very worn shells 

were found. These are known shallow water 

species and their presence in the samples is 

considered accidental. Alive on IVFC 

(Martins, 2004). Patella aspera has been incorrectly 

synonymized with P. ulyssiponensis 

Gmelin, 1791. Weber & Hawkins (2005) consider 

both genetically distinct, the name P. 

aspera referring to the Macaronesian populations 

whereas P. ulyssiponenis is applied to 

those in the continental coasts. 

Superfamily LOTIOIDEA Gray, 1840 

Family Lotiidae Gray, 1840 

Tectura virginea (O.F. Müller, 1776) 

(Figure 3) 

Remarks: Worn shells, in various degrees of 

preservation were found. The abundance, 

size and freshness of some specimens indicate 

that they could live on nearby rocky habitats. 

Ávila et al., 2000. Depth range: 0-100 m (P&G); 

this study: 14-360 m. Common alive on IVFC 

(Martins, 2004). 

Family Lepetidae Gray, 1840 

Propilidium exiguum (Thompson, 1844) 

(Figure 4) 

Remarks: Only one specimen found. Depth 

range: 7-600 m (P&G); 1.480-2.190 m (MM&B, 

as P. ancyloide Forbes, 1849); this study: 156- 

360 m. 

Suborder VETIGASTROPODA Salvini- 

Plawén, 1980 

Superfamily FISSURELLOIDEA Fleming, 1822 

Family Fissurellidae Fleming, 1822 

Emarginula sp. 

(Figure 5) 

Remarks: Only one specimen found, probably


E. rosea Monterosato in Locard, 1892, or a juvenile 

of E. guernei Dautzenberg & Fischer, 1896. 

Depth range: 0-90 m (P&G); 0 m (MM&B); this 

study: 117-234 m. 

Family Scissurellidae Gray, 1847 

Sinezona cingulata (O.G. Costa, 1861) 

(Figure 6) 

Remarks: Only one specimen found. Bullock et 

al., 1990 as Scissurella crispata Fleming, 1828; 

Bullock, 1995 as Schismope fayalensis 

Dautzenberg, 1889; Ávila et al., 2000. Depth 

range: 900 m (MM&B); this study: 117-145 m. 

Superfamily HALIOTOIDEA Rafinesque, 1815 

Family Haliotidae Rafinesque, 1815 

Haliotis coccinea Reeve, 1846 

(Figure 7) 

Remarks: Only small and worn shells were 

found. It is possible that the presence of shells 

in deeper water is accidental. Ávila et al., 2000. 

Depth range: 2-25 m (P&G; MM&B); this study: 

30-360 m. Common alive on IVFC (Martins, 

2004). 

?Haliotis sp. 

(Figure 8) 

Remarks: The only specimen collected had two 

holes topically similar to those of Haliotis and 

could be a juvenile. However, the shell morphology 

differs from that typical of the genus, 

namely the flattened columellar lip, and, therefore, 

it is only tentatively referred to Haliotis. 

Superfamily LEPETELLOIDEA Dall, 1882 

Family Lepetellidae Dall, 1882 

Lepetella laterocompressa (de Rayneval & Ponzi, 

1854) 

(Figure 9) 

Remarks: Common in small fractions of deeper 

samples. Depth range, this study: 99-234 m. 

Family Addisoniidae Dall, 1882 

Addisonia excentrica (Tiberi, 1855) 

(Figure 10) 

Remarks: Only one specimen found. Depth 

range: 370-3.307 m (MM&B); this study: 117- 

234 m. 

Superfamily TROCHOIDEA Rafinesque, 1815 

Family Trochidae Rafinesque, 1815 

Clelandella azorica Gofas, 2005 

(Figures 11-12) 

Remarks: Live specimens found only when the 

dredge hit hard surface. Endemic. Depth 

range, this study: 30-360 m; alive: 144-198 m. 

Clelandella sp. 

(Figure 13) 

Remarks: Rare. Depth range: 117-234 m. 

Jujubinus pseudogravinae Nordsieck, 1973 


Remarks: Although live specimens were not collected, 

some shells were fresh, indicating that 

they could live in nearby habitats. Endemic. 

Ávila et al., 2000. Depth range: 0-200 m (P&G, 

MM&B, as J. exasperatus (Pennant, 1777)); this 

study: 18-360 m. Common alive on IVFC 


Gibbula delgadensis Nordsieck, 1982 


Remarks: Uncommon. Not collected alive but 

some shells were fresh, indicating that they 

could live in nearby habitats. Endemic. Depth 

range, this study: 38-234 m. Common alive on 

IVFC (Martins, 2004). 

Gibbula magus (Linnaeus, 1758) 


Remarks: Common on sandy bottoms. The 

specimens from the Azores are smaller than 

their European conspecifics. Bullock et al., 

1990; Ávila et al., 2000. Depth range: 0-70 m 

(P&G); this study: 14-360 m; alive: 18-360 m. 

Common alive on IVFC (Martins, 2004) 

Margarites sp. 

(Figure 23) 

Remarks: A few rolled specimens collected. 

Depth range, this study: 99-198 m. 

Family Solariellidae Powell, 1951 

Solariella azorensis (Watson, 1886) 


Remarks: Relatively common. Some ultrajuvenile 

specimens collected (Figure 26) fit 

the description provided by Adam & 

Knudsen (1969) for Rhodinoliotia roseotincta 

(Smith, 1871); since specimens of the latter 

species were not available for comparison, a 

decision on synonymy is not warranted.

24 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE I 

1. Leptochiton cimicoides (Monterosato, 1879) (Sta37) 

2. Acanthochitona fascicularis (Linnaeus, 1767) (Sta31) 

3. Tectura virginea (O.F. Müller, 1776) (Sta28) 

4. Propilidium exiguum (Thompson, 1844) (Sta41) 

5. Emarginula sp. (Sta37) 

6. Sinezona cingulata (O.G. Costa, 1861) (Sta28) 

7. Haliotis coccinea Reeve, 1846 (Sta26) 

8. ?Haliotis cf. coccinea Reeve, 1846 (juvenile) (Sta29) 

9. Lepetella laterocompressa (Rayneval & Ponzi, 1854) (Sta38) 

10. Addisonia excentrica (Tiberi, 1855) (Sta37) 

11. Clelandella azorica Gofas, 2005 (Sta29) 

12. Clelandella azorica Gofas, 2005 (Sta53) 

13. Clelandella sp. (Sta37) 

14. Jujubinus pseudogravinae Nordsieck, 1973 (Sta44) 

15. Jujubinus pseudogravinae Nordsieck, 1973 (Sta27)

MARTINS ET AL: INFRALITORAL MOLLUSCS OFF VILA FRANCA DO CAMPO 25

26 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE II 

16. Gibbula delgadensis Nordsieck, 1982 (Sta6) 

17. Gibbula delgadensis Nordsieck, 1982 (Sta46) 

18. Gibbula magus (Linnaeus, 1758) (Sta44) 

19. Gibbula magus (Linnaeus, 1758) (Vila Franca do Campo, 1991) 




23. Margarites sp. (Sta29) 

24. Solariella azorensis Watson, 1886 (Sta29) 

25. Solariella azorensis Watson, 1886 (ultrajuvenile) (Sta12) 

26. Solariella azorensis Watson, 1886 (ultrajuvenile) (Sta12) 

27. Calliostoma hirondellei Dautzenberg & Fischer, 1896 (Sta37) 

28. Calliostoma lividum Dautzenberg 1927 (Sta37) 

29. Calliostoma lividum Dautzenberg 1927 (Sta45) 

30. Cirsonella gaudryi (Dautzenberg & Fisher, 1896) (Sta37) 

31. Tricolia pullus azorica (Dautzenberg, 1889) (Sta56) 

32. Tricolia pullus azorica (Dautzenberg, 1889) (Sta38)


28 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE III 

33. Bittium cf. latreillii (Payraudeau, 1826) (Sta1) 

34. Bittium cf. latreillii (Payraudeau, 1826) (Sta40) 

35. Bittium latreillii (Payraudeau, 1826) (Sta32) 

36. Fossarus ambiguus (Linnaeus, 1758 (Sta28) 

37. Fossarus ambiguus (Linnaeus, 1758 (Sta2) 

38. Cheirodonta pallescens (Jeffreys, 1867) (Sta1) 




42. Similiphora similior (Bouchet & Guillemot, 1978) (Sta18) 





47. Marshallora adversa (Montagu, 1803) (Sta13) 


49. Marshallora adversa (Montagu, 1803) Sta56)


30 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE IV 






55. Marshallora cf. adversa (Montagu, 1803) (Sta32) 

56. Monophorus sp. (Sta1) 

57. Monophorus erythrosoma (Bouchet & Guillemot, 1978) (Sta54) 




61. Pogonodon pseudocanaricus (Bouchet, 1985) (Sta1) 

62. Monophorus thiriotae Bouchet, 1985 (Sta32) 



65. Strobiligera brychia (Bouchet & Guillemot, 1978) (Sta37)


32 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE V 

66. Metaxia cf. abrupta (Watson, 1880) (Sta38) 

67. Cerithiopsis tubercularis (Montagu, 1803) (Sta2) 

68. Cerithiopsis tubercularis (Montagu, 1803) (Sta56) 

69. Cerithiopsis tiara (Monterosato, 1874) (Sta38) 

70. Cerithiopsis jeffreysi Watson, 1885 (Sta28) 

71. Cerithiopsis scalaris Locard, 1892 (Sta1) 




75. Cerithiopsis minima (Brusina, 1865) (Sta40) 

76. Cerithiopsis minima (Brusina, 1865) (Sta15) 

77. Cerithiopsis cf. minima (Brusina, 1865) (Sta2) 

78. Cerithiopsis cf. minima (Brusina, 1865) (Sta1) 

79. Cerithiopsis fayalensis Watson, 1886 (Sta28) 

80. Cerithiopsis fayalensis Watson, 1886 (Sta37) 

81. Krachia cf. guernei (Dautzenberg & Fischer, 1896) (Sta37)


34 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE VI 

82. Epitonium turtonis (Turton, 1819) (Sta44) 

83. Epitonium clathrus (Linnaeus, 1758) (Sta44) 

84. Epitonium pulchellum (Bivona, 1832) (Sta12) 



87. Epitonium celesti (Aradas, 1854) (Sta7) 




91. Punctiscala cerigottana (Sturany, 1896) (Sta38) 

92. Opaliopsis atlantis (Clench & Turner, 1952) (Sta7) 

93. Cirsotrema cf. cochlea (Sowerby, 1844) (Sta31) 

94. Cirsotrema cf. cochlea (Sowerby, 1844) (Sta18) 

95. Acirsa subdecussata (Cantraine, 1835) (juvenile) (Sta1) 

96. Opalia hellenica (Forbes, 1844) (Sta48) 

97. Opalia sp. 1 (Sta27) 

98. Opalia sp. 2 (Sta38)


36 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE VII 

99. Melanela bosci Payraudeau, 1826 (Sta37) 

100. Melanella cf. crosseana (Brusina, 1886) (Sta13) 

101. Melanella cf. crosseana (Brusina, 1886) (Sta2) 

102. Melanella cf. trunca (Watson, 1897) (Sta40) 

103. Parvioris microstoma (Brusina, 1864) (Sta2) 

104. Parvioris sp. (Sta2) 

105. Crinophteiros collinsi (Sykes, 1903) (Sta37) 

106. Sticteulima jeffreysiana (Brusina, 1869) (Sta12) 

107. Vitreolina sp. (Sta2) 

108. Vitreolina curva (Monterosato, 1884) (Sta1) 

109. Vitreolina curva (Monterosato, 1884) (Sta32) 

110. Pelseneeria minor Koehler & Vaney, 1908 (Sta28) 

111. Skeneopsis planorbis (Fabricius, 1870) (Sta1)


38 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE VIII 

112. Rissoa guernei Dautzenberg, 1889 (Sta27) 



115. Rissoa sp. 1 (Sta28) 

116. Rissoa sp. 2 (Sta28) 

117. Setia subvaricosa Gofas, 1991 (Sta27) 

118. Setia subvaricosa Gofas, 1991 (Sta27) 

119. Setia cf. quisquiliarum (Watson 1886) (Sta41) 

120. Crisilla postrema (Gofas, 1991) (Sta27) 

121. Crisilla cf. postrema (Sta27) 

122. Crisilla cf. postrema (Sta28) 

123. Pseudosetia azorica Bouchet & Warén, 1993 (Sta18) 

124. Pseudosetia azorica Bouchet & Warén, 1993 (Sta38) 

125. Cingula trifasciata (Adams, 1798) (Sta2) 

126. Manzonia unifasciata Dautzenberg, 1889 (Sta1) 



129. Onoba moreleti Dautzenberg, 1889 (Sta12) 



132. Onoba moreleti Dautzenberg, 1889 (Sta37)


40 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE IX 

133. Alvania angioyi van Aartsen, 1982 (Sta1) 

134. Alvania angioyi van Aartsen, 1982 (Sta28) 

135. Alvania poucheti (Dautzenberg, 1889) (Sta1) 



138. Alvania mediolittoralis Gofas, 1989 (Sta1) 

139. Alvania punctura (Montagu, 1803) (Sta37) 

140. Alvania sp. (?tarsodes Watson, 1886) (Sta1) 

141. Alvania sleursi (Amati, 1987) (Sta1) 

142. Alvania sleursi (Amati, 1987) (Sta37) 

143. Alvania cancellata (da Costa, 1778) (Sta1) 





148. Alvania platycephala Dautzenberg & Fisher, 1896 (Sta41) 




152. Alvania cimicoides (Forbes, 1844) (Sta37) 

153. Alvania cimicoides (Forbes, 1844) (Sta37) 

154. Alvania cf. cimicoides (Forbes, 1844) (Sta41)


42 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE X 

155. Caecum wayae Pizzini & Nofroni, 2001 (Sta12) 

156. Caecum wayae Pizzini & Nofroni, 2001 (Sta18) 

157. Talassia cf. tenuisculpta (Watson 1873) (Sta37) 

158. Talassia cf. tenuisculpta (Watson 1873) (Sta27) 

159. Capulus ungaricus (Linnaeus, 1758) (Sta7) 

160. Lamellaria perspicua (Linnaeus, 1758) (Vila Franca do Campo, 1991) 

161. Lamellaria perspicua (Linnaeus, 1758) (Sta56) 

162. Trivia pulex (Solander in J.E. Gray, 1828 (Vila Franca do Campo, 1991) 

163. Trivia pulex (Solander in J.E. Gray, 1828 (Sta56) 

164. Trivia pulex (Solander in J.E. Gray, 1828 (Sta56) 

165. Trivia candidula (Gaskoin, 1835) (Sta44) 

166. Erato sp. (juvenile) (Sta2) 

167. Erato sp. (juvenile) (Sta38) 

168. Aperiovula juanjosensii Perez & Gomez, 1987 (Sta38) 

169. Notocochlis dillwynii (Payraudeau, 1826) (Sta53) 

170. Natica prietoi (Hidalgo, 1873) (Sta15) 



173. Natica cf. prietoi (Hidalgo, 1873) (Sta57)


44 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XI 

174. Atlanta peronii Lesueur, 1817 (Sta13) 

175. Protatlanta souleyeti (E.A. Smith, 1888) (Sta13) 

176. Ocenebra erinaceus juvenile (Linnaeus, 1758) (Sta27) 

177. Ocenebra sp. (Sta56) 

178. Ocinebrina aciculata (Lamarck,1822) (Sta40) 

179. Ocinebrina cf. aciculata (Payraudeau, 1826) (Sta41) 

180. ? Ocinebrina aciculata (Lamarck,1822) (Sta40) 

181. ?Ocinebrina cf. aciculata (Payraudeau, 1826) (Sta27) 

182. ?Ocinebrina sp. (Sta15) 

183. Orania fusulus (Brocchi, 1814) (Sta1) 



186. Trophonopsis barvicensis (Johnston, 1825) (Sta29) 




190. Trophonopsis barvicensis (Johnston, 1825) (Sta27)


46 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XII 



193. Trophonopsis cf. muricatus (Montagu,1803) (Sta5) 

194. Trophonopsis cf. muricatus (Montagu,1803) (Sta27) 

195. Coralliophila cf. meyendorfii (Calcara, 1845) (Sta7) 

196. Coralliophila panormitana (Monterosato, 1896) (Sta37) 

197. Stramonita haemastoma (Linnaeus, 1767) (Sta27) 

198. Gibberula vignali (Dautzenberg & Fischer 1896) (Sta41) 

199. Gibberula cf. lazaroi Contreras, 1992 (Sta37) 

200. Mitra cornea Lamarck, 1811 (Sta31) 

201. Pollia dorbignyi (Payraudeau, 1826) (Sta45) 

202. Pollia dorbignyi (Payraudeau, 1826) Ilhéu de Vila Franca do Campo. 

203. Nassarius incrassatus (Ström, 1768) (Sta40) 



206. Nassarius incrassatus (teratology) (Ström, 1768) (Sta40) 

207. Nassarius cf. cuvierii (Payraudeau, 1826). Juvenile (Sta44) 

208. Nassarius recidivus (Martens, 1876) (Sta53) 

209. Columbella adansoni Menke, 1853 (Sta44) 

210. Mitrella pallaryi (Dautzenberg, 1927) (Sta37) 

211. Anachis avaroides Nordsieck, 1975 (Sta44) 

212. Anachis avaroides Nordsieck, 1975 (Sta1)


48 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XIII 

213. Brocchinia clenchi Petit, 1986 (Sta49) 

214. Brocchinia clenchi Petit, 1986 (Sta1) 

215. Mitromorpha azorensis Mifsud, 2001 (Sta46) 

216. Mitromorpha azorensis Mifsud, 2001 (Sta1) 

217. Bela nebula (Montagu, 1803) (Sta56) 






223. Mangelia cf. costata (Donovan, 1804) (Sta25) 







230. Raphitoma purpurea (Montagu, 1803) (Sta56) 

231. Raphitoma linearis (Montagu, 1803) (Sta40)


50 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XIV 

232. Raphitoma cf. aequalis (Jeffreys, 1867) (Sta56) 









241. Raphitoma sp. (Sta58) 

242. Pleurotomella gibbera Bouchet & Warén, 1980 (Sta29) 



245. Pleurotomella cf. gibbera Bouchet & Warén, 1980 (Sta29) 

246. Teretia teres (Reeve, 1844) (Sta28) 

247. Teretia teres (Reeve, 1844) (Sta27)


52 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XV 

248. Crassopleura maravignae (Bivona, 1838) (Sta56) 



251. Haedropleura septangularis (Montagu, 1803) (Sta15) 

252. Haedropleura septangularis (Montagu, 1803) (Sta56) 

253. Philippia krebsi (Mörch, 1875) (Sta46) 

254. Pseudotorinia architae (O.G. Costa, 1841) (Sta32) 

255. Pseudomalaxis zanclaeus (Philippi, 1844) (Sta37) 

256. Mathilda cochlaeformis Brugnone, 1873 (Sta29) 

257. Mathilda retusa Brugnone, 1873 (Sta37)


54 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XVI 

258. ?Rissoella sp. 1 (Sta41) 

259. ?Rissoella sp. 2 (Sta2) 

260. Omalogyra atomus (Philippi, 1841) (Sta28) 

261. Odostomella doliolum (Philippi, 1844) (Sta37) 

262. Odostomella doliolum (Philippi, 1844) (Sta32) 

263. Chrysallida cf. flexuosa (Monterosato, 1874 ex Jeffreys) (Sta41) 

264. Odostomia bernardi Aartsen, Gittenberger & Goud, 1998 (Sta58) 

265. Odostomia cf. verhoeveni Aartsen, Gittenberger & Goud, 1998 (Sta1) 

266. Odostomia duureni Aartsen, Gittenberger & Goud, 1998 (Sta15) 

267. Odostomia cf. striolata Forbes & Hanley, 1850 (Sta18) 

268. Eulimella sp. (Sta38) 

269. Turbonilla rufa (Philippi, 1836) (Sta1) 

270. Turbonilla lactea (Linnaeus, 1758) (Sta56) 

271. Turbonilla sp. 1 (Sta38) 




275. Ebala nitidissima (Montagu, 1803) (Sta28) 

276. Ebala nitidissima (Montagu, 1803) (Sta18)


56 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XVII 

277. Colpodaspis pusilla Sars, 1870 (Sta7) 

278. Retusa truncatula (Bruguière, 1792) (Sta27) 

279. Retusa truncatula (Bruguière, 1792) (Sta1) 

280. Haminoea cf. orteai Talavera, Murillo & Templado, 1987 (Sta58) 

281. Atys macandrewi E.A. Smith, 1872 (Sta56) 

282. Atys sp. (Sta15) 

283. Philine approximans Dautzenberg & Fischer, 1896 (Sta13) 

284. Philine sp. (Sta18) 

285. ?Chelidonura africana Pruvot-Fol, 1953 (Sta37) 

286. Cavolinia inflexa (Lesueur, 1813) (Sta56) 

287. Cavolinia inflexa (Lesueur, 1813) (Sta29) 

288. Cavolinia tridentata (Forskal, 1775) (Sta37) 

289. Diacria trispinosa (Lesueur, 1821) (Sta50) 

290. Diacria trispinosa (Lesueur, 1821) (Sta12) 

291. Cuvierina atlantica (Bé, MacClintock & Currie, 1972) (Sta38) 

292. Clio pyramidata Linnaeus, 1767 (Sta29) 

293. Clio pyramidata Linnaeus, 1767 (Sta12) 

294. Limacina cf. helicina (Phipps, 1774) (Sta32) 

295. Limacina inflata (d’Orbigny, 1836) (Sta1) 

296. Umbraculum umbraculum (Lightfoot, 1786) (Sta31) 

297. Tylodina perversa (Gmelin, 1791) (Sta28) 

298. Williamia gussonii (O.G. Costa, 1829) (St12)


58 AÇOREANA 

2009, Sup. 6: 15-103 

Endemic. Depth range, this study: 14-360 m; 

alive from 14-207 m. 

Calliostomatidae Thiele, 1924 

Calliostoma hirondellei Dautzenberg & Fisher, 

1896 

(Figure 27) 

Remarks: Rare. Depth range, this study: 72- 

234 m. 

Calliostoma lividum Dautzenberg, 1927 


Remarks: Sometimes subtidal in the Azores. 

Endemic. Ávila et al., 2000 as C. cf. conulus 

(Linnaeus, 1758). Depth range: 20-200 m 

(P&G and MM&B, as C. conulum (L.)); this 

study: 30-360 m. Alive on IVFC (Martins, 

2004). 

Superfamily TURBINOIDEA Rafinesque, 

1815 

Family Turbinidae Rafinesque, 1815 

Cirsonella gaudryi (Dautzenberg & Fisher, 

1896) 

(Figure 30) 


234 m. 

Family Phasianellidae Swainson, 1840 

Tricolia pullus azorica (Dautzenberg, 1889) 


Remarks: Some specimens were fresh, indicating 

that they could live in nearby habitats. 

Endemic. Bullock et al., 1990, Bullock, 1995 

and Knudsen, 1995 as T. pullus (Linnaeus, 

1758); Ávila et al., 2000. Depth range: intertidal-35 

m (P&G, as T. pullus (L.)); this study: 

14-360 m. Common alive on IVFC (Martins, 

2004). 

Order APOGASTROPODA Salvini-Plawén & 

Haszprunar, 1987 

Suborder CAENOGASTROPODA Cox, 1959 

Superfamily CERITHIOIDEA Fleming, 1822 

Family Cerithiidae Fleming, 1822 

Bittium cf. latreillii (Payraudeau, 1826) 


Remarks: This form of Bittium has been questionably 

ascribed to B. latreillii, and awaits further 

study to clarify its taxonomic status. 

Shells are very common. Bullock et al., 1990 

and Bullock, 1995 as B. reticulatum (da Costa, 

1779); Ávila et al., 2000. Depth range: littoral 

(MM&B); this study: 14-360 m; alive at 38 m. 

Common alive on IVFC (Martins, 2004). 

Bittium latreillii (Payraudeau, 1826) 

(Figure 35) 

Remarks: Very rare. This specimen conforms to 

the description of B. latreillii. Depth range: littoral 

(MM&B); this study: 180 m. 

Family Planaxidae Gray, 1850 

Fossarus ambiguus (Linaeus, 1758) 


Remarks: Occurs regularly in the samples. 

Houbrick, 1990; Bullock, 1995; Knudsen, 1995; 

Ávila et al., 2000. Depth range: littoral 

(MM&B); this study: 30-180 m. Common alive 

on IVFC (Martins, 2004). 

Superfamily TRIPHOROIDEA Gray, 1847 

Triphoridae Gray, 1847 

NOTE: Triphorids were present in most samples. 

No living specimens were collected, 

although some shells appeared to be fresh; 

commonly, they exhibited clear signs of predation. 

The species of this family are difficult to 

identify without information on the animal 

and the larval shell; the granulation of the 

spiral and basal cords is often less distinct and 

intermediate. Although tentatively identified 

to species, they are however illustrated 

profusely to record their variability, and 

to provide a basis for further, perhaps more 

accurate identifications. Information on 

www.naturamediterraneo.com, based on 

Bouchet & Guillemot (1978) and Bouchet 

(1984), was helpful for species identification. 

Cheirodonta pallescens (Jeffreys, 1867) 


Remarks: Shell light-brown to whitish, uniformly 

coloured; protoconch bi-carinated; spiral 

cord 4 and basal cords smooth; supranumerary 

cords on last whorl. Depth range, this study: 

30-145 m. 

Similiphora similior (Bouchet & Guillemot, 1978) 


Remarks: Shell brownish, variously coloured; 1 st 

whorl of protoconch uni-carinated, remaining 

bi-carinated; spiral cord 4 granulated, basal


cords smooth; supranumerary cords on last 

whorl. Depth range, this study: 14-360 m. 

Marshallora adversa (Montagu, 1803) 


Remarks: Shell brownish, variously coloured; 

protoconch bi-carinated; spiral cord 4 and 

basal cords smooth; tubercles on last whorl 

elongated; absence of supranumerary cords on 

last whorl. Depth range, this study: 30-234 m. 

Marshallora cf. adversa (Montagu, 1803) 

(Figure 55) 

Remarks: The presence of brownish markings 

between tubercles is reminiscent of M. thiriotae, 

but other characters indicate affinity with M. 

adversa. Depth range, this study: 46 m. 

Monophorus sp. 

(Figure 56) 

Remarks: Depth range, this study: 40-135 m. 

Monophorus erythrosoma (Bouchet & Guillemot, 

1978) 


Remarks: Shell brownish, unicoloured or variously 

coloured; protoconch bi-carinated; spiral 

cord 4 granulated, basal cords 1-2 granulated, 3 

smooth; supranumerary cords on last whorl. 


Monophorus thiriotae Bouchet, 1985 


Remarks: Shell brownish, variously coloured; 

protoconch bi-carinated; spiral cord 4 granulated, 

basal cords 1-2 granulated, 3 smooth; 

absence of supranumerary cords on last whorl; 

brownish intertuberculary markings. Depth 

range, this study: 30-207 m. 

Pogonodon pseudocanaricus (Bouchet, 1985) 

(Figure 61) 

Remarks: Shell whitish, brown vertical markings; 

protoconch redish, bi-carinated; spiral cord 4 

granulated, basal cords 1-2 granulated. Only 

one apparently fresh specimen, but with last 

whorl crushed. Depth range, this study: 57 m. 

Strobiligera brychia (Bouchet & Guillemot, 

1978) 

(Figure 65) 

Remarks: Tubercles of spiral cord 1 smaller than 

those of remaining cords. Rare. Depth range, 

this study: 117-234 m. 

Metaxia cf. abrupta (Watson, 1880) 

(Figure 66) 

Remarks: Only one specimen collected, apparently 

a juvenile. Depth range, this study: 129- 

207 m. 

Family Cerithiopsidae Adams H. & A., 1853 

Cerithiopsis tubercularis (Montagu, 1803) 


Remarks: Relatively rare. Depth range: littoral- 

100 m (MM&B); this study: 41-207m. 

Cerithiopsis tiara (Monterosato, 1874) 

(Figure 69) 


207 m. 

Cerithiopsis jeffreysi Watson, 1885 

(Figure 70) 


207 m. 

Cerithiopsis scalaris Locard, 1892 


Remarks: Shells uncommon but some specimens 

appeared fresh. Depth range, this study: 

57-234 m. 

Cerithiopsis minima (Brusina, 1865) 


Remarks: Uncommon. Depth range, this study: 

38-207 m. 

Cerithiopsis cf. minima (Brusina, 1865) 



57-135 m. 

Cerithiopsis fayalensis Watson, 1886 


Remarks: Shells uncommon but some specimens 

appeared fresh. Depth range, this study: 

99-234 m. 

Krachia cf. guernei (Dautzenberg & Fischer, 

1896) 

(Figure 81) 


234 m.

60 AÇOREANA 

2009, Sup. 6: 15-103 

Superfamily JANTHINOIDEA Gray, 1847 

Family Epitoniidae S.S. Berry, 1910 (1812) 

Gyroscala lamellosa (Lamarck, 1822) 

Remarks: Only one fragment. Depth range: 

Infralittoral to 620 m (MM&B); this study: 56 m. 

Epitonium turtonis (Turton, 1819) 

(Figure 82) 

Remarks: Rare. Depth range: 5-70 m (P&G); this 

study: 66m. 

Epitonium clathrus (Linnaeus, 1758) 

(Figure 83) 

Remarks: Rare. Depth range: 5-70 m (P&G); this 

study: 66-72 m. 

Epitonium pulchellum (Bivona, 1832) 


Remarks: Not uncommon. Depth range: 20-40 m 

(P&G; MM&B); this study: 58-145 m. 

Epitonium celesti (Aradas, 1854) 


Remarks: Not uncommon. Depth range: 50-1250 

m (P&G; MM&B); this study: 57-207 m. 

Punctiscala cerigottana (Sturany, 1819) 

(Figure 91) 

Remarks: Rare. Only one damaged specimen 

found. Depth range: 50-600 m (MM&B); this 

study: 129-207 m. 

Opaliopsis atlantis (Clench & Turner, 1952) 

(Figure 92) 

Remarks: Rare. Only one damaged specimen 

found. Depth range: 810-825 m (MM&B); this 

study: 167-189 m. 

Cirsotrema cf. cochlea (Sowerby, 1844) 


Remarks: Uncommon. The Azorean specimens 

are stouter and more globose than the illustrations 

consulted (P&G). Depth range: 

Infralittoral-60 m (MM&B); this study: 40-318 m; 

alive: 66-72 m. 

Acirsa subdecussata (Cantraine, 1835) 

(Figure 95) 

Remarks: Only one fresh specimen collected. 

Depth range: 12-500 m (P&G; MM&B); this study: 

57 m. 

Opalia hellenica (Forbes, 1844) 

(Figure 96) 

Remarks: Uncommon. Ávila et al., 2000. Depth 

range: 20-770 m (MM&B); this study: 63-234 m; 

alive: 66-81 m. 

Opalia sp. 1 

(Figure 97) 


108 m. 

Opalia sp. 2 

(Figure 98) 

Remarks: Only one shell collected, with broken 

tip. Depth range, this study: 129-207 m. 

Superfamily EULIMOIDEA Philippi, 1853 

Family Eulimidae Philippi, 1853 

Melanella bosci Payraudeau, 1826 

(Figure 99) 

Remarks: Rare. Depth range: 10-150 m (P&G); 


Melanella cf. crosseana (Brusina, 1886) 

(Figures 100-101) 


73-145 m. 

Melanella cf. trunca (Watson, 1897) 

(Figure 102) 


38 m. 

Parvioris microstoma (Brusina, 1864) 

(Figure 103) 

Remarks: Rare. Depth range, this study: 135 m. 

Parvioris sp. 

(Figure 104) 


207 m. 

Crinophteiros collinsi (Sykes, 1903) 

(Figure 105) 


234 m. 

Sticteulima jeffreysiana (Brusina, 1869) 

(Figure 106) 


121 m.


Vitreolina sp. 

(Figure 107) 


Vitreolina curva (Monterosato, 1884) 

(Figures 108-109) 

Remarks: Rare. Depth range, this study: 57-180 m. 

Pelseneeria minor Koehler & Vaney, 1908 

(Figure 110) 

Remarks: Only one specimen collected. Depth 

range: 90-185 m (MM&B); this study: 117-145 m. 

Superfamily LITTORINOIDEA Children, 1834 

Family Littorinidae Children, 1834 

Littorina striata King & Broderip, 1832 

Remarks: This is a supralittoral species, and 

its presence in the dredged material is accidental. 

Melarhaphe neritoides (Linnaeus, 1758) 

Remarks: This is a supralittoral species, and its 

presence in the dredged material is accidental. 

Family Skeneopsidae Iredale, 1915 

Skeneopsis planorbis (Fabricius, 1870) 

(Figure 111) 

Remarks: Uncommon. Reported as very common 

(Bullock et al., 1990; Bullock, 1995). Knudsen, 1995; 

Ávila et al., 2000. Depth range: Infralittoral to 70m 

(P&G; MM&B); this study: 32-145 m. Common 

alive on IVFC (Martins, 2004). 

Superfamily RISSOOIDEA Gray, 1847 

Family Rissoidae Gray, 1847 

Rissoa guernei Dautzenberg, 1889 

(Figures 112-114) 

Remarks: Uncommon but some specimens 

appeared fresh. Endemic. Reported as very common 

(Bullock et al., 1990; Ávila, 2000). Gofas, 1990; 

Knudsen, 1995; Ávila et al., 2000. Depth range, this 

study: 32-234 m. Common alive on IVFC 


Rissoa sp. 1 

(Figure 115) 


Rissoa sp. 2 

(Figure 116) 


Setia subvaricosa Gofas, 1990 

(Figures 117-118) 

Remarks: Rare. Endemic. This species is 

common in 15-20 m (Gofas, 1990; Ávila, 2000). 

Ávila et al., 2000. Depth range, this study: 99- 

135 m. Collected alive at IVFC (Martins, 2004). 

Setia cf. quisquiliarum (Watson 1886) 

(Figure 119) 

Remarks: Rare. Endemic. Depth range, this 

study: 237-360 m. 

Crisilla postrema (Gofas, 1990) 

(Figure 120) 

Remarks: Rare. Endemic. Infralittoral to 20 m; 

collected alive at IVFC (Gofas, 1990). Bullock et 

al., 1990 as Rissoa pulcherima (Jeffreys, 1848); 

Bullock, 1995 as Alvania postrema. Depth range, 

this study: 56-360m. 

Crisilla cf. postrema (Gofas, 1990) 

(Figures 121-122) 


207 m. 

Pseudosetia azorica Bouchet & Warén, 1993 

(Figures 123-124) 


study: 72-207 m. 

Cingula trifasciata (Adams, 1798) 

(Figure 125) 

Remarks: Rarely collected. Specimens eroded, 

probably transported from the intertidal, 

where it is common. Depth range, this study: 

38-180 m. 

Manzonia unifasciata Dautzenberg, 1889 

(Figures 126-128) 

Remarks: Relatively common; probably transported 

from infralittoral zone where it is very 

common from 0-10 m (Ávila, 2003). Bullock et 

al., 1990 and Bullock, 1995 as M. crassa 

(Kanmacher, 1798); Gofas, 1990; Knudsen, 

1995; 1995; Ávila et al., 2003. Depth range, this 

study: 237-360 m. Collected alive at IVFC 


Onoba moreleti Dautzenberg, 1889 

(Figures 129-132) 

Remarks: Uncommon. Endemic. Depth range, 

this study: 95-234 m.

62 AÇOREANA 

2009, Sup. 6: 15-103 

Alvania angioyi van Aartsen, 1982 

(Figures 133-134) 

Remarks: Rare. Endemic. Infralittoral to 20 m 

(Gofas, 1990). Bullock et al., 1990 as A, watsoni 

(Schwartz MS) Watson, 1873; Bullock, 1995; 

Knudsen, 1995; Ávila et al. 2000. Depth range, 

this study: 57-360 m. Collected alive at IVFC 


Alvania poucheti Dautzenberg, 1889 

(Figures 135-137) 

Remarks: Relatively common; some specimens 

appeared fresh. Endemic. Gofas, 1990; 

Bullock et al., 1990; Bullock, 1995; Knudsen, 

1995; Ávila et al. 2000. Depth range, this 

study: 38-207 m. Collected alive at IVFC 


Alvania mediolittoralis Gofas, 1989 

(Figure 138) 

Remarks: Rare. Common intertidally (Gofas, 

1990). Depth range, this study: 30-207 m. 

Alvania punctura (Montagu, 1803) 

(Figure 139) 

Remarks: Very rare. Depth range, this study: 

117-234 m. 

Alvania sp. (?tarsodes Watson, 1886) 

(Figure 140) 

Remarks: Very rare. Depth range, this study: 

57 m. 

Alvania sleursi (Amati, 1987) 

(Figures 141-142) 

Remarks: Common; some specimens appeared 

fresh. Infralittoral, common at 20 m (Gofas, 

1990; Knudsen, 1995). Depth range, this study: 

30-234 m. 

Alvania cancellata (da Costa, 1778) 

(Figures 143-147) 


fresh. Infralittoral, most common at 20 m 

(Gofas, 1990; Knudsen, 1995). Depth range, 


Alvania platycephala Dautzenberg & Fischer, 

1896 

(Figures 148-151) 


study: 129-360 m. 

Alvania cimicoides (Hoenselaar & Goud, 

1998) 

(Figures 152-153) 


360 m. Collected alive at 117-234. 

Alvania cf. cimicoides (Hoenselaar & Goud, 

1998) 

(Figure 154) 

Remarks: Only one specimen collected. Depth 


Family Caecidae Gray, 1850 

Caecum wayae Pizzini & Nofroni, 2001 

(Figures 155-156) 

Remarks: Common. Depth range, this study: 

57-171 m. 

Superfamily VANIKOROIDEA Gray, 1840 

Family Vanikoridae Gray, 1840 

Talassia cf. tenuisculpta (Watson, 1873) 

(Figures 157-158) 


Superfamily CAPULOIDEA Fleming, 1822 

Family Capulidae Fleming, 1822 

Capulus ungaricus (Linnaeus, 1758) 

(Figure 159) 

Remarks: Rare. Depth range: sublittoral to 850 

m (P&G). Depth range, this study: 117-189 m. 

Superfamily VELUTINOIDEA Gray, 1850 

Family Velutinidae Gray, 1850 

Lamellaria perspicua (Linnaeus, 1758) 

(Figures 160-161) 

Remarks: Uncommon. Depth range: littoral to 

200 m (P&G; MM&B.) Depth range, this study: 

58-135 m. Collected alive at about 20 m, in previous 

workshop. 

Family Triviidae Troschel, 1863 

Trivia pulex (Solander in J.E. Gray, 1828) 

(Figures 162-164) 


30-234 m. Ávila et al., 2000. Collected alive at 

about 20 m, during previous workshop. 

Trivia candidula (Gaskoin, 1835) 

(Figure 165) 


fresh. Intertidal (MM&B). Ávila et al., 2000. 

Depth range, this study: 12-360 m.


Erato sp. 

(Figures 166-167) 

Remarks: Rare; the specimens herein represented 

are tentatively identified as juveniles. 

Depth range, this study: 135-207 m 

Family Ovulidae Feming, 1822 

Aperiovula juanjosensii Perez & Gomez, 1987 

(Figure 168) 

Remarks: Only a fragment was collected. 


Superfamily NATICOIDEA Guilding, 1834 

Family Naticidae Guilding, 1834 

Notocochlis dillwynii (Payraudeau, 1826) 

(Figure 169) 

Remarks: Rare. Depth range: 1-25 m (MM&B; 

P&G); this study: 318 m 

Natica prietoi Hidalgo, 1873 

(Figures 170-172) 

Remarks: Common, variable. Previously cited 

as Natica adansoni de Blainville, 1825 (Ávila et 

al., 2000). Depth range: infralittoral to 200 m 

(MM&B); this study: 14-360 m; alive: 18- 

318 m. 

Natica cf. prietoi Hidalgo, 1873 

(Figure 173) 


Superfamily TONNOIDEA Suter, 1913 

Family Tonnidae Suter, 1913 

Phalium undulatum (Gmelin, 1791) 

Remarks: Only fragments. Depth range: 8-80 

m (P&G); infralittoral to 115 m (MM&B); this 

study: 32-207 m. 

Superfamily PTEROTRACHEOIDEA 

Rafinesque, 1814 

Family Atlantidae Rang, 1829 

Atlanta peroni Lesueur, 1817 

(Figure 174) 

Remarks: Pelagic. 

Protatlanta souleyeti (E.A. Smith, 1888) 

(Figure 175) 

Remarks: Pelagic. 

Superfamily MURICOIDEA Rafinesque, 1815 

Family Muricidae Rafinesque, 1815 

Ocenebra erinaceus (Linnaeus, 1758) 

(Figure 176) 

Remarks: Rare. Only juveniles collected. Depth 

range: intertidal to150 m (P&G); this study: 32- 

207 m. 

Ocenebra sp. 

(Figure 177) 


58-189 m. 

Ocinebrina aciculata (Lamarck, 1822) 

(Figure 178) 

Remarks: Relatively common. Ávila et al., 2000. 

Depth range: intertidal to at least 25 m (P&G); this 

study: 30-360 m. 

Ocinebrina cf. aciculata (Lamarck, 1822) 

(Figure 179) 

Remarks: Relatively common. Depth range, this 

study: 38-360 m. 

?Ocinebrina cf. aciculata (Lamarck, 1822) 

(Figures 180-181) 


45-207 m. 

?Ocinebrina sp. 

(Figure 182) 


Orania fusulus (Brocchi, 1814) 

(Figures 183-185) 

Remarks: Depth range: 95-920 m (MM&B); 100- 

150 m (P&G); this study: 30-360 m; alive: 30-57 m. 

Trophonopsis barvicensis (Johnston, 1825) 

(Figures 186-192) 

Remarks: Commonly found. Depth range: 440-550 

m (MM&B); this study: 32-207 m. 

Trophonopsis cf. muricatus (Montagu, 1803) 

(Figures 193-194) 

Remarks: Commonly found. Ávila et al., 2000. 

Depth range: 10-200 m, with records down to 

2000 m (MM&B; P&G); this study: 41-108 m. 

Coralliophila cf. meyendorfii (Calcara, 1845) 

(Figure 195) 

Remarks: Uncommon. Depth range: infralittoral 

to circalittoral (MM&B); this study: 30-189 m. 

Coralliophila panormitana (Monterosato, 1896) 

(Figure 196) 

Remarks: Rare. Depth range: below low tide to 

640m (P&G); this study: 99-234 m.

64 AÇOREANA 

2009, Sup. 6: 15-103 

Stramonita haemastoma (Linnaeus, 1767) 

(Figure 197) 

Remarks: Uncommon; specimens rolled, mostly 

juvenile, probably transported from the littoral. 

Knudsen, 1995 as Thais haemastoma floridana 

(Conrad, 1837); Ávila et al., 2000. Depth range: 

intertidal to 3 m (P&G); this study: 32-360 m. 

Alive on IVFC (Martins, 2004). 

Family Cysticidae Stimpson, 1865 

Gibberula vignali (Dautzenberg & Fischer 1896) 

(Figure 198) 

Remarks: Rare. Depth range, this study: 156-360m. 

Gibberula cf. lazaroi Contreras, 1992 

(Figure 199) 

Remarks: Rare. Could be a juvenile of G. vignali, 

but resembles Gibberula lazaroi, described from the 

intertidal of Pico and Terceira islands (Contreras, 

1992). Depth range, this study: 117-234 m. 

Family Mitridae Swainson, 1831 

Mitra cornea Lamarck, 1811 

(Figure 200) 

Remarks: Uncommon; specimens rolled, mostly 

juveniles. Knudsen, 1995 as M. nigra (Gmelin, 

1791); Ávila et al., 2000. Depth range: intertidal 

to 40 m (MM&B, as M. cornicula (Linnaeus); 

this study: 30-207 m; alive at IVFC (Martins, 

2004). 

Superfamily BUCCINOIDEA Rafinesque, 1815 

Family Buccinidae Rafinesque, 1815 

Pollia dorbignyi (Payraudeau, 1826) 

(Figures 201-202) 

Remarks: one fresh fragment. Depth range: in and 

just above littoral zone (MM&B; P&G); this study: 

30 m; alive at IVFC (Martins, 2004). 

Family Nassariidae Iredale, 1916 

Nassarius incrassatus (Ström, 1768) 

(Figures 203-206) 

Remarks: Common, mostly fragmented shells. 

Ávila et al., 2000. Depth range: intertidal to 200 m 

(MM&B; P&G); this study: 18-.360 m. 

Nassarius cf. cuvierii (Payraudeau, 1826) 

(Figure 207) 

Remarks: One juvenile specimen collected. 

Depth range: in and just above littoral zone 

(MM&B; P&G); this study: 66 m. 

Nassarius recidivus (Martens, 1876) 

(Figure 208) 


Family Columbellidae Swainson, 1840 

Columbella adansoni Menke, 1853 

(Figure 209) 

Remarks: Uncommon, mostly fragmented shells; 

some juveniles appeared fresh. Knudsen, 1995; 

Ávila et al., 2000. Depth range: 4-1402 m (MM&B, 

as C. rustica (Linnaeus)); this study: 30-360 m; 

common alive at IVFC (Martins, 2004). 

Mitrella pallaryi (Dautzenberg, 1758) 

(Figure 210) 

Remarks: Rare. Depth range: 40-200 m (MM&B; 

P&G); this study: 129-360 m. 

Anachis avaroides Nordsieck, 1975 

(Figures 211-212) 

Remarks: Common. Ávila et al., 2000. Depth 

range: infralittoral (MM&B), this study: 30-207 m; 

common alive at IVFC (Martins, 2004). 

Superfamily CANCELLARIOIDEA Forbes & 

Hanley, 1851 

Family Cancellariidae Forbes & Hanley, 1851 

Brocchinia clenchi Petit, 1986 

(Figures 213-214) 

Remarks: Frequent in some samples. Depth range, 

this study: 40-07 m; alive: 58-81 m. 

Superfamily CONOIDEA Fleming, 1822 

Family Conidae Fleming, 1822 

Mitromorpha (Mitrolumna) azorensis Mifsud, 2001 

(Figures 215-216) 

Remarks: Uncommon. Bullock, 1995 as Mitrolumna 

olivoidea (Cantraine, 1835); Ávila et al., 2000. 

Depth range, this study: 30-207 m 

Bela nebula (Montagu, 1803) 

(Figures 217-222) 

Remarks: Common throughout the sampled depth 

range. Ávila et al., 2000. Depth range: 10-30 m 

(P&B); this study: 14-360 m; alive: 38-169 m. 

Mangelia cf. costata (Montagu, 1803) 

(Figures 223-229) 

Remarks: Common. Depth range: infralittoral 

40 m (MM&B); this study: 14-360 m; alive: 66- 

189 m.


Raphitoma purpurea (Montagu, 1803) 

(Figure 230) 


30-207 m 

Raphitoma linearis (Montagu, 1803) 

(Figure 231) 

Remarks: Uncommon. Depth range: Intertidal to 

200 m (P&G); this study: 38-234 m. 

Raphitoma cf. aequalis (Jeffreys, 1867) 

(Figures 232-240) 

Remarks: Common. Depth range: intertidal 

(MM&B); this study: 14-360 m; alive: 46-58 m. 

Raphitoma sp. 

(Figure 241) 


Pleurotomella gibbera Bouchet & Warén, 1980 

(Figures 242-244) 

Remarks: Rare. Depth range, this study: 72- 234 m. 

Pleurotomella cf. gibbera Bouchet & Warén, 1980 

(Figure 245) 


Teretia teres (Reeve, 1844) 

(Figures 246-247) 

Remarks: Rare. Depth range: 30-1385 (MM&B); 


Family Drilliidae Olsson, 1964 

Crassopleura maravignae (Bivona, 1838) 

(Figures 248-250) 

Remarks: Common at deeper sandy bottoms. 

Ávila et al., 2000. Depth range, this study: 14-360 

m; alive at: 58-234 m. 

Family Turridae H. & A. Adams, 1853 

Haedropleura septangularis (Montagu, 1803) 

(Figures 251-252) 


range: 7-70 m (P&G); this study: 30-207 m 

Subclass HETEROBRANCHIA Gray, 1840 

Order HETEROSTROPHA Fischer P., 1885 

Superfamily ARCHITECTONICOIDEA Gray, 

1850 

Family Architectonicidae Gray, 1850 

Philippia krebsi (Mörch, 1875) 

(Figure 253) 


Pseudotorinia architae (Costa O.G., 1841) 

(Figure 254) 

Remarks: Rare. Ávila et al., 2000. Depth range, 

this study: 180-234 m; alive at: 180m. 

Pseudomalaxis zancleus (Philippi, 1844) 

(Figure 255) 

Remarks: Rare. Depth range: 644 m (MM&B); 


Superfamily MATHILDOIDEA Dall, 1889 

Family Mathildidae Semper, 1865 

Mathilda cochlaeformis Brugnone, 1873 

(Figure 256) 

Remarks: Rare; only one fresh, broken shell. Depth 

range: 1205 m (MM&B); this study: 144-198 m. 

Mathilda retusa (Brocchi, 1814) 

(Figure 257) 

Remarks: Rare; only one specimen collected. 


Superfamily RISSOELLOIDEA Gray, 1850 

Family Rissoellidae Gray, 1850 

?Rissoella sp. 1 

(Figure 258) 


? Rissoella sp. 2 

(Figure 259) 


Superfamily OMALOGYROIDEA Sars, 1878 

Family Omalogyridae Sars, 1878 

Omalogyra atomus (Philippi, 1841) 

(Figure 260) 

Remarks: Rare; only one specimen; however, 

the rarity of minute species can be seen as an 

artefact of sampling, for most of the fine fraction 

was discarded. Bullock et al. (1990) reported 

it to be very common on algal samples. 

Bullock, 1995; Knudsen, 1995; Ávila et al., 2000. 

Depth range: intertidal to 20 m (MM&B); this 

study: 117-145 m. 

Superfamily PYRAMIDELLOIDEA Gray, 1840 

Family Pyramidellidae Gray, 1840 

Odostomella doliolum (Philippi, 1844) 

(Figures 261-262) 

Remarks: Rare; one very fresh shell collected. 

Ávila et al., 2000. Depth range: 10-800 m 

(MM&B); this study: 180-234 m.

66 AÇOREANA 

2009, Sup. 6: 15-103 

Chrysallida cf. flexuosa (Monterosato, 1874 ex 

Jeffreys) 

(Figure 263) 


300 m. 

Odostomia bernardi Aartsen, Gittenberger & 

Goud, 1998 

(Figure 264) 

Remarks: Common in fine fractions. Endemic. 

Depth range, this study: 30-360m. 

Odostomia cf. verhoeveni Aartsen, Gittenberger 

& Goud, 1998 

(Figure 265) 


30-207 m. 

Odostomia duureni Aartsen, Gittenberger & 

Goud, 1998 

(Figure 266) 


Odostomia cf. striolata Forbes & Hanley, 1850 

(Figure 267) 


234 m. 

Eulimella sp. 

(Figure 268) 


207 m. 

Turbonilla rufa (Philippi, 1836) 

(Figure 269) 


234 m. 

Turbonilla lactea (Linnaeus, 1758) 

(Figure 270) 

Remarks: Regularly present. Ávila, 2000. 

Depth range: Infralittoral to 80 m (MM&B); this 

study: 30-207 m. 

Turbonilla sp. 1 

(Figure 271) 



(Figure 272) 


234 m. 


(Figure 273) 


207 m. 


(Figure 274) 


207 m. 

Family Murchisonellidae Casey, 1905 

Ebala nitidissima (Montagu, 1803) 

(Figures 275-276) 

Remarks: Uncommon. Depth range, this 

study: 57-145 m. 

Subclass OPISTHOBRANCHIA Milne- 

Edwards, 1848 

Order CEPHALASPIDEA Fischer P., 1883 

Family Diaphanidae Odhner, 1914 

Colpodaspis pusilla Sars, 1870 

(Figure 277) 


189 m. 

Family Retusidae Thiele, 1925 

Retusa truncatula (Bruguière, 1792) 

(Figures 278-279) 

Remarks: Common. Mikkelsen, 1995. Depth 

range: below low tide to 200 m (P&G); this 

study: 38-145 m. 

Family Haminoeidae Pilsbry, 1895 

Haminoea cf. orteai Talavera, Murillo & 

Templado, 1987 

(Figure 280) 

Remarks: Uncommon. Mikkelsen, 1995. 


Atys macandrewi E.A. Smith, 1872 

(Figure 281) 

Remarks: Uncommon. Mikkelsen, 1995. 


Atys sp. 

(Figure 282) 


Family Philinidae Gray, 1850 

Philine approximans Dautzenberg & Fisher, 1896 

(Figure 283) 

Remarks: Rare. Depth range, this study: 86-234 m.


Philine sp. 

(Figure 284) 


Family Aglajidae Pilsbry, 1895 

?Chelidonura africana Pruvot-Fol, 1953 

(Figure 285) 


234 m. 

Order THECOSOMATA de Blainville, 1824 

Family Cavoliniidae Gray, 1850 

Cavolinia inflexa (Lesueur, 1813) 

(Figures 286-287) 

Remarks: Common; it is a pelagic species. 

Cavolinia tridentata (Forskal, 1775) 

(Figure 288) 

Remarks: Common; it is a pelagic species. 

Diacria trispinosa (Lesueur, 1821) 

(Figures 289-290) 

Remarks: Rare; it is a pelagic species. 

Cuvierina atlantica (Bé, MacClintock & Currie, 

1972) 

(Figure 291) 


Clio pyramidata (Lesueur, 1821) 

(Figures 292-293) 


Family Limacinidae Gray, 1840 

Limacina cf. helicina (Phipps, 1774) 

(Figure 294) 

Remarks: Rare. Limacina helicina is a boreal, 

pelagic species; the specimens of the Azores, 

however, resemble those reported to this 

species elsewhere (Rolán, 2005). 

Limacina inflata (d’Orbigny, 1836) 

(Figure 295) 


Order NOTASPIDEA Fischer P., 1883 

Family Umbraculidae Dall, 1889 

Umbraculum umbraculum (Lightfoot, 1786) 

(Figure 296) 


52-198 m. 

Family Tylodinidae Gray, 1847 

Tylodina perversa (Gmelin, 1791) 

(Figure 297) 


145 m. 

Subclass PULMONATA Cuvier, 1817 

Superfamily SIPHONARIOIDEA Gray, 1827 

Family Siphonariidae Gray, 1827 

Williamia gussonii (O.G. Costa, 1829) 

(Figure 298) 



Order ARCHAEOPULMONATA 

Superfamily ELLOBIOIDEA Pfeiffer, 1854 

Family Ellobiidae Pfeiffer, 1854 

Ovatella vulcani (Morelet, 1860) 

Remarks: Only one shell; this is a supratidal 

pulmonate. 

Pedipes pedipes Bruguière, 1789 

Remarks: Only one shell; this is a supratidal 

pulmonate. 

Class BIVALVIA Linnaeus, 1758 

Order PTEROMORPHIA Beurlen, 1944 

Superfamily ARCOIDA Stoliczka, 1871 

Family Arcidae Lamarck, 1809 

Arca tetragona Poli, 1795 

(Figure 299) 

Remarks: Common. This species lives on hard 

substrates. Bullock, 1995; Ávila et al., 2000. 

Depth range: intertidal to 900 m (MM&B); low 

tide to 120 m (P&G); this study: 30-360 m. 

Asperarca nodulosa (O.F. Müller, 1776) 

(Figure 300) 

Remarks: Rare. Depth range: low tide to 1000 m 

(P&G); intertidal to 3300 m (MM&B); this 

study: 117-234 m. 

Family Noetiidae Stewart, 1930 

Bathyarca philippiana (Nyst, 1848) 

(Figures 301-302) 

Remarks: Rare. Depth range: around 150 m (P&G); 

60-1200 m (MM&B); this study: 117- 360 m. 

Family Limopsidae Dall, 1895 

Limopsis minuta Philippi, 1836 

(Figure 303) 

Remarks: Rare. Depth range: 40-1400m

68 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XVIII 

299. Arca tetragona Poli, 1795. Loose valves (Sta8) 

300. Asperarca nodulosa (O.F. Müller, 1776). Loose valves (Sta37) 

301. Bathyarca philippiana (Nyst, 1848) (Sta41) 

302. Bathyarca philippiana (Nyst, 1848) (juvenile) (Sta37) 

303. Limopsis minuta Philippi, 1836 (Sta1) 

304. Gregariella semigranata (Reeve, 1858) (Sta28) 

305. Rhomboidella prideauxi (Leach, 1815) (Sta13) 

306. Pecten jacobeus (Linnaeus, 1758) (Sta26) 

307. Pecten jacobeus (Linnaeus, 1758) (juvenile) (Sta18)


70 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XIX 

308. Aequipecten commutatus (Monterosato, 1875) (Sta29) 

309. Aequipecten commutatus (Monterosato, 1875) (Sta23) 

310. Aequipecten opercularis (Linnaeus, 1758) (Sta8) 

311. Aequipecten opercularis (Linnaeus, 1758) (Sta5) 

312. Bractechlamys corallinoides (d’Orbigny, 1840) (Sta26) 



315. Palliolum incomparabile (Risso, 1826) (Sta29) 

316. Palliolum incomparabile (Risso, 1826) (Sta29) 

317. Palliolum incomparabile (Risso, 1826) (Sta7)


72 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XX 

318. Chlamys flexuosa (Poli, 1795) (Sta7) 

319. Talochlamys pusio (Linnaeus, 1758) (Sta26) 

320. Talochlamys pusio (Linnaeus, 1758) (Sta26) 

321. Pododesmus patelliformis (Linnaeus, 1761) (Sta56) 

322. Limaria hians (Gmelin, 1791) (Sta56) 

323. Neopycnodonte cochlear (Poli, 1795) (Sta55) 

324. Myrtea spinifera (Montagu, 1803) (Sta37) 

325. Lucinoma borealis (Linnaeus, 1767) (Sta37) 

326. Lucinoma borealis (Linnaeus, 1767) (Sta37) 

327. Thyasira flexuosa (Montagu, 1803) (Sta7) 

328. Diplodonta berghi (Dautzenberg & Fischer, 1897) (Sta32) 



331. Diplodonta trigona (Scacchi, 1835) (Sta56) 

332. Diplodonta trigona (Scacchi, 1835) (Sta29)


74 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXI 

333. Chama gryphoides (Linnaeus, 1758) (Sta55) 

334. Chama gryphoides (Linnaeus, 1758) (Sta37) 

335. Kurtiella pellucida (Jeffreys, 1881) (Sta27) 

336. Kurtiella pellucida (Jeffreys, 1881) (Sta7) 

337. Basterotia clancula von Cosel, 1995. Loose valves (juvenile) (Sta28) 

338. Basterotia clancula von Cosel, 1995. Loose valves (Sta12) 

339. Cardita calyculata (Linnaeus, 1758) (Sta24)


76 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXII 

340. ?Crassatina sp. (Sta46) 

341. Papillicardium papillosum (Poli, 1791) (Sta47) 






347. Parvicardium vroomi van Aartsen, Menkhorst & Gittenberger, 1984. Loose 

valves (Sta27)


78 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXIII 

348. Tellina incarnata Linnaeus, 1758 (Sta57) 

349. Tellina pygmaea Lovén, 1846 (Sta48) 





354. Arcopagia balaustina (Linnaeus, 1758) (Sta43) 



357. ?Tellina sp. (Sta28)


80 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXIV 

358. Gari costulata (Turton, 1822) (Sta15) 

359. Gari costulata (Turton, 1822) (Sta31) 

360. Ervilia castanea (Montagu, 1803) (Sta26) 

361. Ervilia castanea (Montagu, 1803) (Sta31) 

362. Azorinus chamasolen (da Costa, 1778) (juvenile) (Sta44) 

363. Azorinus chamasolen (da Costa, 1778) (Sta40)


82 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXV 

364. Coralliophaga lithophagella (Lamarck, 1819) (Sta37) 

365. Coralliophaga lithophagella (Lamarck, 1819) (Sta37) 

366. Coralliophaga lithophagella (Lamarck, 1819) (juvenile) (Sta53) 

367. Coralliophaga lithophagella (Lamarck, 1819) (juvenile) (Sta37) 

368. Venus casina Linnaeus, 1758 (Sta54) 

369. Venus verrucosa Linnaeus, 1758 (Sta1) 

370. Venus verrucosa Linnaeus, 1758 (Sta26) 

371. Globivenus effossa (Philippi, 1836) (Sta41)


84 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXVI 

372. Timoclea ovata (Pennant, 1777) (Sta48) 



375. Gouldia minima (Montagu, 1803) (Sta5) 





380. Callista chione (Linnaeus, 1758) (Sta40)


86 AÇOREANA 

2009, Sup. 6: 15-103 

PLATE XXVII 

381. Hiatella arctica (Linnaeus, 1767) (Sta32) 

382. Hiatella arctica (Linnaeus, 1767) (Sta18) 

383. Gastrochaena dubia (Pennant, 1777) (Sta32) 

384. Nototeredo norvagica (Spengler, 1792) (Sta12) 

385. Teredora malleolus (Turton, 1822) (Sta1) 

386. Xyloredo sp. (Sta56) 

387. Xyloredo sp. (Sta56) 

388. Thracia papyracea (Poli, 1791) (Sta13) 

389. Cardiomya costellata (Deshayes, 1835) (Sta23) 

390. Cuspidaria atlantica Allen & Morgan, 1981 (Sta38)


88 AÇOREANA 

2009, Sup. 6: 15-103 

(MM&B); continental shelves down to 900 m 

(P&G); this study: 57 m. 

Superfamily MYTILOIDA de Férussac, 1822 

Family Mytilidae Rafinesque, 1815 

Gregariella semigranata (Reeve, 1858) 

(Figure 304) 

Remarks: Common. Morton, 1995 as Trichomusculus 

semigranatus. This species lives on 

hard substrata. Depth range, this study: 30- 

360m; alive: 66 m. 

Rhomboidella prideauxi (Leach, 1815) 

(Figure 305) 


180 m. 

Superfamily PTERIOIDA Newell, 1965 

Family Pectinidae Rafinesque, 1815 

Pecten jacobeus (Linnaeus, 1758) 

(Figures 306-307) 

Remarks: Uncommon. Depth range: 25-250 m 

(P&G); this study: 40-360 m; alive: 72 m. 

Aequipecten commutatus (Monterosato, 1875) 

(Figures 308-309) 


range: 30-250 m; this study: 40-360 m; alive: 

180 m. 

Aequipecten opercularis (Linnaeus, 1758) 

(Figures 310-311) 


range: low tide to 400 m (P&G; MM&B); this 

study: 38-360 m; alive: 41-171 m. 

Bractechlamys corallinoides (d’Orbigny, 1840 

(Figures 312-314) 

Remarks: Common. Depth range: 3-100 m 

(P&G); this study: 38-360 m. 

Palliolum incomparbile (Risso, 1826) 

(Figures 315-317) 


range: 10-250 m (P&G); infralittoral to 2000 m 

(MM&B); this study: 30-360 m. 

Chlamys flexuosa (Poli, 1795) 

(Figure 318) 

Remarks: Very rare. Ávila et al., 2000. Depth 

range: 30-250 m (P&G); this study: 45-189 m. 

Talochlamys pusio (Linnaeus, 1758) 

(Figures 319-320) 

Remarks: Common. Ávila et al., 2000 as 

Crassadoma pusio. Depth range: from a few 

meters to 150 m (P&G, as Hinnites distortus 

(da Costa, 1778)); 100-2300 (MM&B, as Chlamys 

distorta (da Costa, 1778)); this study: 30- 

360 m. Collected alive at IVFC (Martins, 

2004). 

Family Anomiidae Rafinesque, 1815 

Pododesmus patelliformis (Linnaeus, 1761) 

(Figure 321) 

Remarks: Common. Depth range: intertidal to 

50 m (P&G); intertidal to 450 m (MM&B); this 

study: 14-360 m. 

Family Limidae Rafinesque, 1815 

Limaria hians (Gmelin, 1791) 

(Figure 322) 


range: low tide to 100 m (P&G); low tide to 450 

m (MM&B); this study: 30-360 m. Collected 

alive at IVFC (Martins, 2004). 

Superfamily OSTREOIDA de Férussac, 1822 

Family Gryphaeidae Vyalov, 1936 

Neopycnodonte cochlear (Poli, 1795) 

(Figure 323) 


range: 45-250 m (P&G); this study: 41-360 m; 

alive: 66-162 m. 

Order HETERODONTA Neumayr, 1884 

Superfamily VENEROIDA H. & A. Adams, 

1857 

Family Lucinidae Fleming, 1828 

Myrtea spinifera (Montagu, 1803) 

(Figure 324) 



Lucinoma borealis (Linnaeus, 1767) 

(Figures 325-326) 

Remarks: Common. Bullock, 1995. Range: 

intertidal to 500 m (P&G); intertidal to 1500 m 

(MM&B); this study: 30-360 m; alive: 95-351 m. 

This species was collected alive at IVFC about 

20 cm deep into the sandy substratum (pers. 

obs., AMFM).


Family Thyasiridae Dall, 1900 

Thyasira flexuosa (Montagu, 1803) 

(Figure 327) 


infralittoral to 2190 m (MM&B); this study: 

135-360 m. 

Family Ungulinidae Adams, H. & A., 1857 

Diplodonta berghi (Dautzenberg & Fischer, 

1897) 

(Figures 328-330) 

Remarks: Uncommon. Depth range: 130-1360 

m (MM&B); this study: 38-234 m. 

Diplodonta trigona (Sacchi, 1835) 

(Figures 331-332) 


30-360 m; alive: 30-198 m. 

Family Chamidae Lamarck, 1809 

Chama gryphoides (Linnaeus, 1758) 

(Figures 333-334) 

Remarks: Common. Depth range: intertidal to 

200 m (P&G; MM&B); this study: 41-360 m; 

alive: 162-234 m. 

Family Montacutidae Clark, 1855 

Kurtiella pellucida (Jeffreys, 1881) 

(Figures 335-336) 


40-360 m; alive: 99-207 m. 

Family Sportellidae Dall, 1899 

Basterotia clancula von Cosel, 1995 

(Figures 337-338) 


18-360 m. 

Family Carditidae Fleming, 1828 

Cardita calyculata (Linnaeus, 1758) 

(Figure 339) 

Remarks: Common. Bullock, 1995; Ávila et al., 

2000. Depth range: low tide to 200 m (MM&B; 

P&G); this study: 30-360 m; alive: 38-117 m. 

Alive at IVFC (Martins, 2004). 

Family Crassatellidae Férussac, 1822 

?Crassatina sp. 

(Figure 340) 

Remarks: Rare. One valve, tentatively assigned 

to this genus. Depth range, this study: 

56 m. 

Family Cardiidae Lamarck, 1809 

Papillicardium papillosum (Poli, 1791) 

(Figures 341-346) 

Remarks: Very common. Ávila et al., 2000. 

Depth range: infralittoral to 60 m (MM&B); this 

study: 45-360 m; alive: 45-318 m. 

Parvicardium vroomi van Aartsen, Menkhorst & 

Gittenberger, 1984 

(Figure 347) 



Family Tellinidae de Blainville, 1814 

Tellina incarnata Linnaeus, 1758 

(Figure 348) 


range: intertidal to 85 m (MM&B; P&G); 

this study: 18-360 m; alive: 18-72 m. This 

species was taken alive at IVFC about 20 cm 

deep into the sandy substrata (pers. obs., 

AMFM). 

Tellina pygmaea Lovén, 1846 

(Figures 349-353) 

Remarks: Very common. Depth range: intertidal 

to 150 m; this study: 18-360 m. Collected 

alive throughout its range. Authors report T. 

donacina Linnaeus, 1758 as living in the Azores 

(Ávila et al., 2000); however, all the specimens 

herein collected conform to the representation 

of T. pygmaea instead. 

Arcopagia balaustina Linnaeus, 1758 

(Figures 354-355) 

Remarks: Common. Depth range: infralittoral 

to 750 m (MM&B; P&G); this study: 46-360 m; 

alive: 66-360 m. 

? Tellina sp. 

(Figure 356) 

Remarks: Rare; only one valve, tentatively 

assigned to this genus. Depth range, this 

study: 117-145. 

Family Psammobiidae Fleming, 1828 

Gari costulata (Turton, 1822) 

(Figures 357-359) 


range: infralittoral to 150 m, most abundant 35- 

60 m (MM&B; P&G); this study: 18-360 m; 

alive: 18-66 m.

90 AÇOREANA 

2009, Sup. 6: 15-103 

Family Semelidae Stoliczka, 1870 (1850) 

Ervilia castanea (Montagu, 1803) 

(Figures 360-361) 

Remarks: The commonest bivalve in the Azores 

(Morton, 1990). Bullock, 1995; Ávila et al., 2000. 

Depth range: 30-100 m (P&G); intertidal to 

1800 m (MM&B); this study: 18-360 m; collected 

alive throughout its range. Alive at IVFC 


Family Solecurtidae d’Orbigny, 1846 

Azorinus chamasolen (da Costa, 1778) 

(Figures 362-363) 

Remarks: Rare. Depth range, this study: 38-66 

m; alive: 66 m. 

Family Trapezidae d’Orbigny, 1846 

Coralliophaga lithophagella (Lamarck, 1819) 

(Figures 364-367) 

Remarks: Uncommon. Depth range: 33-200 m 

(MM&B; P&G); this study: 57-360 m; alive: 162- 

234 m. 

Family Veneridae Rafinesque, 1815 

Venus casina Linnaeus, 1758 

(Figure 368) 


range: 5-200 m (P&G); this study: 32-360 m; 

alive: 32-180 m. 

Venus verrucosa Linnaeus, 1758 

(Figures 369-370) 

Remarks: Uncommon. Depth range: intertidal 

to 100 m (MM&B; P&G); this study: 46-360; 

alive: 32-66 m. 

Globivenus effossa (Philippi, 1836) 

(Figure 371) 

Remarks: Rare. Ávila et al., 2000. Depth range: 

50-300 m (P&G); this study: 156-360; taken 

alive throughout the range. 

Timoclea ovata (Pennant, 1777) 

(Figures 372-374) 


Depth range: 4-200 m (P&G); this study: 46-360 

m; alive: 30-360 m. 

Gouldia minima (Montagu, 1803) 

(Figures 375-379) 


Depth range: intertidal to 200 m (MM&B; 

P&G); this study: 18-360 m; alive: 38- 

360 m. 

Callista chione (Linnaeus, 1758) 

(Figure 380) 


range: intertidal to 180 m (MM&B; P&G); this 

study: 18-360 m; alive: 38-81 m. 

Superfamily MYOIDA Stoliczka, 1870 

Family Hiatellidae Gray, 1824 

Hiatella arctica (Linnaeus, 1767) 

(Figures 381-382) 

Remarks: Uncommon; possibly young specimens 

of this species. Ávila et al., 2000. Depth 

range: intertidal to 1400 m (MM&B; P&G); this 

study: 72-207 m. 

Family Gastrochaenidae Gray, 1840 

Gastrochaena dubia (Pennant, 1777) 

(Figure 383) 

Remarks: Rare; one valve tentatively assigned to 

this species. Depth range, this study: 180 m. 

Family Teredinidae Rafinesque, 1815 

Nototeredo norvagica (Spengler, 1792) 

(Figure 384) 

Remarks: Rare. This species lives in drifting 

wood. 

Teredora malleolus (Turton, 1822) 

(Figure 385) 

Remarks: Rare. This species lives in drifting 

wood. 

Family Xylophagidae Purchon, 1941 

Xyloredo sp. 

(Figures 396-387) 

Remarks: Common. This species lives in 

sunken wood. Depth range, this study: 32- 

360 m. 

Order ANOMALODESMATA Dall, 1889 

Superfamily PHOLADOMYOIDA Newell, 

1965 

Family Thraciidae Stoliczka, 1870 

Thracia papyracea (Poli, 1791) 

(Figure 388) 


range: intertidal to 50 m (MM&B; P&G as T. 

phaseolina (Lamarck, 1818)); this study: 30-360 

m; alive: 32-198 m.


Family Cuspidariidae Dall, 1886 

Cardiomya costellata (Deshayes, 1833) 

(Figure 389) 

Remarks: Common. Depth range: 18-200 m 

(P&G); infralittoral to 1400 m (MM&B); this 

study: 40-360 m; alive: 46-198 m. 

Cuspidaria atlantica Allen & Morgan, 1981 

(Figure 390) 


207 m. 

Class CEPHALOPODA Schneider, 1784 

Subclass COLOIDEA Bather, 1788 

Order OCTOPODA Leach, 1818 

Suborder INCIRRATA Grimpe, 1916 

Family Octopodidae d’Orbigny, 1840 

Octopus vulgaris Cuvier, 1797 

(Figure 391) 

Remarks: One very young specimen collected 

alive on a crevice of a dredged rock. Depth 

range, this study: 150 m. 

ACKNOWLEDGEMENTS 

We thank Serge Gofas, Henk Dijsktra, 

Emílio Rolán and Marco Oliverio for their 

kind contributions to the identities of many 

species. Sérgio P. Ávila was supported by 

grant SFRH/BPD/22913/2005 (FCT – 

Fundação para a Ciência e Tecnologia) of the 

Portuguese government. IMAR-DOP/UAc 

(UI&D #531 and ISR LA#9) is funded by 

FCT/MCTES– Lisbon and DRCT/Azores 

through pluri-annual and programmatic 

funding schemes (part FEDER). 

LITERATURE CITED 

ADAM, W., & J. KNUDSEN, 1969. 

Quelques genres de mollusques 

prosobranches marins inconnus ou 

peu connus de l’Afrique 

Occidentale. Bulletin de l’Institut 

royal des Sciences naturelles de 

Belgique, 44(27): 1-69. 

ÁVILA, S.P., 2000. The shallow-water 

Rissoidae (Mollusca, Gastropoda) 

of the Azores and some aspects of 

their ecology. Iberus, 18(2): 51-76. 

ÁVILA, S.P., 2003. The littoral molluscs 

(Gastropoda, Bivalvia and 

Polyplacophora) of São Vicente, 

Capelas (São Miguel Island, 

Azores): ecology and biological 

associations to algae. Iberus, 21(1): 

11-33. 

ÁVILA, S.P., J.M.N. AZEVEDO, J.M. 

GONÇALVES, J. FONTES & F. 

CARDIGOS, 2000. Checklist of the 

shallow-water marine molluscs of 

the Azores: 2 - São Miguel island. 

Açoreana, 9(2): 139-173. 

BOUCHET, P., 1984. Les Triphoridae 

de Mediterranée et le proche 

Atlantique (Mollusca, Gastropoda). 

Lavori Società Italiana di 

Malacologia, 21: 5-58. [not seen] 

BOUCHET, P., & H. GUILLEMOT, 

1978. The Triphora perversa complex 

in Western Europe. Journal of 

Molluscan Studies, 44: 344-356. 

BULLOCK, R.C., 1995. The distribution 

of the molluscan fauna associated 

with the intertidal coralline 

algal turf of a partially submerged 

volcanic crater, the Ilhéu de Vila 

Franca, São Miguel, Azores. In: 

MARTINS, A.M.F. (ed.), The marine 

fauna and flora of the Azores. 

Proceedings of the Second International 

Workshop of Malacology and 

Marine Biology, Vila Franca do 

Campo, São Miguel, Azores. 

Açoreana, Supplement [4]: 9-55. 

BULLOCK, R.C., TURNER, R.D. & 

R.A. FRALICK, 1990. Species richness 

and diversity of algal - associated 

micromolluscan communities 

from São Miguel, Açores. In: 



Proceedings of the First International 

Workshop of Malacology São Miguel, 

Azores. Açoreana, Supplement [2]: 

39-58.

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2009, Sup. 6: 15-103 

CONTRERAS, J.A., 1992. Una nuova 

Gibberula dalle Isole Azzorre 

(Gastropoda, Marginellidae). La 

Conchiglia, 262: 44-45. 

GOFAS, S., 1990. The littoral 

Rissoidae and Anabathridae of São 

Miguel, Azores. In: MARTINS, 

A.M.F. (ed.), The marine fauna and 

flora of the Azores. Proceedings of the 

First International Workshop of 

Malacology São Miguel, Azores. 


HOUBRICK, J., 1990. Anatomy, 

reproductive biology and systematic 

position of Fossarus ambiguous 

(Linné) (Fossarinae: Planaxidae: 

Prosobranchia). In: MARTINS, 

A.M.F. (ed.), The marine fauna and 

flora of the Azores. Proceedings of the 




KNUDSEN, J., 1995. Observations on 

reproductive strategy and zoogeography 

of some marine 

Prosobranch Gastropods (Mollusca) 

from the Azores. In: 








MACEDO, M.C.C., M.I.C. MACEDO 

& J.P. BORGES, 1999. Conchas 

Marinhas de Portugal, 516 pp. 

Editorial Verbo, Lisboa. 

MARTINS, A.M.F., 2004. The Princess’ 

Ring, 99 pp. Intermezzo-Audiovisuais, 

Lda., Lisboa. 

MIKKELSEN, P.M., 1995. Cephalaspids 

of the Azores. In: 



Proceedings of the Second 


and Marine Biology, Vila Franca do 



MORTON, B., 1990. The biology and 

functional morphology of Ervilia 

castanea (Bivalvia: Tellinacea) from 

the Azores. In: MARTINS, A.M.F. 

(ed.), The marine fauna and flora 

of the Azores. Proceedings of the 




MORTON, B, 1995. The biology and 

functional morphology of Trichomusculus 

semigranatus (Bivalvia: 

Mytiloidea) from the Azores. In: 








POPPE, G.T., & Y. GOTO, 1991-1993. 

European Seashells, vol. 1 [1991] 

(Polyplacophora, Caudofoveata, 

Solenogastra, Gastropoda), 352 pp; 

vol. 2 [1993] (Scaphopoda, 

Bivalvia, Cephalopoda), 221 pp. 

Verlag Christa Hemmen, 

Wiesbaden. 

ROLÁN, E., 2005. Malacological Fauna 

from the Cape Verde Archipelago, 455 

pp. ConchBooks, Hackenheim.


FIGURE 391. Octopus vulgaris Cuvier, 1797 (Sta42)

94 AÇOREANA 

2009, Sup. 6: 15-103 

TABLE 1. Distribution of the species dredged during the workshop. See text for station descrip- 

TAXA 

STATIONS 

3 

9 

2 

5 

3 

4 

1 

6 

3 

5 

1 

9 

POLYPLACOPHORA 

Lepidochiton cimicoides 

Acanthochitona fascicularis v v v v v 

4 

5 

5 

7 

5 

8 

3 

6 

4 

0 

0 

6 

0 

5 

1 

4 

1 

5 

2 

0 

3 

1 

4 

6 

0 

1 

5 

6 

3 

0 

4 

8 

GASTROPODA 

Patella candei v v v v v v 

Patella ulyssiponensis v v v v v v v 

Tectura virginea v v v v v v v v v v v v 

Propilidium exiguum 

Emarginula sp. 

Sinezona cingulata 

Haliotis coccinea v v f f v 

? Haliotis sp. 

Lepetella laterocompressa 

Addisonia excentrica 

Clelandella azorica 

v 

Clelandella sp. 

Jujubinus pseudogravinae v v v v v v v v v v v v v v 

Gibbula delgadensis v v v v 

Gibbula magus v + v v + + + + + + v + v v + v 

Margarites sp. 

Solariella azorensis + f f + 

Calliostoma lividum v f v f f 

Calliostoma hirondellei 

Cirsonella gaudryi 

Tricolia pullus azorica f v v v v f v v v v 

Bittium cf. latreillii v v v v v + v v v v v v v v v 

Bittium latreillii 

Fossarus ambiguus v f 

Cheirodonta pallescens v v v v v v 

Similiphora similior v v v v v v v v v 

Marshallora adversa v v v v v v 

Marshallora cf. adversa 

v 

Monophorus sp. v v v 

Monophorus erythrosoma v v v v v 

Monophorus thiriotae v v v v v 

Pogonodon pseudocanaricus 

f 

Strobiligera.brychia 

Metaxia cf. abrupta 

Cerithiopsis tubercularis v v v v 

Cerithiopsis tiara 

Cerithiopsis jeffreysi 

Cerithiopsis scalaris 

v 

Cerithiopsis minima v v v 

Cerithiopsis cf. minima 

v 

Cerithiopsis fayalensis 

Krachia cf. guernei 

Gyroscala lamellosa 

f 

Epitonium turtonis 

Epitonium clathrus 

Epitonium pulchellum 

v 

Epitonium celesti v v 

Punctiscala cerigottana


tion. f, fragment; v, rolled shell or single valve; +, with live animal or both valves. 

4 

4 

5 

0 

5 

4 

1 

8 

4 

7 

4 

9 

1 

3 

2 

7 

2 

3 

2 

4 

2 

1 

1 

2 

0 

2 

2 

8 

4 

2 

2 

6 

0 

8 

3 

2 

0 

7 

2 

9 

3 

8 

3 

3 

3 

7 

4 

3 

5 

2 

5 

3 

5 

1 

4 

1 

5 

5 

v v f v v v 

+ 

f v v v v v f f f 

f v v v v f v v v v v 

v v v v v v v v v v v v v v v v v v 

v 

v 

v 

v v f v v v v f v v 

v 

v v v v v 

v 

v v v v v + v v f v v 

f 

v 

v v v v v v f f f v v v v f v v v v v 

v v v v v v v v 

v v + v v v v v v v v v v v f f v v v f v v + 

v v v 

f v + v v v + f + + f v 

v v f v f v f f v v v 

v v v 

v 

f v f v v v v v v v v v v v 

v v v v v v v v v v v v v v v v v v v v v 

v 

v v v v v 

v v v v v 

v v v v v v v v v v v v 

v v v v v v v v v v v v v 

v 

v 

v 

v v v v v 

v v v v 

v 

v 

v v v 

v 

v 

v 

v v v v 

v v v 

v 

v v v v 

v v 

v 

v 

v 

v v v v 

v v v v v 

v

96 AÇOREANA 

2009, Sup. 6: 15-103 


STATIONS 

TAXA 

Acirsa subdecussata 

Opalia hellenica 

3 

9 

2 

5 

3 

4 

1 

6 

3 

5 

1 

9 

4 

5 

5 

7 

Opalia sp. 1 

Opalia sp. 2 

Melanella bosci 

Melanella cf. crosseana 

Melanella cf. trunca v v 

Parvioris microstoma 

Parvioris sp. 

Crinophteiros collinsi 

Sticteulima jeffreysiana 

Vitreolina sp. 

Vitreolina curva 

v 

Pelseneeria minor 

Littorina striata v f v 

Melarhaphe neritoides 

v 

Skeneopsis planorbis v v 

Rissoa guernei v v v 

Rissoa sp. 1 

Rissoa sp. 2 

Setia subvaricosa 

Setia cf. quisquiliarum 

Crisilla postrema v v v 

Crisilla cf. postrema 

Pseudosetia azorica 

Cingula trifasciata v f 

Manzonia unifasciata v v v v 

Onoba moreleti 

Alvania angioyi v v 

Alvania poucheti v v 

Alvania mediolittoralis v v v 

Alvania punctura 

Alvania sp. (?tarsodes) 

v 

Alvania sleursi v v v v v v 

Alvania cancellata v v v v v v v v v v v 

Alvania platycephala 

Alvania cimicoides 

Alvania cf. cimicoides 

Caecum wayae 

v 

Talassia cf. tenuisculpta 

Capulus ungaricus 

Lamellaria perspicua 

v 

Trivia pulex v v v v v v v v f 

Trivia candidula v v v v v v v v v 

Erato sp. 

Aperiovula juanjosensii 

Notocochlis dillwynii 

Natica prietoi v + v + + + v v + + v v v f v 

Natica cf. prietoi 

v 

Phalium undulatum f f 

Atlanta peronii 

Protatlanta souleyeti 

Ocenebra erinaceus v v 

5 

8 

3 

6 

4 

0 

0 

6 

0 

5 

1 

4 

1 

5 

2 

0 

3 

1 

4 

6 

0 

1 

v 

5 

6 

3 

0 

4 

8 

v


tion. f, fragment; v, rolled shell or single valve; +, with live animal or both valves. (cont.) 

4 

4 

5 

0 

5 

4 

1 

8 

4 

7 

4 

9 

1 

3 

2 

7 

2 

3 

2 

4 

2 

1 

1 

2 

0 

2 

2 

8 

4 

2 

2 

6 

0 

8 

3 

2 

0 

7 

2 

9 

3 

8 

3 

3 

3 

7 

4 

3 

5 

2 

5 

3 

5 

1 

4 

1 

5 

5 

+ + + v v v 

v 

v 

v v v 

v v v 

v 

v 

v 

v 

v 

v 

v v 

v 

f 

f 

v v v 

v v v v v v v v v v 

v 

v v v v 

v v 

v v 

v v v v v v 

v v v 

v 

v 

v v 


v v f v v 

v v v v v v v v v 

v v v v v v v 

v v v v v v 

v 


v v v v v v v v v v v v v v v v 

v v v 

v v + v 

v 

v v v v v 

v 

v 

v v 

v v v 

v v v v v f f f f 

v v v v v v v v v v v f 

v 

v 

f 

+ 

+ v v v + v + v v v f f f f + v v 

f f f 

v 

v 

v v v

98 AÇOREANA 

2009, Sup. 6: 15-103 


STATIONS 

TAXA 

Ocinebrina cf. aciculata 

3 

9 

2 

5 

3 

4 

1 

6 

3 

5 

1 

9 

4 

5 

5 

7 

?Ocinebrina cf. aciculata v v v v 

?Ocinebrina sp. 

v 

Orania fusulus + + v v + 

Trophonopsis barvicensis v v v v v v v 

Trophonopsis cf. muricatus 

v 

Coralliophila cf. meyendorffi v v v v 

Carolliophila panormitana 

Stramonita haemastoma v v v v v 

Gibberula vignali 

Gibberula cf. lazaroi 

Mitra cornea v v v v v 

Pollia dorbignyi 

f 

Nassarius incrassatus f v v v v f v v v v v v 

Nassarius cf. cuvierii 

Nassarius recidivus 

Columbella adansoni v v v v f f f v v f 

Mitrella pallaryi 

Anachis avaroides v v v v v v 

Brocchinia clenchi v v v v v v + 

Mitromorpha azorensis v v v v v v 

Bela nebula v v v + v v v v v v + + v 

Mangelia cf. costata v v v v 

Raphitoma purpurea v v v v 

Raphitoma linearis v v 

Raphitoma cf. aequalis v v v v v v + v v v + v f 

Raphitoma sp. v v v 

Pleurotomella gibbera 

Pleurotomella cf. gibbera 

Teretia teres 

Crassopleura maravignae v v v v v v v v v + v 

Haedropleura septangularis v v v v v v v v 

Philippia krebsi 

v 

Pseudotorinia architae 

Pseudomalaxis zanclaeus 

Mathilda cochlaeformis 

Mathilda retusa 



Omalogyra atomus 

Odostomella doliolum 

Chrysallida cf. flexuosa 

Odostomia bernardi v v v v v 

Odostomia cf. verhoeveni v v v 

Odostomia duureni 

v 

Odostomia cf. striolata 

Eulimella sp. 

Turbonilla rufa 

v 

Turbonilla lactea v v f v v 





5 

8 

3 

6 

4 

0 

v 

0 

6 

0 

5 

1 

4 

1 

5 

2 

0 

3 

1 

4 

6 

0 

1 

5 

6 

3 

0 

4 

8



4 

4 

5 

0 

5 

4 

1 

8 

4 

7 

4 

9 

1 

3 

2 

7 

2 

3 

2 

4 

2 

1 

1 

2 

0 

2 

2 

8 

4 

2 

2 

6 

0 

8 

3 

2 

0 

7 

2 

9 

3 

8 

3 

3 

3 

7 

4 

3 

5 

2 

5 

3 

5 

1 

4 

1 

5 

5 

v v v 

v v v v v 

f v v v v v f f v 

v v v v v v v v v v 

v 

v v v 

v 

v 

v v v v v f v f 

v 

v 

v v v v v 

v f v v f v f v v v v f v f 

v 

f f v f v 

v f v v v v v v v f v v f 

f v f 

v v v v v v v v v v v v v v 

v + v + v v v v v v v v v 

v v v v 

+ + v + + v v v + f + v v + v v v v v v 

+ v v v v + v v v v v 

v f f v 

v v v v v v v v f v v v v 

v v v v v v v v v v v v v v v v v 


v v v v v v 

v 

v v v 

v + v + v v v v v v f v v v v v v + v v v 

v v v v v v 

v 

+ v v v 

v 

f 

v 

v 

v 

v 

v f v 

v 


v v v v v v 

v 

v v v v 

v 

v 

v 

v 

v 

v 

v 

v

100 AÇOREANA 

2009, Sup. 6: 15-103 


TAXA 

STATIONS 

3 

9 

2 

5 

3 

4 

1 

6 

3 

5 

1 

9 

4 

5 

Retusa truncatula v v 

Haminoea cf. orteai v v 

Atys macandrewi 

v 

Atys sp. 

v 

Philine approximans 

Philine sp. 

?Chelidonura africana 

Cavolinia inflexa v v v v v v v v v v 

Cavolinia tridentata 

Diacria trispinosa f v f 

Cuvierina atlantica 

Clio pyramidata v v 

Limacina cf. helicina 

Limacina inflata 

v 

Umbraculum umbraculum 

v 

Tylodina perversa 

Williamia gussonii 

v 

Ovatella vulcani 

Pedipes pedipes 

5 

7 

5 

8 

3 

6 

4 

0 

0 

6 

0 

5 

1 

4 

1 

5 

2 

0 

3 

1 

4 

6 

0 

1 

5 

6 

3 

0 

4 

8 

BIVALVIA 

Arca tetragona v f v v v v v v v v 

Asperarca nodulosa 

Bathyarca philippiana 

Limopsis minuta 

v 

Gregariella semigranata v v v v v v v v v v v v 

Rhomboidella prideauxi 

Pecten jacobaeus f v 

Aequipecten commutatus f v v v 

Aequipecten opercularis v v + + f + + v v f v 

Bractechlamys corallinoides v v v v f v v v 

Palliolum incomparabile f v v f 

Chlamys flexuosa 

v 

Talochlamys pusio v v v v v v v v v v v v 

Pododesmus patelliformis v v v v v v v v v 

Limaria hians v f v v v v v f v 

Neopycnodonte cochlear v v v 

Myrtea spinifera 

Lucinoma borealis v v v v v v 

Thyasira flexuosa 

Diplodonta berghi v v v 

Diplodonta trigona + + + v v v v + v 

Chama gryphoides v v 

Kurtiella pellucida v v 

Basterotia clancula v v v v v v v 

Cardita calyculata v + v v v v v v v 

?Crassatina sp. 

v 

Papillicardium papillosum + + + v + + + v + v + + v 

Parvicardium vroomi v v v 

Tellina incarnata + + + v v v v v v v v v 

Tellina pygmaea + + + + + + + + + + + + + + + 

Arcopagia balaustina 

f 

?Tellina sp.



4 

4 

5 

0 

5 

4 

1 

8 

4 

7 

4 

9 

1 

3 

2 

7 

2 

3 

2 

4 

2 

1 

1 

2 

0 

2 

2 

8 

v v v v v 

v f v 

v v f v v v 

4 

2 

2 

6 

0 

8 

3 

2 

0 

7 

2 

9 

3 

8 

3 

3 

3 

7 

4 

3 

5 

2 

5 

3 

5 

1 

4 

1 

5 

5 

v v v v 

v 

v 

v v v v v v v v v v v v v v v v v v v v 

v f v v 

v v v v f f 

v v v f v v v 

v v v v v v v v v v v v v 

v v v 


v v v f 

v 

v v v v 

v 

v 

v v v v v v v v v v v v v v v v v v v v v 

v 

v v 

+ v v v v v v v v v v f v v v v v v 

v 

v 

+ v f f v f v f 

v v + v v f v v v v v f v 

+ v v + v v v v + v v v + v + v v v v v v v f 

v v v v v v v v v v v v v v v v f v v 

+ v v v v v v + + v v v v v v 

v 

v 

v v v v v v v v v v v v v v v v v v v v 

v v v v v v v v v v v v v v v 

v v v v v v v v v v v v f v f v v f v v 

+ v f v v v v v v v v v + 

v 

v v v v v v v + + v + + + v + + + + v 

v v v v v 

v v v v v v v 

+ v v v v v + v + v + + v v v + v v v v 

v v v + v v v + v v v + 

v + v v v + + v v v v 

v v v v v v v v v v v v 

v + v v + v + v v v v v v v v v v v 

+ + + + + v + + + + + + v v v + v v + v + v v 

v v v v v v v v v v v 

v v v + v v v v v v v v v v v v 

+ + + + + + + + + v + + + v v + v v v + v + v + 

+ v v + + + + + + f + + + + + v + v + 

v

102 AÇOREANA 

2009, Sup. 6: 15-103 


STATIONS 

TAXA 

Azorinus chamasolen 

Coralliophaga lithophagella 

3 

9 

2 

5 

3 

4 

1 

6 

3 

5 

1 

9 

4 

5 

5 

7 

Venus casina + v v v v v v + v + v 

Venus verrucosa + + 

Globivenus effossa 

Timoclea ovata v + + + + + + + + + + + + + 

Gouldia minima v + + + + + + + v + + v 

Callista chione v v + v v v v v v v + v v 

Hiatella arctica 

Gastrochaena dubia 

Nototeredo norvagica 

Teredora malleolus v v 

Xyloredo sp. v v v v v v 

Thracia papyracea v + v + v v v v v v v v 

Cardiomya costellata v v + + + v 

Cuspidaria atlantica 

5 

8 

3 

6 

4 

0 

v 

0 

6 

0 

5 

1 

4 

1 

5 

2 

0 

3 

1 

4 

6 

0 

1 

v 

5 

6 

3 

0 

4 

8 

CEPHALOPODA 

Octopus vulgaris



4 

4 

5 

0 

5 

4 

1 

8 

4 

7 

4 

9 

1 

3 

2 

7 

2 

3 

2 

4 

2 

1 

1 

2 

0 

2 

2 

8 

4 

2 

+ 

v v + v + v v v + 

v v + v v + v v v v + + v v v + v v v v v 

+ v v v v v v v v v 

v v f v + 

+ + + + + + + + + + + + + v v + + + + + + + v + 

v + v + v + + + + v + + + v v + v v + + v + v 

v + + v v + v v v v v v v v v v v v v 

v v v v v v 

v 

v v 

v 

v v v v v v v v v v v v v v v v v v 

v v v + v + v v v v v v v v v + v v v v 

v v + + v + + + + v + v + + v f v f 

v 

2 

6 

0 

8 

3 

2 

0 

7 

2 

9 

3 

8 

3 

3 

3 

7 

4 

3 

5 

2 

5 

3 

5 

1 

4 

1 

5 

5 

+


ASPECTS OF THE BIOLOGY AND FUNCTIONAL MORPHOLOGY OF TIMOCLEA 

OVATA (BIVALVIA: VENEROIDEA: VENERINAE) IN THE AÇORES, PORTUGAL, 

AND A COMPARISON WITH CHIONE ELEVATA (CHIONINAE) 

Brian Morton 


e-mail: prof_bmorton@hotmail.co.uk 

ABSTRACT 

Timoclea ovata occurs in the Açorean offshore seabed down to ~200 metres depth. 

Here, however, it only grows to half (10 mm) the shell length of conspecifics in European 

continental waters. In the Açores also, T. ovata is drilled by a naticid predator - probably 

Natica prietoi. This is the first report of naticid drilling predation upon T. ovata. 

The shell and the organs of the mantle cavity and visceral mass, and their ciliary currents, 

are described (for the first time) and compared specifically with representatives of 

the Chioninae (within which it was traditionally placed), including the Western Pacific 

Bassina calophylla and the Western Atlantic Chione elevata (also illustrated herein). 

Anatomically all three species are similar to each other, reflecting the inherent conservatism 

of the venerid bauplan. An interesting aspect of the complex surface architectures 

of the shells of T. ovata, B. calophylla and C. elevata is that they are not successful in protecting 

individuals from, in particular, naticid predation even though one supposes that 

this is what they are for. That is, in the predator-prey “arms race”, naticids are clearly winning. 

Notwithstanding, the complex shell architecture may also fulfill other functions 

such as stabilizing these shallow burrowers in soft sediments. 

In other bivalve lineages, “success” has been achieved through reproductive and/or 

anatomical specializations. However, the two most widely distributed and, possibly, most 

“successful” modern bivalve lineages are the Mytiloidea and heterodont Veneroida that 

are generally but not exclusively dominant on rocky and soft marine habitats, respectively. 

This success has been achieved by reproductive and anatomical conservatism. Thus, 

one can take virtually any mytilid or any venerid and they will be, as this study demonstrates 

for Timoclea ovata, generally similar to other representatives of their respective families. 

RESUMO 

Timoclea ovata ocorre nos fundos costeiros dos mares dos Açores até uma 

profundidade de ~200 metros. Aqui, porém, cresce apenas até metade (10 mm) do 

comprimento de concha dos seus conspecíficos em águas continentais Europeias. 

Também nos Açores T. ovata é perfurada por um predador naticídeo – provavelmente 

Natica prietoi. Este é o primeiro registo de predação por perfuração de um naticídeo em T. 

ovata. 

Descrevem-se (pela primeira vez) a concha e os órgãos da cavidade palial e massa 

visceral, e as suas correntes ciliares, e comparam-se especificamente com outros 

representantes dos Chioninae, incluindo Bassina calophylla do Pacífico Oeste e Chione 

elevata do Atlântico Oeste (também ilustrada aqui). Anatomicamente as três chioninas são 

muito semelhantes entre si, reflectindo o conservantismo inerente do bauplan venerídeo. 

Um aspecto interessante da complexa arquitectura da superfície da concha chionina, 

incluindo T. ovata, é não ser muito bem sucedida em proteger os seus representantes de, 

em particular, predação por naticídeos embora se pense ser esta a sua função. Isto é, na 

“corrida às armas” de predador-presa, os naticídeos estão claramente a ganhar. Não

106 AÇOREANA 

2009, Sup. 6: 105-119 

(Figure 5B). Jones (1979) compared the 

anatomy of Chione cancellata (Linnaeus, 

1767) with those of other chionines. 

Morton & Knapp (2004) re-examined 

some aspects of the anatomy of the 

Atlantic C. elevata and compared it with 

the Pacific Bassina calophylla, specifically 

with regard to how the shell architecture, 

in particular, protects (or rather does not) 

the contained animal from drilling predators 

of the Naticidae. 

Timoclea ovata has a wide distribution 

from northern Norway and Iceland south 

to Angola (West Africa). It is also recorded 

from the Canary Islands, the Açores 

and the Mediterranean and Black Sea. 

Despite this wide distribution there is, as 

noted above, little known about the 

anatomy of T. ovata and little also about 

its basic biology. Labrune et al. (2007) 

described changes in the species composition 

of the macrofauna of the Bay of 

Banyuls-sur-Mer in the Mediterranean 

between 1967 and 2003 noting that the 

greatest changes were in the T. (as Venus) 

ovata community between 1967-1968 and 

1994. The size frequency distribution of a 

population of T. ovata from the Pliocene of 

Volpedo, Italy, was described by Benigni 

& Corselli (1981) and Dauvin (1985) 

undertook a study of the population 

dynamics of a Recent population of the 

same species from the Bay of Moraix in 

the Mediterranean noting there to be 

pluri-annual variations in recruitment, 

growth and production, thereby explainobstante, 

a arquitectura da concha chionina pode também exercer outras funções tais 

como estabilizar estes escavadores superficiais nos sedimentos finos. 

Noutras linhagens de bivalves, o “sucesso” foi conseguido mediante especializações 

reprodutivas e/ou anatómicas. No entanto, as duas linhagens de bivalves modernos mais 

largamente distribuídas e, possivelmente, melhor sucedidas são os Mytiloidea e os 

heterodontes Veneroida que são geralmente mas não exclusivamente dominantes em 

habitats marinhos rochoso e mole, respectivamente. Tal sucesso foi conseguido mediante 

conservantismo reprodutivo e anatómico. Assim, pode tomar-se virtualmente qualquer 

mitilídeo ou qualquer venerídeo e eles serão, como este estudo demonstra para os 

Chioninae, respectivamente muito semelhantes aos seus outros representantes. 

INTRODUCTION 

Timoclea ovata (Pennant, 1777) was classified 

by Keen (1969) as a member of 

the Chioninae Frizzell, 1936 (Veneroidea, 

Veneridae), representatives of which have 

been studied anatomically by Ansell 

(1961), Jones (1979), Morton (1985) and 

Morton & Knapp (2004). Morton (1985) 

compared Bassina calophylla (Philippi, 

1836) (Chioninae) with Irus irus 

(Linnaeus, 1758) (Tapetinae Adams & 

Adams, 1852). Subsequently, however, 

Coan et al. (1997) synonymized the 

Chioninae with the Venerinae 

Rafinesque, 1815. Kappner & Bieler 

(2006) have, most recently, and on the 

basis of a much broader gene sequencing 

study, argued, however, that the 

Chioninae, with Chione cancellata 

(Linnaeus, 1767) as its type species, is a 

distinct entity from the Venerinae as proposed 

by Canapa et al. (2003). Kappner & 

Bieler (2006), however, have also demonstrated 

that Timoclea Brown, 1827 should 

be placed within the Venerinae and not 

within the Chioninae. 

Ansell (1961) compared the anatomy 

of the British species of Veneracea 

(Veneroidea) but, surprisingly, had little 

to say about Timoclea (as Venus) ovata noting 

only that, in common with Gafrarium 

minimum (Montagu, 1803), the inner apertures 

of the siphons possessed membranes 

that, in the case of the inhalant, 

directed the incoming water dorsally

MORTON: TIMOCLEA OVATA IN THE AZORES 107 

ing the observations of Labrune et al. 

(2007). Anfossi & Brambilla (1981) noted 

that T. ovata has been a member of the 

Mediterranean’s detrital biocenosis since 

at least the Pleistocene. In terms of predation, 

only Mienis (2003) has noted that 

the starfish Astropecten aranciacus 

(Linnaeus, 1758) preys upon T. ovata. 

This study was the first to be undertaken 

on Timoclea ovata in the Açores, a 

species initially recorded from there by 

Morton (1967). The study’s aims were 

three fold: (i), to obtain support for (or 

not) the proposal of Kappner & Bieler 

(2006) that T. ovata should be placed in the 

Venerinae; (ii), to document information 

on the biology of this species in the 

remote mid-Atlantic Açores and (iii), to 

provide a picture of its anatomy that 

might explain facets of its biology and 

give clues to the success of the venerid 

bauplan. 

MATERIALS AND METHODS 

Biology 

For ten days from 17-26 July 2006, the 

sea bed off the southern coast of the 

island of São Miguel, Açores, was 

sampled using a benthic box dredge at six 

stations to the east and west of Ilhéu de 

Vila Franca do Campo. Station details are 

described by Martins et al. (2009). Station 

depths ranged from –50 to -250 metres 

C.D.. All living and empty shells of 

Timoclea ovata (plus any living naticids) 

were sorted from the samples. These 

were analyzed in the following manner. 

Living individuals of Timoclea ovata 

were measured along their greatest 

lengths using vernier calipers to the nearest 

0.5 mm. Empty shell valves were 

identified and both left and right ones 

were measured in the same manner. 

Empty valves were also examined for 

drill holes. Where these were encountered, 

the following records were made 

of: (i), which valve and (ii), the location of 

each drill hole was plotted on master 

illustrations of the left and right valves. 

Statistical analyses 

The dataset comprising the numbers 

of living, empty and drilled shell valves of 

Timoclea ovata among the six stations was 

tested for normality and homogeneity of 

variances using the Shapiro-Wilk test and 

Levene statistic, respectively, both at the 

p = 0.05 level of significance before 

ANOVA. One-way ANOVA’s were performed 

on the dataset to test the null 

hypothesis that there were no significant 

differences in these variables among locations. 

Where differences were detected, 

Student’s Newman-Keuls (SNK) tests 

were carried out to identify where the differences 

lay. The shell lengths of living T. 

ovata and empty and drilled valves were 

also compared using a one-way ANOVA 

and the Student’s Newman-Keuls (SNK) 

test. 

Anatomy 


were observed alive in aquaria. Other living 

animals were dissected and drawings 

made of the anatomy, notably the organs 

of the mantle cavity. The ciliary currents 

in the mantle cavity were detected using a 

suspension of carmine in seawater. 

Living individuals of Chione elevata from 

Florida were also examined in the same 

way. 

RESULTS 

Biology 


were collected from all six stations and 

there was no significant difference in the 

numbers collected between them. Figure 

1 illustrates length frequency histograms 

for A, living individuals, B and C, empty 

left right shell valves, respectively, and D

108 AÇOREANA 

2009, Sup. 6: 105-119 

and E, drilled left and right shell valves, 

again respectively. Living shells ranged 

in shell length from 1.5 mm to 9.0 mm, as 

did, approximately, the empty and drilled 

valves. One empty right valve was 10 mm 

in length. 

shell length showed that the valves of 

empty and drilled individuals, irrespective 

of whether they were the right or left 

valves, did not significantly differ in 

terms of mean shell length (p >0.05) 

whereas the valves of living individuals 

were significantly smaller than the empty 

valves (except for the drilled left valves, 

probably due to the small sample size). 

Figure 2 shows outline drawings of 

the left and right shell valves and the 

numbers and the distribution pattern of 

naticid drill holes made in them. There 

were over twice as many in the right 

(n=33) as in the left (n=15) valves and the 

pattern of distribution was unusual in 

that they were not near the umbones (an 

often favoured location for drilling gastropod 

predators, as will be discussed). 

Rather, they seemed to be located mostly 

over the position of the pallial line, that is, 

where the thick, recessed, mantle edge is 

united to the shell internally. 

FIGURE 1. Timoclea ovata. Five histograms 

showing the size distributions of A, living individuals; 

B, empty left and C, right shell valves; 

C, and D, drilled left and right shell valves, 

respectively. 

The results of the ANOVA show that 

there were significant differences 

between the collected Timoclea ovata shells 

of the various categories, that is, living, 

empty and drilled valves, in term of their 

mean shell lengths (F = 6.96, p


posteriorly, so that the beaks are distinctly 

prosogyrate (Figure 3A) and result 

from a strong tangential pattern of 

growth. This gives the shell its equilateral 

shape that increases during ontogeny 

so that the juvenile is more oval, the adult 

more angular. 

FIGURE 3. Timoclea ovata. Five stylized views 

of the shell and illustrated from A, the right 

side; B, dorsally; C, ventrally; D, anteriorly and 

E, posteriorly. (x—-y is the approximate position 

of the greatest width to the shell.) For 

other abbreviations see page 119. 

In dorsal view (Figure 3B), there is a 

small anterior, heart-shaped, lunule (LU) 

(as defined by Carter, 1967) each valve 

here also interlocking by means of marginal 

denticles as in Chione elevata 

(Morton & Knapp, 2004). Unlike other 

venerids, however, the lunule is not well 

defined because it is sculptured like the 

remainder of the shell but is defined by a 

light indentation and is coloured slightly 

differently. The more oval juvenile shell 

(JS) is also illustrated. There is no distinct 

posterior escutcheon because the ligament 

is internal, although it can be seen 

as a thin, black, line extending posterior 

to the umbones about one quarter of the 

way towards the posterior margin. The 

ventral shell valve margins (Figure 3C) 

are interlocked virtually everywhere 

along the extent of the shell margin by the 

expanding radial ribs. The shell valves of 

T. ovata are thus very difficult to separate, 

again like those of C. elevata (Morton & 

Knapp, 2004). Figures 3B & C also show 

the approximate position of the greatest 

shell width (x-y). It lies just posterior to 

the umbones above the ligament. 

The shell is illustrated in anterior view 

in Figure 3D and again shows the lunule 

(LU) and juvenile shell (JS). The greatest 

width to the shell when seen from this 

perspective (x-y) is dorsal to the mid 

point of the dorso-ventral height of the 

shell. A similar situation is seen when the 

shell is illustrated from the posterior perspective 

(Figure 3E). 

The shell of Timoclea ovata is illustrated 

in internal view in Figure 4A. The left 

valve (Figure 4A) has a hinge plate with 

an elongate posterior cardinal tooth and 

two other larger teeth more centrally 

placed. These are here interpreted as a 

central cardinal and robust anterior (but 

more centrally located, unlike the elongate 

posterior cardinal tooth) cardinal 

tooth, all arising from the umbo (U). 

There are no lateral teeth. The ligament 

(L) is internal and opisthodetic. Also well 

defined are anterior (AA) and a larger 

posterior (PA) adductor muscle scars. 

These are connected by a thick pallial line 

(PL) that has a similarly thick, short, pallial 

sinus (PS). Both are deeply inset from 

the valve margin and this is characterized 

by a scalloped edge internal to which, 

extending virtually all the way round are 

marginal denticles (MD) that interlock 

with those of the other valve. The interi-

110 AÇOREANA 

2009, Sup. 6: 105-119 

or of the shell is polished and is typically 

white although in those individuals with 

an external coloration, the interior reflects 

this as an orange-lilac stain. The hinge 

plate of the right valve similarly also has 

three (a central cardinal, elongate posterior 

lateral and more central and robust 

anterior lateral) teeth (Figure 4B). 

FIGURE 4. Timoclea ovata. A, An interior view 

of the left shell valve and B, an interior view of 

the right hinge plate. For abbreviations see 

page 119. 

FIGURE 5. Timoclea ovata. An individual illustrated 

in its life position in the sediment. 

The external appearance of the shell is 

illustrated from the right side in Figure 5. 

Each valve has a stout sculpture of ~50 

ribs that radiate from the umbones and 

because each valve has strong commarginal 

lamellae too, there are nodes on the 

ribs giving the shell a rough, file like, feel 

to the touch. As noted above the shell is 

often uniformly coloured light brown 

although those of most individuals are 

patterned with streaks and blotches of 

pink, red or brown. And some individuals 

have two radiating (antero- and postero-ventrally) 

bands of pigmentation, as 

illustrated in Figure 5. 

Tebble (1966) records that Timoclea 

ovata has a maximum shell length of 19 

mm while Dauvin (1985) identified a figure 

of 15.1 mm for this species in the 

Mediterranean. As noted above, however, 

the largest Açorean individual 

collected had a shell length of but 10 mm. 

The siphons 

living individual of Timoclea ovata is 

illustrated in Figure 5 from the right side. 

Anteriorly, there is a large digging foot. 

Posteriorly, there is a pair of separated 

siphons. The exhalant siphon is conical 

and a ring of 12 short tentacles sub-apically 

surrounds its transparent coneshaped 

aperture. About 12 yellow-brown 

stripes arise from between each tentacle 

and extend inwards. The inhalant siphon 

is much larger in diameter and is fringed 

apically by a circlet of ~24 long siphonal 

tentacles. As with the exhalant about 24 

yellow-brown stripes extend inwards. 

Where each stripe unites with the tentacular 

rings, there is a darker brown spot.


Mid-ventrally, the mantle possesses a line 

of papillae and pallial fusions, where they 

occur, are of the inner folds only, that is, 

type A (Yonge, 1982). The siphons are 

illustrated in greater detail in Figure 6. 

FIGURE 7. Timoclea ovata. The ciliary currents 

of the left mantle lobe after after removal of the 

left shell valve and mantle and the body. For 

abbreviations see page 119. 

FIGURE 6. Timoclea ovata. A detail of the 

siphons. 

The ciliary currents of the organs of the mantle 

cavity 

The ciliary currents of the left mantle 

lobe of Timoclea ovata are illustrated in 

Figure 7. The pallial currents sweep particles 

of material in a clockwise direction 

towards the antero-dorsal regions of the 

mantle cavity and then downwards, so 

that particles end up in a ventral marginal 

rejection tract that transports such 

unwanted material towards the base of 

the inhalant siphon (IS) where it accumulates 

as balls of pseudofaecal matter (PM). 

These little balls are periodically ejected 

from the mantle cavity, via the inhalant 

siphon, by sharp contractions of the 

adductor muscles that create the pallial 

pressure necessary to do so. 

The right ctenidium of Timoclea ovata 

is illustrated in Figure 8A. Each ctenidium 

is homorhabdic, eulamellibranchiate 

and comprises two unequal demibranchs. 

The inner demibranch (ID) is large and 

extends anteriorly from beneath the posterior 

adductor muscle (PA) up into the 

sub-umbonal cavity and ends on the postero-ventral 

face of the anterior adductor 

muscle (AA). The outer demibranch 

(OD) is dorso-ventrally short and and, as 

with the inner, extends anteriorly from 

beneath the posterior adductor muscle to 

a position just posterior of the hinge plate, 

beneath the ligament (L). This demibranch 

is thus foreshortened anteriorly. 

The ciliary currents of the right ctenidium 

are illustrated in transverse section 

in Figure 8B. The ciliary currents are of 

Type C (1) (Atkins, 1937), typical of many 

eulamellibranchs and, specifically, Venus 

fasciata (da Costa, 1778), Dosinia lupinus 

(Linnaeus, 1758), Venerupis aurea (Gmelin, 

1791) and Venerupis rhomboides (Pennant, 

1777) (Ansell, 1961). Ctenidia with a ciliation 

of type C (1) have oralward acceptance 

tracts located in the ctenidial axis 

and in the ventral marginal food groove 

of the inner demibranch (ID) only. Hence, 

particles filtered by the ascending lamella 

of the outer demibranch (OD) pass downward 

(although some may dorsally be 

carried anteriorly) (Figure 8A) and on 

reaching the ventral margin of this demi-

112 AÇOREANA 

2009, Sup. 6: 105-119 

FIGURE 8. Timoclea ovata. A, The ciliary currents 

of the ctenidium as seen from the right 

side after removal of the right shell valve 

and mantle. B, A diagrammatic transverse 

section through the right ctenidium showing 

the ciliary currents. For abbreviations see 

page 119. 

branch, turn, and pass upwards on the 

descending lamella towards the ctenidial 

axis. In the food groove of the ctenidial 

axis they pass anteriorly. Ciliary currents 

on the inner demibranch are largely 

downward towards the ventral margin 

that has an anteriorly directed food 

groove. Because the outer demibranch is 

anteriorly foreshortened, at its anterior 

terminus, particles arriving here in the 

ctenidial axis pass on to the descending 

lamella of the inner demibranch. Thus, 

particles arrive at the ctenidial labial palp 

junction in (i), the ventral marginal food 

groove of the inner demibranch and (ii), 

in the ctenidial axis tract, also of the inner 

demibranch. 

The ctenidial-labial palp junction is of 

Category II (Stasek, 1964) and the palps 

(LP) are small, each possessing no more 

than five pleats that would, in the typical 

bivalve, have a sorting function, either 

accepting or rejecting particles of possible 

food according to size. Only those particles 

passing directly into the oral grooves 

of the palps from the acceptance tracts in 

the ctenidial axes, an arrangement that 

characterizes bivalves with a 

ctenidial/labial palp junction of Category 

II (Stasek, 1964), are forwarded directly to 

the mouth. The reduced size and sorting 

ability of the labial palps may be a reflection 

of the low numbers and narrow size 

limits of particles in the Açorean waters of 

the mid-Atlantic. Similarly, small labial 

palps have been recorded for Fragum erugatum 

(Tate, 1889), an inhabitant of oligotrophic, 

high salinity waters in Australia 

(Morton, 2000). Throughout its wide geographical 

range, however, T. ovata may 

occur in a variety of sediment types 

although it seems to prefer well-sorted 

gravels, as in the Açores, in which case 

the small labial palps are an adaptation to 

sediments and overlying waters low in 

particulates. 

The ciliary currents of the right side of 

the visceral mass of Timoclea ovata are 

illustrated in Figure 9. On the surface of 

the right side of the visceral mass, the ciliary 

currents move material in a clockwise 

direction, anteriorly above and posteriorly 

below. Eventually, all such 

currents become directed downwards 

and feed into a rejection tract that passes 

accumulated material posteriorly where 

it falls off the posterior edge of the visceral 

mass onto the mantle. Ciliary currents 

on the foot (F) also pass material into this 

rejection tract. Particles that fall off the 

visceral mass are subjected to the ciliary 

currents of the mantle (Figure 7). 

Also illustrated in Figure 9 are some 

details of the structure of the visceral 

mass. Below and just posterior to the 

hinge plate, below the ligament (L), there 

is a heart (H) and posterior to this the 

brown, paired, kidneys (K). Anterior to 

the hinge plate is the dark brown digestive 

diverticula (DD). The gut has not


FIGURE 9. Timoclea ovata. The ciliary currents of the right side of the visceral mass after removal 

of the right shell valve, mantle and ctenidium. Also illustrated are some details of the structure of 

the visceral mass. For abbreviations see page 119. 

been examined in detail, but a conjoined 

style sac and mid gut (CSS/MG) leaves 

the postero-ventral edge of the stomach 

and passes postero-ventrally into the visceral 

mass. Eventually, the extremely thin 

mid gut (MG) separates from this and 

makes a single, simple loop upwards 

back towards the stomach but then passes 

posteriorly, penetrating the ventricle of 

the heart as the rectum (R) which passes 

over the posterior adductor muscle (PA) 

to terminate on its posterior surface as an 

anus. 

A final anatomical point is that, surprisingly, 

the pedal retractor muscles are 

minute. The posterior pedal retractor 

muscle (Figure 9, PPR) is located next to 

the antero-dorsal edge of the posterior 

adductor (PA) while the anterior pedal 

retractor muscle (APR) is even smaller 

and located on the postero-dorsal edge of 

the anterior adductor muscle (AA). I use 

the word ‘surprisingly’ above, because, 

despite these tiny muscles, the foot is further 

surprisingly, very active. Hence, its 

movements must be largely related to 

hydrostatically induced pressure 

changes, as will be discussed. 

Comparison with Chione elevata 

Aspects of the anatomy and the ciliary 

currents of the ctenidium of Chione elevata 

are illustrated in Figure 10 and as seen 

from the right side after removal of the 

right shell valve and mantle. This drawing 

should be compared with the corresponding 

one for Timoclea ovata (Figure 8). 

In C. elevata, the shell has a lunule (L) 

beneath which are located interlocking 

denticles. The shell also has interlocking

114 AÇOREANA 

2009, Sup. 6: 105-119 

valve margins and there are three hinge 

teeth in each valve, that is, what is here 

interpreted as anterior (ALT) and posterior 

lateral (PLT) and a large central cardinal 

teeth (CT). Jones (1979) also identifies 

three teeth in each valve of Chione cancellata 

(Linnaeus, 1767), C. paphia (Linnaeus, 

1767) and C. undatella (Sowerby, 1835) but 

refers to them as anterior, central and posterior 

cardinal teeth, as decribed above 

for Timoclea ovata. 

Internally, Chione elevata has anterior 

(AA) and posterior (PA) adductor muscles 

and anterior (APR) and posterior 

(PPR) pedal retractor muscles that are 

larger than those of Timoclea ovata. There 

is an extensive pedal gape (PG) as in T. 

ovata, a large digging foot (F) and a mantle 

margin (MM) lined with mantle papillae 

(MP), all, again, as in T. ovata. The 

inhalant (IS) and exhalant (ES) siphons of 

C. elevata are very similar in structure to 

those of T. ovata and both have similarly 

organized ctenidia with ciliary currents of 

Type C (1) (Atkins, 1937). Both, therefore, 

have ctenidial-labial palp junction of 

Category II (Stasek, 1964). The labial 

palps of C. elevata are relatively larger 

than those of T. ovata, possibly because it 

inhabits shallow coastal waters off 

Florida, U.S.A.. 

In conclusion therefore Chione elevata 

and Timoclea ovata are similar anatomically, 

even though gene sequencing places 

FIGURE 10. Chione elevata. The ciliary currents of the ctenidium as seen from the right side after 

removal of the right shell valve and mantle. For abbreviations see page 119.


them in two different sub-families, that is, 

the Chioninae and Venerinae, respectively 

(Kappner & Bieler, 2006). Importantly, 

such similarities now suggest convergence 

between the two thick-shelled and 

surface ornamented genera demonstrating 

the success, at least in part, of the veneroidean 

body plan as a whole and which 

seems to be based around anatomical 

conservatism. 

DISCUSSION 

Poppe & Gotto (1993) record a depth 

distribution of between 4 m to 200 m for 

Timoclea ovata, as does Tebble (1966), 

approximately. It is thus unsurprising that 

no differences in population numbers 

were recorded with depth from the 

Açorean dredge samples (50-200 m). 

Poppe & Gotto (1993) similarly note that T. 

ovata individuals from the northern part of 

the species’ range are larger than southern 

conspecifics. This may explain why the T. 

ovata individuals from Açorean waters are 

small (half the length of northern conspecifics), 

but it may also be because of the 

low levels of nutrients available to this 

species in the depauperate waters of the 

central Atlantic. 

Very little is known about the predator-prey 

relationships of the Açorean 

marine fauna (Morton et al., 1998). In 

terms of the predatory gastropods, on 

rocky shores in the Açores, Thais haemastoma 

(Linnaeus, 1767) drills the intertidal 

mussel Gregariella semigranata (Reeve, 

1858) at the posterior margin (Morton, 

1995a). In Europe, Ansell (1960, 1982) 

demonstrated that Polinices alderi (Forbes, 

1838) drilled the bivalves Venus striatula da 

Costa, 1778 and Tellina tenuis (da Costa, 

1778) whereas in the Açores, P. alderi 

attacked the commonest shallow subtidal 

bivalve, Ervilia castanea (Montagu, 1803), 

by drilling in a stereotypical, posterior, 

position (Morton, 1990a). 

Morton & Harper (2009) have studied 

the drill holes made in the tubes of the 

serpulid polychaete Ditrupa arietina (O.F. 

Müller, 1776) from depths of 50-200 

metres in the Açores and concluded that 

the only predator present in the samples 

from which the tubes were collected was 

Natica prietoi Hidalgo, 1873, formerly 

identified as Natica adansoni de Blainville, 

1825. Since the specimens of Timoclea 

ovata reported upon here came from the 

same dredge samples it seems possible 

(likely) that N. prietoi made the holes in 

the shell of this species too. Confirmation 

of this is, however, required. 

Notwithstanding, Kabat (1990) has 

reviewed the literature on naticid predation 

and there are no records of Timoclea 

ovata as prey. This is thus the first record 

of naticid predation, possibly by N. prietoi 

on T. ovata, although Vermeij (1980) 

reports that Timoclea marica (Linnaeus, 

1758) is drilled (laterally, as with T. ovata) 

by an unknown gastropod in Guam. 

Morton & Knapp (2004) identified an 

almost equal distribution of drill holes 

(and attempts) between the two valves of 

Chione elevata. Most of the drill holes 

were distributed around the postero-dorsal 

region of the shell and there were few 

failed drill holes. Finally, only a very few 

of the drill holes were over the shell 

lamellae and were at inter-lamellar 

spaces. That is, if the lamellae have 

evolved as anti-predation devices, they 

are not very successful, in this case from 

the naticid Naticarius canrena (Linnaeus, 

1758). This is not the case with the also 

heavily sculptured Bassina calophylla 

(Chioninae) in the Indo-West Pacific 

(Ansell & Morton, 1985; 1987), where the 

shell lamellae do protect the bivalve 

inhabitant, except from species of edge 

drilling naticids, that is, Polinices tumidus 

(Swainson, 1844) and Polinices melanostomus 

(Gmelin, 1791). 

Timoclea ovata has a shell that, superfi-

116 AÇOREANA 

2009, Sup. 6: 105-119 

cially, would appear to offer much protection 

against drilling predators. Protective 

characteristics include a relatively stout 

shell with tightly fitting valve margins, 

ventrally interlocking ribs, similarly interlocking 

denticles that occur all around the 

valve margins and large hinge teeth. 

Each adductor muscle is also large, facilitating 

sustained adduction and the pallial 

line is deeply inset within the shell margin. 

A thick shell characteristically protects 

bivalves from drilling predators, for 

example, Corbula crassa Hinds, 1843 in 

Hong Kong (Morton, 1990b), although 

Borzone (1988) showed that a species of 

Polinices, as demonstrated here for N. prietoi 

Hidalgo, 1873 and T. ovata, selectively 

drilled its prey, Venus antiqua King & 

Broderip, 1831, in the thickest region of 

the shell, that is, umbonally. Natica catena 

(da Costa, 1778) similarly bores its prey, 

the subtidal Donax vittatus (da Costa, 

1778) around the umbones (Negus, 1975). 

The anatomies of various representatives 

of the Chioninae have been 

described. These include Bassina calophylla 

(Morton, 1985) and Chione elevata (formerly 

identified as C. cancellata) (Jones, 

1979; Morton & Knapp, 2004) while 

Ansell (1961) has described the anatomies 

of the representatives of the Veneridae 

(including a little about Timoclea ovata) 

that occur in British waters. The most 

obvious feature of the studied representatives 

of the Veneridae is their anatomical 

conservatism. Hence, illustrations of the 

ctenidia within the mantle cavity of T. 

ovata (Venerinae), and C. elevata (illustrated 

herein: Figs 8 & 10, respectively) and B. 

calophylla (Morton, 1985, fig. 10) 

(Chioninae) are virtually identical, differing 

only in labial palp size. It is well 

known that palp size in the Bivalvia is 

related to the degree of sorting necessary 

for the particle load in the inhalant 

stream. Thus, the Hong Kong continental 

shelf species B. calophylla has large palps, 

the Floridian C. elevata has intermediate 

sized palps (Figure 10) while T. ovata has 

tiny palps (Figure 8). That is, because of 

high nutrient loading in continental shelf 

waters, B. calophylla needs big palps to 

reject a surfeit of unwanted particles 

whereas, oppositely, T. ovata in mid- 

Atlantic waters has little need to reject 

anything and has tiny palps. Timoclea 

ovata has also, for the same reason, a 

short, narrow, mid-gut. 

Morton (1995b) pointed out that the 

two most widely distributed and, possibly, 

most “successful” modern bivalve 

lineages are the Mytiloidea and heterodont 

Veneroida that are generally but 

not exclusively dominant on rocky and 

soft marine habitats, respectively. This 

has been achieved by reproductive (virtually 

all representatives being broadcast 

spawners) and anatomical conservatism. 

Thus, one can take virtually any mytilid 

or any venerid and they will be, as this 

study demonstrates for Timoclea ovata, 

similar to the other representatives of 

each order. In other bivalve lineages, 

more limited “success” has been achieved 

through reproductive and/or anatomical 

specialisms (Morton, 1995b). But the true 

inheritors of the bivalve bauplan are the 

modern Mytiloidea and Veneroidea. 

One interesting aspect of this “success”, 

however, is that, as discussed here, 

the complex surface architectures of the 

shell of the chionine Chione elevata and the 

venerine Timoclea ovata have not been successful 

in protecting representatives from, 

in particular, naticid predation even 

though one instinctively supposes that 

that is what it is for. That is, in the predator-prey 

“arms race”, naticids are clearly 

winning but the chionine shell architecture 

may also fulfill other functions such 

as the stability of the shallow burrowing 

Timoclea ovata, and other chionines, in soft 

sediments – a habitat that their shell 

architectures suit them ideally to.



I am grateful to Prof. A.M. Frias Martins 

(University of the Açores) for funding this 

research and for much practical help and 

warm hospitality during my stay on São 

Miguel. I also thank a number of University 

of the Açores students who helped with 

sample sorting and Dr. K.F. Leung (Hong 

Kong) for statistical advice. 


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102-103. 

ABBREVIATIONS USED IN THE 

FIGURES 

AA Anterior adductor muscle or 

scar 

ALT Anterior lateral tooth 

APR Anterior pedal retractor muscle 

AU Auricle (of heart) 

CSS/MG Conjoined style sac and mid 

gut 

CT Cardinal tooth 

DD Digestive diverticula 

ES Exhalant siphon 

F Foot 

H Heart 

ID Inner demibranch 

ILP Inner labial palp 

IS Inhalant siphon 

JS Juvenile shell 

K Kidney 

L Ligament 

LP Labial palp 

LU Lunule 

MD Marginal denticles 

MG Mid gut 

MM Mantle margin 

MP Mantle papillae 

OD Outer demibranch 

OLP Outer labial palp 

PA Posterior adductor muscle or 

scar 

PG Pedal gape 

PL Pallial line 

PLT Posterior lateral tooth 

PM Pseudofaecal mass 

PPR Posterior pedal retractor 

muscle 

PS Pallial sinus 

R Rectum 

U Umbo 

V Ventricle (of heart)


ANATOMY AND BIOLOGY OF MITRA CORNEA LAMARCK, 1811 (MOLLUSCA, 

CAENOGASTROPODA, MITRIDAE) FROM THE AZORES 

M. G. Harasewych 

Dept. of Invertebrate Zoology, MRC-163, National Museum of Natural History, 

Smithsonian Institution, PO Box 37012, Washington, D.C. 20013-7012. USA. 

e-mail: Harasewych@si.edu 

ABSTRACT 

Mitra cornea Lamarck, 1811, a member of the taxonomically complex group of small 

brown miters, is described anatomically, including observations on shell ultrastructure 

and diet. Morphological features confirm its taxonomic placement within the genus Mitra, 

and indicate a closer relationship with the western African Mitra nigra than with the 

Mediterranean Mitra cornicula. Mitra cornea shares the morphological adaptations of the 

anterior alimentary system that have evolved in conjunction with a specialized sipunculan 

diet, and that appear to be fairly uniform within the Mitrinae. Studies on the composition 

and pharmacological effects of the secretions of the salivary glands and hypobranchial 

gland are needed to better interpret the origin and evolutionary pathways that gave rise 

to the extreme trophic specialization of the Mitridae. 

RESUMO 

Descreve-se anatomicamente Mitra cornea Lamarck, 1811, um membro do grupo 

taxonomicamente complexo de pequenas mitras castanhas, incluindo-se observações 

sobre a estrutura da concha e sobre a dieta. As características morfológicas confirmam a 

sua localização taxonómica no género Mitra e indicam um relacionamento mais chegado 

com Mitra nigra, da África ocidental, do que com a mediterrânea Mitra cornicula. Mitra 

cornea possui as adaptações morfológicas da porção anterior do sistema alimentar que 

evoluíram juntamente com uma dieta especializada de sipúnculos, e que parece ser 

bastante uniforme dentro dos Mitrinae. São necessários estudos sobre a composição e 

efeitos farmacológicos das secreções das glândulas salivares e da glândula hipobranquial 

para melhor se interpretar a origem e os percursos evolutivos que fizeram aparecer a 

especialização trófica extrema dos Mitridae. 

INTRODUCTION 

The Mitridae comprise a diverse and 

cosmopolitan family of predatory 

neogastropods that are common in the 

tropics, but are also present in temperate 

seas. Mitrids usually occur at intertidal 

to subtidal depths, but extend into the 

bathyal zone (Cernohorsky, 1976; 

Ponder, 1998). Taylor (1993) noted that 

Mitridae are perhaps the most trophically 

specialized family of Neogastropoda, 

as all species studied to date have been 

found to feed exclusively on Sipuncula 

(e.g., Kohn, 1970; Fukuyama & 

Nybakken, 1983; Loch, 1987; Taylor, 

1989; 1993). 

The mitrid fauna of the Azores is not 

diverse, with the family represented by 

two Recent species. Mitra zonata Marryat, 

1818, is easily distinguished by its large 

size and two-toned periostracum that is 

lighter in color along the apical portion of 

the shell above the suture. Cernohorsky 

(1976:367) regarded this Recent taxon to 

be a subspecies of the Pliocene Mitra 

(Mitra) fusiformis (Brocchi, 1814). He 

reported it to be limited to the western 

Mediterranean and northwestern Africa, 

but it has since been documented to occur

122 AÇOREANA 

2009, Sup. 6:121-135 

in the Azores by Burnay & Martins (1988). 

Martins (2004:94) noted that this species 

lives on offshore, sandy bottoms. 

A smaller species that inhabits the 

intertidal and subtidal rocky substrates of 

the Azores has a long and complicated 

taxonomic history. Multiple names have 

been applied to various phenotypes of 

small, smooth, dark brown mitrids that 

inhabit the Mediterranean and eastern 

Atlantic. Rolán et al. (1997) reviewed the 

systematics of the “dark brown 

Mitridae”, concluded that three morphologically 

similar species are involved, and 

apportioned the multiple available names 

among them. According to these authors, 

Mitra cornicula (Linnaeus, 1758) is endemic 

to the Mediterranean Sea, Mitra 

nigra (Gmelin, 1791) ranges from the Cape 

Verde Islands to Angola, while the broad 

ranging Mitra cornea Lamarck, 1811, overlaps 

the range of the other two, occurring 

in the western Mediterranean Sea, the 

Macaronesian Archipelagos (Azores, 

Canaries, and Cape Verde Islands) and 

along the west coast of Africa as far south 

as Angola. The Mediterranean M. cornicula 

can be easily distinguished by its 

smaller shell size (to 25 mm), brown shell 

and brown periostracum, larger protoconch 

(320 µm) with fewer whorls ( 1¼), 

an animal that is entirely white, and a distinctive 

radula with lateral teeth that 

retain prominent dentition along their 

entire width (Rolan et al., 1997:fig. 16D). 

Mitra nigra and M. cornea are more similar 

in protoconch size, radular morphology, 

and periostracal color. Mitra nigra reaches 

a larger shell size (to 70 mm, compared 

to 40 mm for M. cornea) and has a black 

shell and brown to reddish-violet animal, 

while M. cornea has a light brown to pale 

grey shell and a distinctive white animal 

with yellow stripes along the tentacles 

and lateral margins of the foot (see 

Martins, 2004:65, fig. M). It should be 

noted, however, that this distinctive coloration 

is evident in living animals, but 

alcohol preserved specimens are brownish, 

becoming stained as the hypobranchial 

gland exudate oxidizes to a 

purplish brown. 

Thus, the small brown mitrids of the 

Azores are Mitra cornea Lamarck, 1811. 

Earlier literature (e.g. Dautzenberg, 1889) 

may refer to this species as Mitra fusca 

Reeve, 1844, which, however, is a junior 

synonym of M. cornea and is preoccupied 

by Mitra fusca Swainson, 1831, an Indo- 

Pacific species. Cernohorsky (1969:972, 

fig. 28) designated as lectotype for Mitra 

cornea Lamarck, 1811, a specimen from 

the west coast of Africa, thus fixing its 

type locality. He regarded this taxon to be 

a synonym of M. cornicula, a species that 

is endemic to the Mediterranean and 

readily distinguished from M. cornea on 

the basis of shell and radular morphology. 

Mitra aquitanica Locard, 1892, is an 

unnecessary replacement name for Mitra 

fusca Reeve, 1844 non Swainson, 1831. 

Azorean records of Mitra cornea were, 

until recently, reported as Mitra nigra 

(e.g., Cernohorsky, 1976:371; Knudsen, 

1995:152; Morton et al., 1998:57, 65, 76). 

The present study documents the 

anatomy, shell morphology and biology 

of Mitra cornea from São Miguel, Azores, 

in order to provide a basis for more comprehensive 

studies of the systematics and 

relationships of the three small, smooth 

mitrids of the eastern Atlantic and 

Mediterranean Sea. 


Numerous specimens were collected 

from algae covered intertidal rocks along 

the SW wall of the fishing pier in Vila 

Franca do Campo, São Miguel, Azores 

[N37° 42’ 49.75”, W25 25’ 52.10”] (USNM 

1114338). Additional specimens were collected 

on shallow (< 1 m) subtidal rocky 

ledges along the southern rim of the

HARASEWYCH: ANATOMY AND BIOLOGY OF MITRA CORNEA 123 

crater of Ilhéu de Vila Franca, São Miguel, 

Azores [N37° 42’ 18.75”, W25° 26’ 35.33”] 

(USNM 1114340). 

Specimens were maintained and 

observed in seawater. The shells of a subset 

of specimens were cracked in a vice, 

and the animals irritated with forceps until 

they everted their proboscis. These animals 

were then preserved in 70% ethanol 

and transferred to 95% ethanol for storage. 

Animals were dissected, portions critical 

point dried and examined with a scanning 

electron microscope. 

SYSTEMATICS 

Class GASTROPODA 

Order Neogastropoda Wenz, 1943 

Family Mitridae Swainson, 1831 

Genus Mitra Lamarck, 1798 


Synonymy 


Mitra fusca Reeve, 1844 non Mitra 

fusca Swainson, 1831 

Mitra aquitanica Locard, 1892 

Description 

Shell morphology: Shell (Figures 1, 2) small 

for genus (to 34 mm), thick, biconic, 

fusiform, with elongate aperture and 4 

heavy columellar folds. Protoconch 

increasing from ≈ 110 µm to ≈ 1.2 mm in 4 

¼ evenly rounded, conical whorls, badly 

eroded or missing on most specimens. 

Transition to teleoconch indistinct, marked 

by change in surface from glossy to matte, 

and onset of spiral sculpture consisting of 

narrow, pitted furrows separating adjacent 

broad, low, spiral cords. Teleoconch of up 

to 7+ smooth, evenly convex whorls. 

Suture adpressed, irregular, showing evidence 

of axial lamellae. Spiral sculpture of 

low, abutting spiral cords separated by a 

series of closely spaced pits in early 

whorls, that become sharply incised furrows 

in later whorls, and largely obscure 

except along the anterior third of the final 

whorl. Axial sculpture consists of low, 

nearly obscure lamellae that are most evident 

at the suture of smaller specimens 

(Figure 2) from which the periostracum 

has been removed. Pitting, periostracum, 

and repaired breaks (Figure 2, rb) obscure 

the sculpture of larger specimens. 

Aperture elongated, tapering posteriorly 

beneath suture to form anal sulcus. Outer 

lip smooth, thickened, nearly straight 

along middle portion, rounded and dorsally 

reflected anteriorly to form siphonal 

notch. Inner lip with narrow, glazed 

inductura, four broad columellar folds 

(Figure 3, cf), decreasing in prominence 

from posterior to anterior, and a siphonal 

fold (Figure 3, sf). Siphonal notch broad, 

siphonal fasciole usually absent, but may 

be weak and short in larger specimens. 

External shell color ranges from dark 

chestnut brown to purplish gray and may 

be solid or banded. Aperture is white, 

especially the columellar folds, but brown 

color is visible through the white glaze 

near the anterior and posterior margins of 

the aperture. Periostracum thin, chestnut 

brown. Operculum absent. 

Shell ultrastructure: (Figure 4) Shell composed 

of four distinct crystal layers. The 

innermost layer (Figure 4, in) (≈ 90 µm) 

comprising the glaze is whitish, while the 

remaining layers are golden brown. The 

crystal faces of the next layer (Figure 4, 

per) (≈ 275 µm) are crossed-lamellar, and 

oriented perpendicular to the growing 

edge of the shell. They are also perpendicular 

to the crystal faces of the adjacent 

crossed lamellar layer (Figure 4, par) (≈ 435 

µm), which are parallel to the growing 

edge. A prismatic layer (Figure 4, ou) 

(≈ 120 µm) is outermost. 

External anatomy: (Figure 5) The soft tissues 

comprise 2½ whorls, of which the

124 AÇOREANA 

2009, Sup. 6:121-135 

FIGURES 1-4. Mitra cornea Lamarck, 1811. 1. USNM 1114338, intertidal rocks along the SW wall of the 

fishing pier in Vila Franca do Campo , São Miguel, Azores. 2. USNM 1114340, subtidal (< 1 m) rocky 

ledges along the southern rim of the crater of Ilhéu de Vila Franca, São Miguel, Azores. 3. Shell fractured 

to reveal columella. 4. shell ultrastructure. Surface parallel to growing edge of shell, ¼ whorl behind 

the lip. cf, columellar fold; in, innermost shell layer; ou, outer, prismatic shell layer; par, crossed-lamellar 

aragonite, crystal faces parallel to growing edge; per, crossed-lamellar aragonite, crystal faces perpendicular 

to crowing edge; rb, repaired break; sf, siphonal fold.


FIGURES 5-10. Anatomical features of Mitra cornea Lamarck, 1811. 5. Male specimen, lateral view. 6. 

Roof of mantle cavity. 7. Alimentary system, semi-diagramatic. 8. Lateral view of anterior proboscis, 

opened from right side. 9. Female reproductive system. 10. Male reproductive system. a, anus; ag, albumen 

gland; ae, anterior esophagus; bc, bursa copulatrix; cg, capsule gland; cm, columellar muscle; ct, 

ctenidium; dg, digestive gland; dsg, duct of salivary gland; ep, epiproboscis; fo, female opening; hg, 

hypobranchial gland; ig, ingesting gland; k, kidney; m, mouth; me, mantle edge; ng, nephridial gland; 

nr, nerve ring; os, osphradium; p, penis; pb, proboscis; pc, pericardium; pp, peristomal papillae; pro, 

prostate gland; r, rectum; rg, rectal gland; rmc, rear of mantle cavity; rr, radular ribbon; s, siphon; sg, salivary 

gland; sto, stomach; sv, seminal vesicle; t, testis; td, testicular duct; vd, vas deferens; vor, ventral 

odontophoral retractor muscle.

126 AÇOREANA 

2009, Sup. 6:121-135 

mantle cavity spans ¾ whorl, the kidney 

¼ whorl, and the digestive gland and 

gonad 1½ whorls. The columellar muscle 

is long, narrow, attaching to the shell 1 ¼ 

whorl behind the mantle edge. The foot is 

long, narrow, rectangular (L/W ≈ 2.0), 

with a deep propodial groove along the 

anterior edge. In living specimens, the 

foot is bright white with a narrow, bright 

yellow band along the lateral edges of the 

foot and outer edges of the tentacles as far 

as the eyes. The color is rapidly lost in 

alcohol preserved specimens, which 

become progressively darker brown over 

time. The siphon is long and muscular. 

Mantle cavity: (Figure 6) The arrangement 

of mantle cavity organs is similar to that 

of most neogastropods. The mantle edge 

is thin and smooth, somewhat thickened 

at the right margin. The osphradium 

(Figure 6, os) is bipectinate, large, brownish, 

with 64-73 filaments above the ganglion 

and 52-60 below. The ctenidium 

(Figure 6, ct) is twice a broad and twice as 

long as the osphradium. A large renal 

organ (Figure 5, k), in which the primary 

and secondary lamellae do not interdgitate 

[termed Meronephridiens by Perrier 

(1889)] forms the right, rear wall of mantle 

cavity. The short, broad pericardium 

(Figure 5, pc), with a narrow nephridial 

gland (Figure 5, ng) lies to the left of the 

kidney. The hypobranchial gland (Figure 

6, hg) is broad and thick, occupying much 

of the dorsal roof of the mantle cavity 

between the osphradium and gonoduct. 

It produces a copious viscous secretion 

that is clear at first, but becomes yellowish 

then purple, and finally dark brown. 

The color is alcohol soluble and stains the 

tissues of preserved specimens. 

Alimentary system: (Figure 7) The pleurombolic 

proboscis (Figures 5, 7 pb) is 

moderately long (extends to ≈ 1.5 x Shell 

length), broad, especially distally, enclosing 

a large, compact buccal mass and 

epiproboscis (Figures 5, 7, 8, 13-22, ep), an 

extensible muscular introvert unique to 

mitrids. When retracted, the proboscis is 

slightly folded within proboscis sheath 

and fills nearly the entire cephalic hemocoel. 

The buccal mass is broad, with a 

short, muscular, peristomal rim lined 

with evertable papillae surrounding the 

mouth. The odontophore and radular 

ribbon are large, with strong protractor 

and retractor muscles. 

The radular ribbon (Figures 11, 12) is 

short and broad (L/W ≈ 3.8; L = 1.9 mm, W 

= 500 µm), consisting of 45-57 rows of 

teeth, and may be slightly asymmetrical, 

with left lateral teeth 5-14% wider than 

right in some specimens. The rachidian 

teeth (Figure 11, rt) have five strong, 

curved cusps emerging from a weakly 

trapezoidal basal plate that is broader at 

the posterior end. Each of the five cusps 

has a broad, thick dorsal surface continuous 

with the anterior end of the basal 

plate, and a sharply tapering, knife-like 

posterior-ventral edge that descends to 

the posterior end of the basal plate. The 

central cusp is longest, lateral cusps are 

shortest. The lateral teeth (Figure 11, lat) 

have a complex, semi-recurved basal 

plate that it broad, thin and flat along its 

outer edge, becoming narrower, thicker, 

with its anterior edge raised by up to 35º 

along the inner 1/3 of its length. Each 

tooth has 14-18 cusps. The 2 nd → 4 th from 

the inner edge are strongest, with the 

same knife-like structure as the cusps on 

the rachidian teeth. The remaining teeth 

become more conical and diminish in 

length and size toward the lateral edge, 

with the outermost 0.18 of the basal plate 

lacking discernible cusps. 

The epiproboscis is a long, anteriorly 

tapering, muscular rod, consisting of a 

central core of longitudinal muscles, surrounded 

by circular muscles and encased 

in an inner and outer sheath. From its


FIGURES 11-12. Radula of Mitra cornea Lamarck, 1811. 11. Dorsal view of half row of radular 

teeth. 12. Oblique view of radular teeth. lat, lateral tooth; rt, rachidian tooth. 

opening just below the mouth (Figures 8, 

22, m), the epiproboscis runs mid-ventrally 

beneath the buccal mass, recurving 

dorsally to form a U-shaped bend behind 

the buccal mass to become attached to the 

odontophore by a short, broad retractor 

muscle. The large, ascinous salivary 

glands (Figure 7, sg) are situated above 

the nerve ring (Figure 7, nr) in the anterior 

portion of the cephalic hemocoel, 

generally to the right of the retracted proboscis. 

The ducts from these glands run 

anteriorly alongside the esophagus, joining 

the epiproboscis at the bend, running 

at first ventrally, then medially within the 

epiproboscis, merging into a single opening 

at its tip (Figures 13, 22, sd). 

The anterior esophagus is broad, and 

flat above the buccal mass (Figure 19, ae), 

with low longitudinal ridges, but 

becomes narrower and more circular posterior 

(Figure 21, ae) to the bend in the 

epiproboscis. The esophagus passes 

through the nerve ring without forming a 

distinctive valve of Leiblein, broadens to 

form a crop-like structure, and runs posteriorly 

to join a broad, muscular stomach. 

Neither a gland of Leiblein nor 

accessory salivary glands are present. 

The stomach (Figure 7, sto) has a muscular 

gizzard between the esophagus and 

the closely spaced ducts of the digestive 

glands. The intestinal region has low longitudinal 

ridges that lead to the long 

intestine, which runs along the kidney 

and pericardium before entering the rear 

of the mantle cavity. The rectum (Figure 

7, r) runs alongside the pallial gonoduct, 

forming the anus (Figures 7, 9, a) anterior 

to the gonoduct, but some distance from 

the mantle edge. A long, narrow rectal 

gland (Figure 7, rg) runs along the roof of 

the mantle cavity nearly its entire length, 

before joining the rectum near its anterior 

margin. 

Of the 20 specimens of Mitra cornea 

dissected, 7 were found to have essentially 

intact sipunculans (Golfingia sp. = G. 

margaritaceum fide Morton et al. 1998:76) 

within their gut, usually in the crop or 

stomach. No other recognizable prey was 

found within any of the specimens. 

Male reproductive system: The testis 

(Figure 10, t) is yellowish, lining the 

right ventral side of the digestive gland. 

A duct leading anteriorly expands to 

form the broad, highly convoluted seminal 

vesicle (Figure 10, sv) near the anterior 

margin of the digestive gland. From

128 AÇOREANA 

2009, Sup. 6:121-135 

FIGURES 13-16. Proboscis tip of Mitra cornea Lamarck, 1811, showing extension of the epiproboscis. 

13. Lateral and 14. frontal views of proboscis tip in early stage of extension of epiproboscis. 

15. lateral and 16. frontal views of proboscis tip with epiproboscis extended. Arrows in figure 15 

indicate planes of section for figures 17-21. ep, epiproboscis; m, mouth; pb, proboscis; pp, peristomal 

papillae; s, siphon; sd, salivary duct opening; t, cephalic tentacle. 

there, the renal vas deferens runs along 

the surface of the nephridium without 

giving rise to a gonopericardial duct, 

entering the pallial cavity where it 

expands to form the prostate gland 

(Figure 10, pro). The prostate gland is 

open to the pallial cavity along its broad 

posterior portion, but forms a closed 

duct anteriorly that descends to the floor 

of the mantle cavity (Figure 10, vd) and 

runs to the base of the penis (Figures 5, 

10, p), situated behind the right cephalic 

tentacle. The penis is broad basally, 

strongly recurved, with a flagellate 

pseudo-papilla, with the duct opening 

at its tip.


FIGURES 17-22. Proboscis tip of Mitra cornea Lamarck, 1811. 17-21. Transverse sections through proboscis 

tip shown in figure 15. 22. Saggital section through proboscis tip in figure 13. ae, anterior esophagus; 

bv, blood vessel; dop, dorsal odontophoral protractor muscle; ep, epiproboscis; eps, epiproboscis 

sheath; es, epiproboscis sheath; lm, longitudinal muscle; m, mouth; odc, odontophoral cartilage; pp, 

peristomal papillae; pr, peristomal rim; rs, radular sac; rt, radular teeth; sd, salivary duct; vor, ventral 

odontophoral retractor muscle.

130 AÇOREANA 

2009, Sup. 6:121-135 

Female reproductive system: The ovary lies 

along the right side of the digestive gland 

and dominates the uppermost visceral 

whorls. The oviduct runs anteriorly from 

the ovary, along the kidney and pericardium 

without giving rise to a gonopericardial 

duct, before entering the rear of the 

mantle cavity. The pallial gonoduct 

(Figure 9) consists of an albumen gland, 

ingesting gland, capsule gland and bursa 

copulatrix. The albumen gland (Figure 

9, ag) is tall, narrow, and glandular, with 

ventral channel. The ingesting gland 

(Figure 9, ig) is situated between the 

albumen gland and long capsule gland 

(Figure 9, cg). The large, muscular bursa 

copulatrix (Figure 9, bc) is situated above 

the female opening (Figure 9, fo) and 

above and anterior to the capsule gland. 

A prominent ventral pedal gland is situated 

along the ventral mid-line of the 

foot, just anterior to the bursa copulatrix. 

Reproductive Biology: Knudsen (1995:153, 

fig. 13) illustrated the egg capsule and 

pre-hatching larva of this species [as 

Mitra nigra], noting that development 

was pelagic in this species. It was not 

known whether the larva develops into a 

sinusigera larva during the pelagic 

phase. 

DISCUSSION 

As the number of mitrids species that 

have been studied anatomically increases, 

all have been found to have a highly 

specialized anterior alimentary system 

with a broad, extensible, muscular proboscis 

and a uniquely evolved muscular 

epiproboscis that is extended through 

the mouth (Figures 13, 14, pp), and 

through which pass the ducts of the salivary 

glands, joining into a single duct 

before emptying at its tip (Figures 13, 14, 

sd). In mitrids, the accessory salivary 

glands and gland of Leiblein are absent 

and the valve of Leiblein is greatly 

reduced or absent. These anatomical 

structures are adaptations to a specialized 

diet consisting nearly exclusively of 

sipunculans (see Taylor, 1993 and references 

therein). The diet of more than 30 

species of mitrids has been studied, yet 

there is but a single report of one individual 

of one species, feeding on a 

nemertean (Fukuyuma and Nybakken, 

1983). 

The earliest fossil record for family 

Mitridae dates to the basal Late 

Cretaceous [Cenomanian / Turonian] 

(Tracey et al. 1993:152), with most modern 

genera diverging during the Miocene 

(Cernohorsky, 1970:fig. 180). Yet all surviving 

lineages have a well developed 

epiproboscis, and lack accessory salivary 

glands and a gland of Leiblein, suggesting 

an early common origin of this specialized 

body plan, adapted for preying 

on sipunculans. Relatively few neogastropods 

prey occasionally on sipunculans 

[eg., Vasum, Drupa, Bursa] and, other 

than Mitridae, only a few species of 

Drupina feed exclusively on them (Taylor 

1989:271). 

Sipuncula is a small phylum with 

only about 150 species worldwide 

(Cutler, 1994:3). Sipunculans live in a 

variety of habitats, burrowing in rock, 

sand or rubble bottoms at all depths. 

The diversity of Mitridae is somewhat 

greater, at 377 living species 

(Cernohorsky, 1970:Table 1). 

In the tropics, mitrid diversity is high 

and species partition their habitat by 

substrate type (e.g., Taylor, 1989:fig. 7), 

although multiple species can co-occur 

in the same general habitat (e.g. thick 

sand). In the Azores, the two mitrid 

species appear to partition their habitat, 

with M. cornea inhabiting intertidal to 

subtidal reef platforms and boulder rubble, 

while M. zonata occurs in offshore 

sandy substrates. Sipunculan diversity


in the Azores somewhat exceeds mitrid 

diversity. 

Interpretation of the function of the 

epiproboscis has varied over the years. 

Several authors have proposed that the 

epiproboscis serves as a venomous organ 

(Vayssiere, 1901; Cernohorsky, 1970), is 

used for both offense and defense 

(Vayssiere, 1901), or applies the products 

of the salivary glands to the sipunculan 

prey (Risbec, 1928; Ponder, 1972, 1998). 

There has also been some question as to 

whether the salivary gland secretion 

weakens the prey integument, serves as 

a relaxant to prevent prey contraction, or 

acts as an adhesive substance to facilitate 

the removal of sipunculans from their 

burrows (West, 1990:774). 

Several authors have reported on the 

feeding behavior of mitrids with varying 

levels of detail. Loch (1987) reported that 

the epiproboscis is extended during 

feeding and inserted into its prey, and is 

thought to deliver salivary secretions. 

Taylor (1989:262) suggested that the peristomal 

papillae [“circum-oral, brush like 

structure”] can be everted during feeding 

and may function to grip prey, noting 

that, while mitrids have a large radula 

with long, multicuspate lateral teeth, 

these teeth may be used to assist with 

swallowing the prey, as the sipunculans 

and not shredded. 

The most detailed observations of 

feeding behavior were those of West 

(1990), who reported that Mitra idae 

located its prey with its siphon, first 

touching it with the siphon edge and 

cephalic tentacles before extending its 

proboscis. The peristomal rim then 

grasps the prey in a series of rapid eversions 

and contractions and the snail 

retracts its proboscis in attempts to pull 

the sipunculan from the substratum. If 

the prey cannot be extracted and swallowed 

whole, secretions from the salivary 

gland are applied and the radula 

used to rasp a hole in the integument. 

The epiproboscis is then inserted into the 

hole, entangling the sipunculan viscera 

and pulling them into the mitrid buccal 

mass. This alternating extension and 

retraction of the epiproboscis is repeated 

multiple times. The proboscis periodically 

further envelopes the sipunculan 

and eventually frees the sipunculan from 

the substratum, by grasping the integument 

with the radula and retracting the 

odontophore. This behavior is characteristic 

of members of the subfamily 

Mitrinae, which includes Mitra cornea. 

Later, West (1991:710) reported on 

the feeding in Mitra catalinae, a member 

of the subfamily Cylindromitrinae. As in 

Mitrinae, the proboscis was extended, 

the peristome flared to grasp the prey 

and the radula was used to rasp a small 

hole in the integument through which 

the epiproboscis was inserted. However, 

in this species, the epiproboscis was 

“rhythmically passed in and out of the 

hole” pumping sipunculan coelomic fluids 

and eggs into the buccal cavity and 

down the esophagus of the mitrid, without 

ingesting either viscera or the entire 

sipunculan. West (1991:710) noted that 

the sipunculans “survived the feeding 

session,” suggesting parasitism rather 

than predation in this mitrid species. 

Ponder (1972:335) raised the question 

of how the epiproboscis might have 

evolved. He suggested that the ducts of 

the salivary glands migrated ventrally, 

and that their openings moved from lateral 

positions on the buccal mass to a 

ventral anterior position at the edge of 

the mouth. The next hypothesized stage 

was their placement on a papilla that 

eventually became invaginated and elongated. 

Ponder noted that the salivary 

glands of mitrids have a second type of 

secretory cell not present in related families, 

and that these cells may produce a 

toxin. It is interesting to note, that

132 AÇOREANA 

2009, Sup. 6:121-135 

although the salivary glands are ascinous 

and typical of salivary glands of other 

neogastropods, the position of the ducts, 

including their becoming fused into a single 

duct before emerging medially at the 

ventral anterior edge of the buccal mass is 

typical of the ducts of accessory salivary 

glands, which are absent in Mitridae. 

West (1991) noted that the epiproboscis 

shows structural and functional affinities 

with other molluscan subradular organs, 

and hypothesized that it developed from 

the musculature of the buccal mass, while 

the sheaths were derived from the walls 

of the buccal cavity. According to the 

Ponder model, the primary function of 

the epiproboscis is the targeted application 

of salivary gland secretions. The 

West model is based on a primarily 

mechanical function of the epiproboscis. 

West (1991:716) commented that the ventral 

migration of the salivary gland ducts 

“to connect with the formative epiproboscis” 

might have changed, or enhanced 

its function, but that it is difficult to assess 

the evolutionary importance of this event 

without knowledge of the function of salivary 

gland secretions”. 

While the morphological adaptations 

of the anterior alimentary system that 

have evolved in conjunction with a specialized 

sipunculan diet are now well 

documented within the Mitridae, nothing 

is known of the chemical adaptations, 

including the composition and physiological 

effects of the secretions of the salivary 

glands, which are delivered to the prey 

through the epiproboscis. The salivary 

glands of related gastropods have been 

shown to produce neurotoxins with 

acetylcholine-like effects that are used to 

overcome prey with a paralytic secretion 

(West et al., 1998). Still others contain proteinaceous 

toxins that are hemolytic and 

lethal (Shiomi et al., 2002). Given that the 

Mitridae represent an adaptive radiation 

following an early [Upper Cretaceous] 

adaptation to a highly specialized diet, 

the structure, specificity and toxicology of 

salivary gland secretions represent a fertile 

area for the study of chemical evolution 

within Mollusca, and interactions of 

predator and prey. 

Similarly, the hypobranchial gland of 

mitrids is voluminous, and noted for producing 

substantial quantities of a 

secretion that oxidizes to form a purple 

pigment. Numerous authors since 

Dubois (1909) have documented that, in 

addition to chromogens, hypobranchial 

gland secretions from various 

neogastropods contain numerous toxic 

and paralytic compounds, including 

choline esters, serotonin and various 

biogenic amines (eg., Shiomi et al., 1998). 

The hypobranchial gland of the neogastropod 

Thais haemastoma (which co-occurs 

with Mitra cornea) contains multiple 

active components, including one that 

produces a stimulatory effect on blood 

pressure, and another that acts as a neuromuscular 

blocking agent of the depolarizing 

type (Hyang & Mir, 1971). Some 

muricids have been observed to use 

hypobranchial gland secretions to 

immobilize prey (Naegel & Alvarez, 

2005:426). Other taxa, including trochids 

(Kelley et al., 2003) and pleurotomariids 

(Harasewych, 2002) have been shown to 

use hypobranchial gland secretions to 

repel predators. 

Despite the exceptionally thick shells, 

periostracum and overgrowth of calcified 

encrusting organisms, cleaned shells of 

many Mitra cornea show evidence of multiple 

repaired breaks (Figure 2, rb). This 

high incidence of unsuccessful predation 

(measured as frequency of shell repair) is 

an indication of high exposure to crushing 

predation as well as of the ability to 

survive such attacks by predators. 

Among shallow water gastropods, 

Vermeij (1989) reported that the incidence 

of unsuccessful predation was greatest in


the Indo-West Pacific and lowest in the 

Atlantic. Frequencies of unsuccessful 

predation of 0.5 breaks per individual 

were uncommon, and never averaged 

more than 1 break per individual for the 

most prey-resistent taxa. Repaired breaks 

on just the last whorl of the shell of Mitra 

cornea range from 0-5 [mean = 1.95, n = 

20], suggesting both an exceptionally 

high rate of attack, and ability to survive 

attack. While this points to the possible 

use of hypobranchial gland secretions as 

a chemical defense against predators, it 

does not preclude the presence of other 

compounds or other uses for the secretory 

products of the hypobranchial gland. 


I am grateful to Prof. António Frias 

Martins and to his staff and students for 

organizing the Third International 

Workshop of Malacology in Vila Franca 

do Campo, São Miguel, Azores. This 

Workshop was a joint organization of 

Sociedade Afonso Chaves and the 

Department of Biology of the University 

of the Azores. Support from FLAD 

(Portuguese-American Foundation for 

Development) is gratefully acknowledged. 

Thanks to Marilyn Schotte for 

assistance with histology and to Yolanda 

Villacampa for Scanning Electron 

Micrographs. 

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COMPARATIVE STUDY OF CHEMICAL DEFENCES FROM TWO ALLOPATRIC 

NORTH ATLANTIC SUBSPECIES OF HYPSELODORIS PICTA (MOLLUSCA: 

OPISTHOBRANCHIA) 

Helena Gaspar 1 , Ana Isabel Rodrigues 1 & Gonçalo Calado 2 

1 Instituto Nacional de Engenharia, Tecnologia e Inovação, I.P. (INETI), Est. Paço do Lumiar, Ef. F, 1649-038 

Lisboa, Portugal. e-mail helena.gaspar@ineti.pt 

2 Faculdade de Engenharias e Ciências Naturais, Universidade Lusófona de Humanidades e Tecnologias. 

Av. do Campo Grande, 376 1749 - 024 Lisboa PORTUGAL 

ABSTRACT 

Different mixtures of furanosesquiterpenes characterized the defensive metabolites of 

the two populations of the nudibranchs Hypselodoris picta azorica and Hypselodoris picta 

webbi from the NW Atlantic, studied here for the first time. The known furonosesquiterpenes 

tavacfuran and longifolin were found in both subspecies, whereas micorcionin-1 

was only present in Hypselodoris picta azorica. 

RESUMO 

O estudo de duas populações de nudibrânquios Hypselodoris picta azorica e Hypselodoris 

picta webbi do Atlântico NW, descrito pela primeira vez neste artigo, mostrou que os seus 

metabolitos de defesa são misturas diferentes de furanosesquiterpenos. Em ambas as 

populações foram encontrados os metabolitos tavacfurano e longifolina, tendo a 

microcionina-1 sido identificada apenas na subespécie Hypselodoris picta azorica. 

INTRODUCTION 

Nudibranchs are shell-less opisthobranch 

molluscs that, in the course 

of evolution have developed several 

defensive strategies which include the 

use of secondary metabolites to avoid 

predation (Wägele, 2006). These compounds 

can be accumulated from 

dietary sources, biosynthesised de novo 

or bio-transformed from dietary 

metabolites in order to provide the 

nudibranchs with more effective defensive 

allomones or non toxic compounds 

(Cimino, 1993, 1999). Most species of 

the genus Hypselodoris (Family 

Chromodoridae) store their allomones 

in high levels of concentrations, on special 

mantle glands denominated MDFs 

(Mantle Dermal Formations). These 

glands are strategically located near the 

vital organs, gills and rhinophores, or 

even ventrally along the mantle margin. 

They are regarded as the development 

of a chemical defensive mechanism in 

the course of opisthobranchs’ evolution 

(Wägele, 2006; García-Gómez, 1990; 

Fontana, 1993). 

Previous chemo-ecological studies of 

Hypselodoris picta (previously known as 

H. webbi or Glossodoris valenciennesi) 

showed that this species, like other 

Hypselodoris, accumulates in the mantle 

(especially in MDFs) furanosesquiterpenes, 

longifolin (1) being usually the 

main metabolite (Figure 1; Table 1). This 

compound, initially isolated from the 

terrestrial Japanese plant Actinodaphne 

longifolia (Hayashi, 1972 fide Guella, 

1985), was the major metabolite found in 

a Mediterranean population of the 

marine sponge Dysidea fragilis (Avila, 

1991) and has antifeedant and ichthyodeterrent 

activities (Fontana, 1993). 

Ecological experiments performed with 

Mediterranean populations of H. picta 

showed the transference of sponge 

metabolites into the MDFs and supported 

the dietary origin of Hypselodoris’ 

allomones (Fontana, 1994b).

138 AÇOREANA 

2009, Sup. 6: 137-143 

FIGURE 1. Chemical structures of furanosesquiterpenes present in different populations of 

Hypselodoris picta. 

TABLE 1 – Furanosesquiterpenes present in different Mediterranean populations of Hypselodoris picta.

GASPAR ET AL: CHEMICAL DEFENCES OF HYPSELODORIS PICTA 139 

Hypselodoris picta (Schultz, 1836) 

seems to be widespread in the Atlantic 

Ocean. So far, five geographic subspecies 

are assigned, H. picta azorica being 

restricted to the Azorean Archipelago, 

whereas H. picta webbi spreads from the 

Caribbean to the Canary Islands and the 

Iberian Peninsula (Ortea et al., 1996). The 

colour pattern varies consistently (Figure 

2), but their internal anatomy is still very 

similar (Alejandrino & Valdés, 2006). No 

genetic data are available to date. In this 

paper, we report a comparative study of 

chemical defences of these two allopatric 

subspecies of Hypselodoris picta from the 

North Atlantic, H. picta webbi from the 

Algarve (SW coast of Europe) and H. picta 

azorica from Azores, with the aim to clarify: 

(1) what are the defensive metabolites 

of H. picta azorica; (2) and if the two subspecies 

have the same chemical profile. 


Biological material. The molluscs were collected 

by SCUBA in two stations: 

Hypselodoris picta azorica (6 specimens July 

2006, 2 specimens September 2008) at 

Ilhéu de Vila Franca do Campo, S. Miguel 

Island, Azores, Portugal, and Hypselodoris 

picta webbi (2 specimens July 2006, 2 specimens 

August 2008) at Portimão, Algarve, 

Portugal. The taxonomic identification of 

H. picta was made by one of us (G.C.). 

Voucher specimens (preserved in 

absolute ethanol) are deposited at 

“Instituto Português de Malacologia”, 

Portugal, reference numbers IPM.MO.005 

and IPM.MO.006, respectively. 

Chemical analysis. Silica-gel chromatography 

was performed using precoated 

Merck F 254 

plates and Silica gel Merck 

Kieselgel 60. 1 H NMR spectra were 

acquired in CDCl 3 

or C 6 

D 6 

on a Bruker 

AMX-300 operating at 300 MHz. 

Two frozen nudibranchs of each subspecies, 

collected in 2006, were dissected 

and differentiated in mantle dermal formations 

(MDFs), the remaining of mantle 

and digestive glands. All the dissected 

sections were separately extracted three 

times with acetone by sonication. The 

concentrated extracts were partitioned 

between diethyl ether and water. The 

resulting diethyl ether extracts were compared 

by TLC (Thin Layer Chromatography). 

The Erlich positive metabolite, 

with R f 

= 0.2 in n-hexane, present in 

both subspecies, was isolated from the 

mantle extract (4 mg) of one specimen of 

H. picta webbi (fresh weight 8.3 g). The 

mantle extract was chromatographed 

on a silica-gel column packed with 

n-hexane and eluted with a gradient of 

n-hexane/diethyl ether. Fractions eluted 

with n-hexane were concentrated to give 

2 mg of compound 1 (Figure 1). 

FIGURE 2. Live specimens of the Hypselodoris picta studied herein: A - H. p. azorica (photo Peter 

Wirtz); B - H p. webbi (photo Rita Coelho).

140 AÇOREANA 

2009, Sup. 6: 137-143 

We were unable to isolate the main 

furanosesquiterpene presented in H. picta 

azorica collected in 2006 because, due to 

its high instability, it had suffered degradation 

during the isolation procedure. 

Each frozen mollusc (4 specimens) 

collected in 2008 was separately 

immersed in acetone and extracted by 

sonication (3 times, 2 min). The concentrated 

extracts were partitioned between 

diethyl ether and water. The resulting 

mantle extracts were compared by TLC 

and 1 H NMR analysis. 

RESULTS 

We have chemically investigated for 

the first time the nudibranch Hypselodoris 

picta azorica and a population of the nudibranch 

Hypselodoris picta webbi from the 

NW Atlantic. The TLC analysis of diethyl 

ether extracts revealed the presence of at 

least two Erlich positive compounds in 

both subspecies. H. picta webbi showed 

two main Erlich positive metabolites (R f 

= 

0.2 and 0.3 in n-hexane) and H. picta azorica 

additionally showed the presence of 

one or two less polar compounds (R f 

= 0.7 

and 0.8 in n-hexane). Each analysed specimen 

showed the same metabolite pattern 

in the extracts obtained from the different 

anatomic parts. The less polar metabolite 

was identified as microcionin-1 (7) by 

TLC comparison with an authentic sample 

(Figure 1). We were unable to isolate 

the furanosesquiterpene with R f 

= 0.7, the 

main metabolite of H. picta azorica collected 

in 2006, due to its instability. The component 

with R f 

= 0.2, isolated from one 

specimen of H. picta webbi collected in 

2008, was identified as longifolin (1). The 

1 

H NMR signals (either in CDCl 3 

or C 6 

D 6 

) 

of this metabolite were identical to those 

reported in the literature (Guella, 1985). 

Longifolin was also identified by NMR as 

the main furanosesquiterpene in the mantle 

extract obtained from H. picta azorica 

collected in 2008 (Figure 3a). The analysis 

of 1 H NMR spectrum (Figure 3b) of the 

mantle extract from H. picta webbi, collected 

in 2008, allowed the identification of 

tavacfuran (11), the compound with R f 

= 

0.3, as the main furanosesquiterpene 

(Figure 1). The 1 H NMR signals attributed 

to both compounds in the NMR 

spectra of mantle extracts (Figure 3) are 

similar to those previous reported data 

(Guella, 1985). The results concerning the 

furanosesquiterpenes are summarized in 

Table 2. 

TABLE 2 – Furanosesquiterpenes found in Hypselodoris picta in this study.


FIGURE 3. 1 H NMR (300MHz, CDCl 3 

) from diethyl ether extracts of Hypselodoris mantle: a) 

Hypselodoris picta azorica; b) Hypselodoris picta webbi. 

DISCUSSION 

In the mantle of Hypselodoris picta azorica 

and Hypselodoris picta webbi the 

furosesquiterpenes longifolin (1) and 

tavacfuran (11) were identified by 1 H 

NMR (Figure 3). TLC comparison of the 

mantle extracts showed that these typical 

sponge metabolites were found in all 

analysed specimens but in different relative 

proportions (Table 2). Longifolin (1) 

has usually been found as the main 

metabolite of H. picta from Mediterranean 

(Table 1). Tavacfuran (11), a compound 

first reported on a mixture of 

Mediterranean sponges (Guella, 1985)

142 AÇOREANA 

2009, Sup. 6: 137-143 

and already isolated from other 

Hypselodoris species (Fontana, 1993), was 

now found for the first time in H. picta. 

The TLC analysis allowed the identification 

of microcionin-1 (7) in all the specimens 

of H. picta azorica. It is worth noting 

that a Mediterranean population of 

H. picta is able to transfer microcionin-1 

(7) and other sponge metabolites into 

MDFs (Fontana, 1994b). 

The high concentration of furanosesquiterpenes, 

such as longifolin (1) 

and tavacpallescencin (11) in the mantle 

(including MDFs), suggests that, as in 

other Hypselodoris, they play an important 

role as defensive metabolites. This 

study showed that the chemical profiles 

of H. picta azorica and H. picta webbi are 

different, as expected, since previous 

studies support the idea that most of 

these furanosesquiterpene compounds 

are accumulated from dietary sources 

(Avila, 1991; Fontana, 1994b). However, 

the presence of longifolin (1) in H. picta 

subspecies living at distant geographical 

areas could suggest two different scenaria: 

(a) the ability to biosynthesized de 

novo this allomone or (b) the ability to 

select sponges rich in longifolin (1) 

(Avila, 1991; Cimino 1993). Therefore, 

the origin of longifolin in H. picta still 

remains to be clarified. 


We thank Professor António Frias 

Martins for the invitation to participate 

in the 3 rd Workshop of Marine Biology, 

held in Vila Franca do Campo, where 

some of the specimens were collected. 

We thank Rita Coelho (IPM) and Lucas 

Cervera (University of Cadiz) for collecting 

the other specimens. We are 

deeply grateful to FCT (VERMEJJ – 

PTDC/MAR/65854/2006) for financial 

support. 


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from Italian and Spanish Coasts. 

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Chemical Defense of Four Mediterranean 

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474. 

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in Marine Molluscs. In: 

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products – diversity and biosynthesis. 

Topics in current chemistry vol. 167: 78- 

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Chemical defense and evolutionary 

trends in biosynthetic capacity among 

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9: 187-307. 

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G. CIMINO, 1993. Defensive 

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Hypselodoris (Gastropoda: Nudibranchia) 

from the Cantabrian Sea. 

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MOLLO, C. CIMINO, C. AVILA, E. 

MARTINEZ & J. ORTEGA, 1994a.


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nudibranchs: A new furnosesquiterpenoid 

from Hypselodoris webbi. 

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E. MOLLO & G. CIMINO, 1994b. 

Transfer of secondary metabolites 

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THE BIOLOGY OF THE ZONING SUBTIDAL POLYCHAETE DITRUPA ARIETINA 

(SERPULIDAE) IN THE AÇORES, PORTUGAL, WITH A DESCRIPTION OF THE 

LIFE HISTORY OF ITS TUBE 

Brian Morton & Andreia Salvador 


e-mail addresses: prof_bmorton@hotmail.co.uk; a.salvador@nhm.ac.uk 

ABSTRACT 

In the Açores, Portugal, the steeply descending continental shelf is characterized at different 

depths by two endobenthic suspension feeding species: the shallower-living (0-~100 

metres) bivalve Ervilia castanea and the deeper-residing serpulid (~100–250 metres) Ditrupa 

arietina. As a dominant member of the continental shelf fauna, D. arietina provides a habitat 

for a number of epibiont species that attach to its tubes anteriorly. These include the 

cemented, introduced, serpulid Hydroides elegans, three species of foraminiferans and a 

number of species of, mostly unidentifiable, bryozoans. Tubes of D. arietina are also drilled 

and, hence, predated by a prosobranch naticid gastropod, possibly Natica prietoi. Ditrupa 

arietina lives for ~2 years with most growth occurring during the first year and with sexual 

maturity also occurring in the first year of life so that, post reproduction and death, its 

tube becomes available for secondary colonization by Aspidosiphon muelleri (Sipuncula). At 

this time, anterior epibionts die, because the sipunculan orientates the tube anterior end 

down, but further epibionts can now colonize the posterior end of the tube. With time and 

wear, the tube slowly degenerates. Throughout its life history, therefore, the tube of D. arietina 

functions as an inhabitable substratum and is thus a locally important Açorean habitat 

for a suite of other epibenthic species. 

Ditrupa arietina is herein recognized as a significant component of the continental shelf 

endobenthos at depths of ~200 metres, mirroring the significance of the bivalve Ervilia castanea 

at generally shallower depths. The ecological importance of these two species is 

hence in urgent need of further detailed study, especially with regard to the productivity 

of the Açorean seabed. 

RESUMO 

Nos Açores, a plataforma continental que desce abruptamente caracteriza-se, a 

diferentes profundidades, por duas espécies endobênticas filtradoras de matérias em 

suspensão: o bivalve de pouca profundidade (0-~100 metros) Ervilia castanea e o serpulídeo 

de maiores profundidades (~100-250 metros) Ditrupa arietina. Elemento dominante da 

fauna da plataforma continental, D. arietina proporciona habitat para uma quantidade de 

espécies epibiontes que se fixam ao seu tubo anteriormente. Incluem-se nestes o 

serpulídeo fixo Hydroides elegans, três espécies de foraminíferos e uma série de briozoários 

na maioria não identificáveis. Os tubos de D. arietina são também perfurados e, por isso, 

predados por um gastrópode prosobrânquio naticídeo, provavelmente Natica prietoi. 

Ditrupa arietina vive por ~2 anos ocorrendo a maior parte do crescimento durante o 

primeiro ano e a maturidade sexual ocorrendo o primeiro ano de vida pelo eu, após a 

reprodução e a morte, o seu tubo fica acessível para colonização secundária por 

Aspidosiphon muelleri (Sipuncula). Por essa altura, os epibiontes anteriores morrem porque 

o sipúnculo orienta para baixo a extremidade anterior do tubo, mas outros epibiontes 

podem agora colonizar a extremidade posterior do tubo. Com o tempo e uso, o tubo 

degenera lentamente. Durante a sua história de via, assim, o tubo de D. arietina funciona 

como substrato habitável e é, por isso, um habitat Açoriano localmente importante para 

uma série de outras espécies.

146 AÇOREANA 

2009, Sup. 6: 145-156 

Como componente significativo do endobentos da plataforma continental a 

profundidades abaixo de ~200 metros, espelhando o significado do bivalve Ervilia castanea 

em profundidades geralmente menores, a importância ecológica destas duas espécies 

necessita estudo ulterior pormenorizado, especialmente no que respeita a produtividade 

do fundo marinho Açoriano. 

INTRODUCTION 

Morton & Britton (1995, table 1) 

showed that offshore from Vila 

Franca do Campo, São Miguel, Açores, 

the abundances of members of the 

benthic community were consistent, save 

for Ditrupa arietina (O.F. Müller, 1776) 

whose numbers differed significantly 

between stations. These authors also 

noted that empty tubes of D. arietina were 

occupied by the sipunculan Aspidosiphon 

muelleri Diesing, 1851, but not by the 

hermit crab Anapagurus laevis (Bell, 1845). 

It is known that many species of tubebuilding 

polychaetes form dense aggregations 

or “patches” within marine softbottom 

habitats (Bolam & Fernandes, 

2002) although in the Bay of Blanes 

(northwest Mediterranean), Ditrupa arietina 

showed seasonal peaks of abundance 

in May and June of 1993, 1994 and 1995 

(Sardá et al., 1999). Similarly, since the 

late 1980’s, D. arietina has increased in 

abundance all along the northwest coast 

of the Mediterranean, numbers reaching 

up to 3,000 individuals·m² and adversely 

affecting the functioning of the coastal 

benthic ecosystem in the region (Gremare 

et al., 1998a, b). 

The strictly dioecious serpulid Ditrupa 

arietina has a reproductive period lasting 

from November to May in the bay of 

Banyuls-sur-Mer in the northwestern 

Mediterranean, with recruitment beginning 

as early as January and ending in 

July but with a peak in April-May 

(Charles et al., 2003) explaining the observation 

by Sardá et al. (1999) of a peak in 

seasonal abundance between May-June in 

the same region. The planktonic life is ~3 

weeks long with metamorphosis being 

completed quickly post-settlement 

(Charles et al., 2003). Ditrupa arietina lives 

for ~2 years, with most growth and sexual 

maturity being achieved during the 

first year (Medernach et al., 2000). 

Hence, although much is known 

about the biology of Ditrupa arietina in the 

Mediterranean, there is nothing known 

about it in the Açores, Portugal, save for 

the paper by Morton & Britton (1995) 

mentioned above. Morton & Britton 

(2000) also showed that the marine flora 

and fauna of the Açores have the 

strongest biogeographic links with the 

Mediterranean. The aim of the present 

paper was thus to determine if, as in the 

Mediterranean, D. arietina occupied a particular 

and similar depth zone. We also 

wished to investigate if there were any 

depth distributions in the Açores between 

living individuals of D. arietina and the 

empty tubes occupied by Aspidosiphon 

muelleri. Finally, it is known that in the 

Mediterranean, tubes of D. arietina, both 

occupied and unoccupied, are colonized 

by an encrusting fauna the major components 

of which are bryozoans (Gambi & 

Jerace, 1997). We wanted to see if this 

associated fauna was replicated in the 

Açores, or would it be, as with other taxa, 

much reduced (Morton & Britton, 2000). 

Finally, we wanted to determine if the 

tube of D. arietina had a life history, once 

the animal that had secreted it died. That 

is, it is known that the tube becomes occupied 

by Aspidosiphon muelleri (Morton & 

Britton, 2000), but what is the complete 

life history of the tube?

MORTON & SALVADOR: THE BIOLOGY OF DITRUPA ARIETINA 147 

TAXONOMIC NOTE 

Most authors, for example, Nelson- 

Smith & Gee (1966), follow Fauvel (1953) 

and recognize a single, worldwide, 

species of Ditrupa, that is, D. arietina (O.F. 

Müller, 1776). ten Hove & Smith (1990), 

however, justified the recognition of a 

separate species, D. gracillima Grube, 

1878, from the Indo-Pacific and deep 

water ecophenotypes of which had previously 

been identified as D. arietina var. 

monilifera. 


For eleven days from 17 - 28 July 2006, 

the sea bed on the southern coast of the 

island of São Miguel, Açores, was sampled 

using a benthic box dredge at six stations 

to the east and west of the islet of 

Ilhéu de Vila Franca do Campo. The stations 

were designated E1, E2, E3 and W1, 

W2 and W3 and located at approximate 

depths of 100 (E1 and W1), 200 (E2 and 

W2) and 250 metres (E3 and W3) (Figure 

1). The reader is referred to Martins et al. 

FIGURE 1. A map of the southern coast of São Miguel Island, showing the six sampling sites to 

the east (E1- E3) and west (W1 - W3) of Ilhéu de Vila Franca do Campo.

148 AÇOREANA 

2009, Sup. 6: 145-156 

(2009) for a full description of the stations 

and their accurate locations. Dredges 

were towed at a standard speed (5 knots) 

for ten minutes and on reaching the surface 

sieved using seawater through a 1 

mm mesh sieve. 

Two 500 ml sub-samples of sediment 

were removed from each dredge sample 

and sorted using dissecting binocular 

microscopes. All living individuals of 

Ditrupa arietina were collected from the 

first sample and the lengths of their tubes 

measured to the nearest 1 mm using a dissecting 

microscope (x10) with a 1 mm 

graduated scale. From these data, it has 

been possible to construct histograms of 

the population structure of D. arietina at 

the six sampling locations (Figure 2). The 

living individuals were also used in simple 

studies of burrowing behaviour. 

All empty tubes and fragments of 

Ditrupa arietina were removed from the 

second sample and measured along their 

greatest lengths to the nearest 1 mm using 

a dissecting microscope (x10) with a graduated 

scale. Each tube and fragment was 

then examined and divided into categories. 

These were: (i), those individuals 

occupied by Aspidosiphon muelleri; (ii), 

those with either anterior or posterior 

encrustations of attached biota and (iii), 

those with a drill hole. Some D. arietina 

tubes that had drill holes were also occupied 

by A. muelleri and had epibiont 

encrustations. Such individuals were 

placed in their own sub-categories. 

Data analysis 

One-factor analysis of variance 

(ANOVA) on untransformed data with a 

statistical significance criterion set at p = 

0.05 was conducted as the statistical technique 

used to examine for any differences 

in the datasets between the six sampling 

stations and hence depth. Any significant 

results obtained by the ANOVA were dif- 

FIGURE 2. Histograms showing the length frequency distribution of living Ditrupa arietina individuals 

at the six stations to the east (E1 - E3) and west (W1 - W3) of Ilhéu de Vila Franca do 

Campo, São Miguel, Açores.


ferentiated using a post-hoc Student’s 

Newman-Kuels (SNK) test to identify 

where the detected differences lay (Zar, 

1984). The data were analyzed using SAS 

(Version 9.1.3). 

RESULTS 

Statistical analyses 

Figure 2 are histograms showing the 

length frequency distribution of living 

Ditrupa arietina individuals at the six stations 

to the east (E1 - E3) and west (W1 - 

W3) of Ilhéu de Vila Franca do Campo, 

São Miguel, Açores. The results of a oneway 

ANOVA on these living individuals 

of D. arietina collected from the three sampling 

depths were significantly different 

(F=68.55; p=0.0031). The results of a posthoc 

Student’s Newman-Keuls test further 

indicated that the numbers of living individuals 

were significantly higher (p>0.05) 

at both the east (E2) and west (W2) ~200 

metre depth stations. 

Figure 3 are histograms showing the 

length frequency distributions of all tubes 

no longer occupied by Ditrupa arietina 

and, at the six stations to the east (E1 - E3) 

and west (W1 - W3) of Ilhéu de Vila 

Franca do Campo, São Miguel, Açores. 

The results of a second one-way ANOVA 

on the dataset indicated that there was no 

significant difference in the total numbers 

of Ditrupa arietina tubes collected at the 

sample depths of ~100, ~200 and ~250 

metres (F=0.36; p=0.7250). Similarly, there 

were no significant differences between 

stations and depths in the incidences of 

tubes that (i), were occupied by 

Aspidosiphon muelleri; (ii), possessed 

encrusting bryozoans and other attached 

organisms either anteriorly or posteriorly, 

nor (iii), in the incidences of predated, 

that is, drilled tubes. A more detailed 

study of the drill holes in the tubes D. 

arietina is reported upon by Morton & 

Harper (2009). 

Behaviour 

When living individuals of Ditrupa 

arietina and tubes occupied by 

Aspidosiphon muelleri were placed on the 

surface of samples of natural sediment 

obtained in the dredges, they both 

FIGURE 3. Histograms showing the length frequency distributions of all Ditrupa arietina tubes 

either empty or occupied by Aspidosiphon muelleri (plus tube fragments) at the six stations to the 

east (E1 - E3) and west (W1 - W3) of Ilhéu de Vila Franca do Campo, São Miguel, Açores.

150 AÇOREANA 

2009, Sup. 6: 145-156 

attempted to re-burrow. Ditrupa arietina 

burrowed with the posterior end of the 

tube down, so that the serpulid’s crown of 

tentacles projected above the sediment 

surface. Conversely, A. muelleri burrowed 

anterior end down so that its head was 

within the sediment. 

Epibionts 

Table 1 gives a list of the epibionts collected 

and identified from the tubes of 

Ditrupa arietina. The most obvious 

epibiont was the cemented serpulid 

Hydroides elegans (Haswell, 1883) that has 

probably been introduced into Açorean 

waters in historical times, possibly 

attached to boats or in ballast waters 

(Morton & Britton, 2000). Other epibionts 

included three species of foraminiferans, 

that is, the bright red Miniacina miniacea 

Pallas, 1776, Cassidulina obtusa 

Williamson, 1858 and Elphidium crispum 

(Linnaeus, 1767), and a number of 

species, mostly unidentifiable, of bryozoans. 

Also present were the egg capsules 

of gastropods (all unidentifiable). 

All the species identified in Table 1 have a 

north-eastern Atlantic/Mediterranean distribution, 

except for M. miniacea which 

occurs in the warmer central Atlantic 

from the Mediterranean to the Caribbean. 

Life history of the Ditrupa arietina tube 

The suggested life history of the tube 

of Ditrupa arietina is illustrated in Figure 

4. A living individual of D. arietina is 

illustrated in Figure 4A. The tube is cylindrical, 

slightly curved and resembles an 

elephant’s tusk or, more appropriately, a 

scaphopod, and anteriorly swollen 

although the mouth typically narrows 

again at its anterior extremity. There are 

numerous constrictions, or flanges, to the 

shell that is also variably patterned with 

circlets of orange-brown pigmentation. 

Because the contained worm is bright red, 

the tubes of living individuals are also 

redder than their empty counterparts. 

The 19 branchial filaments are also bright 

red (sometimes red banded) and the operculum 

is a membranous cup or funnel 

closed distally by a flat, brownish, chiti- 

TABLE 1. A list of encrusting species herein recorded as attached to the tubes of Ditrupa arietina 

from the six stations to the east (E1 - E3) and west (W1 - W3) of Ilhéu de Vila Franca do Campo. 

São Miguel, Açores. 

Phylum Species Notes 

Foraminifera Miniacina miniacea Pallas, 1776 Mediterranean, Caribbean (down to 2000 metres) 

Cassidulina obtusa Williamson, Northern Europe, Mediterranean, Canaries 

1858 

Elphidium crispum (Linnaeus, Mediterranean, Gulf of Cadiz, West Africa 

1767) 

Gymnolaemata: Celleporina hassalli (Johnston, British Isles 

Cheilostomata 1847) 

Cheilostoma 1 

Cheilostoma 2 

Stenolaemata: Crisia cf. eburnea (Linnaeus, Northern Europe, British Isles 

Cyclostomata 1758) 

Tervia irregularis (Meneghini, Northern Europe, Mediterranean 

1844) 

Disporella spp. 

Cyclostome 1 

Cyclostome 2 

Cyclostome 3 

Polychaeta: Hydroides elegans (Haswell, Near cosmopolitan; unintentionally introduced (Morton 

Serpulidae 1883) 

& Britton, 2000)


FIGURE 4. Ditrupa arietina. A, A living individual in its tube and (A¹) in its life position in the sediment; 

B, the anterior end of a tube with encrusting organisms; C, a drill hole (probably) made by 

the naticid Natica prietoi; D, an empty, drilled and anteriorly encrusted, tube occupied by 

Aspidosiphon muelleri and (D¹) in its life position in the sediment; E, an anteriorly encrusted tube 

occupied by A. muelleri; F, a fragment of unoccupied tube colonized by encrusting organisms and 

F, a fragment of tube.

152 AÇOREANA 

2009, Sup. 6: 145-156 

nous plate thickened in the centre to form 

a boss. Living individuals of D. arietina 

(Figure 4, A¹) live in the sediment, posterior 

end down and with the swollen anterior 

end situated above the surface. The 

anterior end of the tube is often encrusted 

with epibionts (Figure 4, B). Figure 4C 

illustrates a drill hole probably made by 

the naticid Natica prietoi Hidalgo, 1873 

(see Morton & Harper, 2009). Following 

death, the tube of D. arietina is occupied 

by Aspidosiphon muelleri (Figure 4D) at 

which time the anterior epibionts also die 

because the sipunculan lives head down 

in the sediment (illustrated in Figure 4 

[D¹]). With the posterior end of the tube 

now projecting above the sediment surface, 

it becomes encrusted by epibionts 

(Figure 4E), such that the tube becomes 

more eroded with age (Figure 4F) until 

only fragments remain (Figure 4G). As 

noted above we could obtain no statistically 

different station and hence depth 

distributions in the various categories of 

D. arietina tubes either occupied by A 

muelleri or encrusted either anteriorly or 

posteriorly. It thus seems that upon the 

death of D. arietina, its tube becomes 

widely distributed throughout the 

Açorean offshore seabed, probably by 

water movements acting upon it. 

DISCUSSION 

Labrune et al. (2007) analyzed the softbottom 

polychaete assemblages of the 

Gulf of Lions in the northwest 

Mediterranean and showed that at virtually 

stations associated with “littoral” fine 

muds and with depths of between 10–20 

metres the fauna was characterized by 

high abundances and high biomasses due 

to the presence of large numbers of 

Ditrupa arietina. Similarly, Cosentino & 

Giacobbe (2006) showed that, also in the 

Mediterranean, maximum mollusc/polychaete 

diversities (H’) occurred at depths 

of between 10–20 metres but that this 

decreased beyond this corresponding to 

the peak in the core population of D. arietina 

at depths of between 30-40 metres 

and abundances of >500 individuals·250·cm². 

Sardá et al. (2000) demonstrated 

that following sand extraction at 

depths of between 10–30 metres in the 

Tordera River, Bay of Blanes in the northwest 

Mediterranean, numbers of D. arietina 

initially remained stable, but numbers 

rose sharply during the following spring 

and autumn possibly due to re-colonization 

of the dredged sea bed. 

In May, that is, spring, coinciding with 

the periods of greatest abundances, 

growth and reproductive period in 

Ditrupa arietina, individuals spent only 

25% of the time feeding compared with 

50% of the time during the rest of the year 

(Jordana et al., 2000). The size spectrum 

of particles filtered by D. arietina was 

rather large and ranged between 1µm- 

50µm with planktonic and benthic 

diatoms, haptophytes, bacteria and 

cyanobacteria being collected and 

absorbed at efficiencies of 84.7, 70.9, 72.3 

and 63.7%, respectively (Jordana et al., 

2001a). Ditrupa arietina selectively filters 

picoplankton this accounting for, on a 

yearly basis, 15% of ingested chlorophylla 

with 95% of this figure being accounted 

for by picoeukaryotes (Jordana et al., 

2001b). The maximum weight specific 

clearance rate for D. arietina was 15.7 

ml.hour¯¹·mg·¯¹, about seven times less 

than for the polychaete Euchone papillosa 

(M. Sars, 1851), because of the latter’s relatively 

larger tentacle crown (Riisgård et 

al. 2002). 

From the above research in the 

Mediterranean, two important facts 

emerge that have relevance to Ditrupa arietina 

in the Açores. First, that D. arietina 

occurs at a relatively shallow depth of 

between 10-30 metres and, second, being 

a benthic suspension feeder it assists in


the clarification of the water column especially 

when the species occurs in large 

numbers. This study demonstrates that 

in the Açores, D. arietina occurs at a depth 

of ~ 200 metres. This distinctive difference 

in depths may be related to light, 

that is, in the Açores, oceanic waters allow 

colonization to a greater depth than in the 

(recently) more turbid Mediterranean. 

Or, such a difference may be related to 

food availability in the form of picoplankton 

that D. arietina feeds on, since this is 

likely to be less abundant in the Açores, 

thereby, somehow, influencing depth 

preference. It is also possible that the two 

factors may be acting synergistically in 

the Açores and/or in concert with other 

factors such as sediment grain size and 

disturbance. 

Morton (1990) studied the bivalve 

Ervilia castanea (Montagu, 1803) collected 

offshore from Vila Franca do Campo, São 

Miguel, and recorded it from depths of 

between 0-40 metres although the species 

is known to occur from ~800-1800 metres 

in the Açores and 130 metres in the 

Canary Islands (Smith, 1885), but possibly 

only as empty shells. This study of 

Ditrupa arietina collected few living E. castanea 

from the depths sampled, but many 

empty valves. Hence, in the Açores, the 

deeply shelving continental shelf (Figure 

1), appears to be characterized at two 

depths by two endobenthic suspension 

feeding species: the shallower living (0- 

~100 metres) bivalve Ervilia castanea and 

the deeper residing serpulid (~100 –250 

metres) D. arietina. 

Gambi et al. (1996) analyzed polychaete 

community structure at 32 stations 

distributed from 2–105 metres depth in 

the southern Tyrrhenian Sea (Italy) and 

showed that Ditrupa arietina was dominant 

at depths of between 26-30 metres 

and enhanced the spatial complexity, 

species composition and diversity and 

community structure of the habitat by 

virtue of the range of diversity of 

epibionts attached to its tube. Gambi & 

Jerace (1997) analyzed the epibionts on D. 

arietina tubes from three bays in the 

southern Tyrrhenian Sea and identified a 

wide range of species, including 

foraminiferans, sponges, hydroids, bryozoans 

and tubiculous polychaetes. 

Epibiosis percentages varied from site to 

site and between years. This study of the 

epibionts attached to the tubes of D. arietina 

identified a number of species 

including, most dominantly, the cemented 

serpulid Hydroides elegans (Haswell, 

1883). Other epibionts included three 

species of foraminiferans and a number of 

species of bryozoans. 

The only predated tubes of Ditrupa 

arietina that could be identified in this 

study appeared to have been drilled by 

the prosobranch naticid gastropod Natica 

prietoi (see Morton & Harper, 2009). 

Ditrupa arietina lives for ~2 years 

(Medernach et al., 2000), with most 

growth occurring during the first year. 

As Morton & Harper (2009) show, tubes 

of many sizes are attacked by the naticid 

although, generally, larger ones were 

favoured. Such a generalization is probably 

related to the relationship between 

predator and prey sizes, but if it is true 

that larger individuals are generally preferred 

by N. prietoi, then clearly, these are 

adults, sexual maturity also occurring in 

the first year of life (Medernach et al., 

2000), so that post reproduction the tube 

becomes available for colonization by 

Aspidosiphon muelleri, anterior epibionts 

then die, but further epibiotic organisms 

can now colonize the posterior end of the 

tube and slowly the tube degenerates. 

Subsequent to the death of the original 

inhabitant, the tube of Ditrupa arietina 

provides secondary accommodation for 

Aspidosiphon muelleri and throughout its 

life history it functions as an inhabitable 

substratum and hence a locally important

154 AÇOREANA 

2009, Sup. 6: 145-156 

Açorean habitat for a suite of epibionts. It 

is also apparent that D. arietina is an overwhelmingly 

dominant and therefore highly 

significant component of the Açorean 

endobenthos at depths of ~200 metres, 

mirroring the significance of the bivalve 

Ervilia castanea at generally shallower 

depths (Morton, 1990). It is finally clear, as 

this study now suggests, that the ecological 

significance of these two species is in 

urgent need of further, much more 

detailed study, especially with regard to 

their role in the productivity of the inshore 

Açorean continental shelf seabed. 


The authors are grateful to Prof. A.M. 

Frias Martins (University of the Açores) for 

funding this research and for much practical 

help and warm hospitality during their 

stay on São Miguel. We are also grateful to 

a number of University of the Açores students 

who helped with sample sorting and 

to Dr. K.F. Leung (Environmental 

Protection Department, Hong Kong SAR 

Government, China) for statistical advice 

and help. Mary Spencer Jones (Natural 

History Museum, London) is thanked for 

identifying the bryozoans. 


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DRILLING PREDATION UPON DITRUPA ARIETINA (POLYCHAETA: 

SERPULIDAE) FROM THE MID-ATLANTIC AÇORES, PORTUGAL 

Brian Morton & E.M. Harper 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK. E-mail: 

prof_bmorton@hotmail.co.uk 

Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, UK. E-mail: 

emh21@cam.ac.uk 

ABSTRACT 

A small, but significant proportion of the empty tubes of the free-living serpulid 

Ditrupa arietina dredged from the seabed off São Miguel, Açores, were punctured by small 

round holes. Analysis of these holes shows that they are typically single, made from the 

outside of the tube and located preferentially, suggesting that they represent the work of 

a predator. The likely identity of the predator is discussed but based on the co-occurrence 

in the dredges and the fact that most of the holes have a distinctive countersunk morphology, 

it is suggested that they were made by the small naticid Natica prietoi. If so, this 

study is the first record of such a predator/prey relationship. 

RESUMO 

Uma pequena mas significativa porção de tubos do serpulídeo livre Ditrupa arietina 

dragados do fundo marinho ao largo de São Miguel, Açores, estavam perfurados por 

pequenos orifícios redondos. Análise desses orifícios mostra que eles são tipicamente 

singulares, feito a partir do exterior do tubo e localizados preferencialmente, sugerindo 

que representam o trabalho de um predador. Discute-se a identidade provável do 

predador mas, com base na co-ocorrência nas dragagens e no facto de que a maioria dos 

orifícios possui uma distinta morfologia chanfrada, sugere-se que foram feitos pelo 

pequeno naticídeo Natica prietoi. Assim sendo, este estudo é o primeiro registo de tal 

relação predador/presa. 

INTRODUCTION 

Although polychaete worms are a 

major source of food for a number of 

predatory taxa including fish, gastropods 

and birds, most of our knowledge is 

focused on predation of the larger errant 

taxa. For example, amongst the predatory 

gastropods, representatives of the 

Muricidae, Columbellidae, Fasciolariidae, 

Vasidae and Buccinidae as well as many 

conoideans are specialist worm predators 

(Pearce & Thorson, 1967; Taylor, 1978a, b, 

1980; Taylor et al., 1980; Shimek, 1984; 

Taylor & Lewis, 1995). Similarly, many 

wading birds are specialist predators 

upon intertidal endobenthic polychaetes 

(Goss-Custard, 1975). 

In contrast, little appears to be known 

about predatory activities upon the tubedwelling 

Serpulidae (Polychaeta) despite 

their cosmopolitan occurrence and frequently 

high abundance. There is, however, 

observational evidence that gastropods 

(Taylor & Morton, 1996; Tan & 

Morton, 1998), fish (Bosence, 1979; 

Witman & Cooper, 1983) and crustaceans 

(Bosence, 1979) feed on serpulids and, in 

the case of the buccinid gastropod Engina 

armillata (Reeve, 1846) in Hong Kong, 

they are its prey of choice (Tan & Morton, 

1998). Additionally, the development of 

calcareous tubes, a well-developed 

“fright response” (Poloczanska et al., 

2004) and, in the case of encrusting 

species, an often cryptic habit suggest

158 AÇOREANA 

2009, Sup. 6: 157-165 

that the evolution of the Serpulidae may 

have been influenced by predation pressure. 

Part of the problem in recognising 

predatory activity on serpulid worms 

undoubtedly results from their typically 

cryptic lifestyles that make direct observations 

difficult. Indirect methods to 

detect predatory activity on serpulids 

must rely on analyses of the predator’s 

gut contents to recognise setae (Taylor & 

Morton, 1996) or the identification of any 

characteristic damage to the tube caused 

by a predator, although for many predators, 

for example E. armillata, which was 

recorded feeding via the prey’s aperture 

(Tan & Morton, 1998), no such characteristic 

damage results. 

In this paper we describe drill holes 

made in the tube of the endobenthic serpulid 

Ditrupa arietina (Müller, 1776) from 

the subtidal seabed off the island of São 

Miguel in the Açores (Portugal). We present 

evidence that these are the result of 

predation and further discuss evidence 

as to the likely culprit(s). 

Ditrupa arietina (Müller, 1776) 

Ditrupa arietina is a widespread 

endobenthic serpulid worm, common in 

the Mediterranean where it achieves 

high densities and there is evidence in 

some areas that its abundance is increasing 

(Gremare et al., 1998a, b; Bolam & 

Fernandes, 2002; Labrune et al., 2007). 

The worm secretes a curved, tuskshaped, 

calcareous tube up to 23 mm 

long and about 3 mm across at its widest 

point. It lives within the sediment, the 

narrowest posterior end down, with an 

anterior bulge (about 10% of the tube) 

exposed above the substratum. This 

anterior bulge houses the circlet of twenty 

feeding tentacles when they are 

retracted and the anterior aperture is 

sealed by a further tentacle bearing an 

operculum. Morton & Salvador (2009, 

figure 4) illustrate a living D. arietina in 

its life position in the sediment. 

The species has been recorded as a 

dominant member of soft-bottom communities 

at depths of 100–250 metres in 

the Açores where the tubes of dead individuals 

also provide domiciles for the 

sipunculan worm Aspidopsiphon muelleri 

Diesing, 1851 (Morton & Britton, 1995; 

Morton & Salvador, 2009). 


During July 2006, large numbers of 

the serpulid polychaete Ditrupa arietina 

were collected from depths of between 

50-200 metres off Vila Franca do Campo, 

São Miguel, Açores. The samples were 

collected and processed as reported in 

Morton & Salvador (2009). Following initial 

observations that some tubes were 

perforated by drill holes, the tubes of 

dead individuals were separated and 

inspected for evidence of such drilling. 

For each holed tube, the following data 

were collected: tube length to the nearest 

1 mm using vernier callipers, number of 

drillholes per tube and the morphology of 

the drillhole. Further observations were 

made on the morphology of the drillholes 

of a small number of specimens by scanning 

electron microscopy (SEM). These 

specimens were prepared by cleaning in 

an ultrasonic bath prior to mounting 

them, coated in gold, for SEM (JEOL 820 – 

University of Cambridge). 

Finally, the mean tube lengths plus 

standard deviations of intact living individuals 

of Ditrupa arientina and those 

with drill holes were compared using a 

t-test. 

RESULTS 

Holed individuals of Ditrupa arietina 

were identified from each of the six sampling 

stations, described by Martins et al.

MORTON & HARPER: DRILLING PREDATION UPON DITRUPA ARIETINA 159 

(2009), but no significant differences in 

numbers were obtained between them 

with regard to the incidence of drilled 

tubes (see Morton & Salvador, 2009 for 

details). In total 5,453 tubes of D. arietina 

were retrieved and examined and from 

which we identified 104 fragments and 

intact empty tubes with holes in them, 

that is, approximately 1.9% of all empty 

tubes examined. The vast majority of the 

tubes were perforated by a single drillhole 

but a few had two or even three complete 

holes. No incomplete holes were 

observed nor any that showed signs of 

repair or healing by the occupant. All 

were drilled perpendicular to the surface 

of the tube and were clearly produced 

from the outside, as evidenced by a wider 

outer diameter. 

All holes in the tubes of Ditrupa arietina 

were small, with an outer diameter of 

less than 700 µm. In outline, the perforations 

were mostly circular (Figure 1A, B) 

but a few were more elliptical (Figure 1C). 

The outer edges of the drillholes were 

clearly defined, and SEM observations of 

the surface of the tube adjacent to the hole 

show no obvious signs of either dissolution 

or rasping. The inner perforations 

were rather more ragged and smaller 

than the outer diameter. Most of the holes 

we have examined had curved walls leading 

to a countersunk morphology, 

although some (Figure 1D) were more 

straight-sided. 

The positions of drill holes in the 

tubes of Ditrupa arietina are shown in 

Figure 2. It is clear that most are located 

in the mid portion of the tube, below the 

anterior bulge. Although they occur all 

around the circumference of the tube, the 

distribution of the holes is not uniform, 

there being a clear preference for the concave 

aspect. 

The length distributions of a sub-sample 

(n = 376) of living Ditrupa arietina 

recorded by Morton & Salvador (2009) are 

FIGURE 1. Scanning electron micrographs of a 

variety of drill holes in the tubes of Ditrupa arietina 

collected from the seabed offshore from 

Vila Franca do Campo, São Miguel, Açores, in 

2006. Scale bars represent 100 µm for A and B 

and 200 µm C and D. 

FIGURE 2. Histograms showing the size distributions 

(tube lengths) of living (white) and 

drilled (black) individuals of Ditrupa arietina 

collected from the seabed offshore from Vila 

Franca do Campo, São Miguel, Açores, in 2006. 

plotted together with the same data for 

drilled individuals in Figure 3. Although 

virtually all sizes of D. arietina tubes were 

drilled (from 5-21 mm tube lengths), the 

mean length of the drilled D. arietina

160 AÇOREANA 

2009, Sup. 6: 157-165 

tubes (n = 102) was 12.7 (S.D. 4.3) whereas 

that of the tubes occupied by living D. 

arietina (n = 378) was 14.9 (S.D. 2.8). The 

result of the t-test showed that the means 

of the two samples were significantly different 

(p =


support of this contention, no marginellids 

are reported from the Açorean fauna 

and the trawls in which the drilled D. arietina 

were collected contained a number 

of individuals of Natica prietoi Hidalgo, 

1873, formerly identified as Natica adansoni 

de Blainville, 1825 (see Gubbioli & 

Nofroni, 1998), and no nudibranchs of 

any species. Further evidence to support 

a naticid origin for the drill holes is the 

preferred position in an area of the tube 

that would be embedded in the sediment 

and hence accessible only to an endobenthic 

predator. Naticids are, moreover, 

known to be highly specific with regard 

to the position on the prey’s exoskeleton 

chosen for attack (Thomas, 1976; Kabat, 

1990). 

Martins et al. (2009), although recording 

one living specimen of Natica dilwinii 

Payraudeau, 1826, dredged at 300 m 

depth off Vila Franca do Campo, consider 

that Natica prietoi is definitively the most 

common naticid recorded alive from the 

Açores, as described in this study. 

Notwithstanding, Morton (1990) recorded 

living Natica intricata (Donovan, 1804) 

from Ilhéu de Vila Franca, São Miguel, 

and was reported by this author to feed, 

by drilling, on the tellinoidean bivalve 

Ervilia castanea Montagu, 1803. Other 

naticids have also been reported to occur 

in the Açores (Table 1), but these are generally 

records of empty shells. Euspira 

pulchella (Risso, 1826) has been recorded 

from the seabed at 2,000 metres depth in 

the Açores by Ávila et al. (1998). Natica 

variabilis Recluz in Reeve, 1855 was 

recorded from the offshore sea-bed of São 

Miguel by Morton & Britton (1995) and 

by Ávila et al. (1998). Natica alderi Forbes, 

1838 was recorded from São Jorge by 

Morton (1967) and, finally, Ávila et al. 

(1998) records Polinices lacteus (Guilding, 

1834). 

One further point which may have 

some bearing on these disparate records 

(except for Natica prietoi that, as this study 

shows, is numerous on the offshore seabed 

at ~ 200 metres depth), is that the 

Caribbean species Natica canrena 

(Linnaeus, 1758) has been recorded from 

the Açores by Morton & Britton (1998), 

Morton et al. (1998) and Ávila et al. (1998), 

an occurrence explained by the former 

authors as a chance event resulting from 

this species having a teleplanic larva 

(Laursen, 1981). The same argument has 

been presented for the occasional recorded 

occurrence of the North African 

Polinices lacteus in the Açores (Laursen, 

1981; Ávila et al., 1998; Morton et al., 

1998). 

Natica prietoi is a small species (~ 10 

mm in shell diameter) and was the only 

naticid collected along with the samples, 

in the same trawls, of Ditrupa arietina 

reported upon by Morton & Salvador 

(2009) and herein. It seems, therefore, 

that Natica prietoi is a plausible candidate 

TABLE 1. A species list of Naticidae recorded from the Açores 

Species Location References 

Natica prietoi Hidalgo, 1873 São Miguel Martins et al., 2009; this study. 

Natica dillwinii Payraudeau, 1826 São Miguel Martins et al., 2009. 

Natica adansoni de Blainville, 1825 São Miguel Ávila et al., 1998. 

Natica intricata (Donovan, 1804) São Miguel Morton, 1990; Ávila et al., 1998. 

Euspira pulchella (Risso, 1826) 2,000 metres Ávila et al., 1998 

Natica variabilis Recluz in Reeve, 1855 São Miguel Morton & Britton, 1995; Ávila et al., 1998. 

Natica alderi (Forbes, 1838) São Jorge Morton, 1967. 

Natica canrena (Linnaeus 1758) 

Natica canrena (Linnaeus 1758) 

Laursen, 1981; Morton et al., 1998; Ávila et 

Polinices lacteus (Guilding, 1834) 

Polinices lacteus (Guilding, 1834) 

al., 1998; Morton & Britton, 2000. 

Laursen, 1981; Ávila et al., 1998; Morton et 

al., 1998.

162 AÇOREANA 

2009, Sup. 6: 157-165 

for the identity of the predator responsible 

for the holes in D. arietina. 

If the above conclusion is correct, this 

is the first record of a naticid attacking a 

serpulid polychaete. We are aware of 

only one other report, by Paine (1963), 

who observed a single individual of 

Neverita duplicata (Say, 1822) attacking the 

sabellid Owenia fusiformis delle Chiaje, 

1841. 

The incidence of multiple, complete, 

drillholes in a small number of tubes is 

curious (Fig. 1D). Simultaneous drilling 

by multiple individuals, although reported 

upon for muricids (Brown & 

Alexander, 1994; Taylor & Morton, 1996), 

is unlikely in naticids because of the manner 

in which such predators handle their 

prey that is (albeit in other species) 

enveloped by the foot of the attacker 

which would preclude access by another 

would-be predator. It is possible, however, 

that they represent sequential attacks, 

the first two of which were failures 

although they were evidently complete, 

in the sense that they perforate the full 

thickness of the tube, they may have been 

non-functional (Kitchell et al., 1986). 

Another possibility is that they were 

caused by another predatory taxon and 

we do note that the multiple holes illustrated 

in Figure 1D are more straightsided 

than are most of the others and thus 

more like muricid drill holes (Carriker, 

1981). Moreover, muricids are known to 

feed in clusters of conspecifics and sympatric 

species (Abe, 1980; Tong, 1986; 

Taylor & Morton, 1996). This raises the 

possibility of there being a muricid predator 

on the offshore seabed that also feeds 

on Ditrupa arietina. 

Notwithstanding, this study appears 

to be the first to record significant drilling 

predation upon Ditrupa arietina although 

Tan & Morton (1998) record the buccinid 

Engina armillata aperturally (but never 

drilling) attacking cemented serpulids in 

Hong Kong. In the Açores, the only significant 

predator captured in the dredges 

with D. arietina was the naticid Natica 

adansoni. However, we conclude, in the 

light of this study, that further research is 

needed to establish the predator-prey 

relationship(s) on and in the Açorean offshore 

seabed. 


The authors are grateful to Prof. A.M. 

de Frias Martins, University of the 

Açores, for organising the workshop at 

which this research was undertaken and 

for providing the boat facilities allowing 

collection of the samples herein reported 

upon. Ms. Andreia Salvador (The 

Natural History Museum, London) is 

thanked for assistance with the sorting of 

the samples. 


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THE PYCNOGONIDS (ARTHROPODA: PYCNOGONIDA) OF SÃO MIGUEL, 

AZORES, WITH DESCRIPTION OF A NEW SPECIES OF ANOPLODACTYLUS 

WILSON, 1878 (PHOXICHILIDIIDAE) 

Roger N. Bamber 1 & Ana Cristina Costa 2 

1 

The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. 

e-mail: roger.bamber@artoo.co.uk 

2 

CIBIO-Pólo Açores, Department of Biology, University of the Azores, 9501-801 Ponta Delgada, São Miguel, Azores, Portugal 

ABSTRACT 

During the Third International Workshop of Malacology and Marine Biology in São 

Miguel, Azores, in July 2006, sampling of algae, and of the littoral and sublittoral benthos 

was undertaken in order to characterize the smaller marine arthropod fauna of this region, 

including pycnogonids. In the event, 50 specimens of Pycnogonida, representing six 

species and including a new species of Anoplodactylus, were collected. In addition, previous 

pycnogonid material collected around São Miguel in 1996 and 1997 was analyzed, 

from 112 samples of which a further 3705 pycnogonid specimens were identified, representing 

eight species, three additional to the above six. All of this material is described 

below. The zoogeography of the fifteen species now recorded from the Azores is analyzed, 

and the likely origins of the Azorean pycnogonid fauna are discussed. 

RESUMO 

Durante o 3º Workshop Internacional de Malacologia e Biologia Marinha em São 

Miguel, Açores, em Julho de 2006, realizaram-se amostragens de algas e o bentos do litoral 

e sublitoral com vista a caracterizar a fauna de artrópodes marinhos de reduzidas 

dimensões da região, incluindo picnogonídeos. Na ocasião foram recolhidos 50 

exemplares de Pycnogonida, representando seis espécies e incluindo uma nova espécie de 

Anoplodactylus. Para além disso, analisou-se material de picnogonídeos, previamente 

recolhido em São Miguel em 1996 e 1997, de 112 amostragens das quais mais 3705 

espécimes de picnogonídeos foram identificados, representando oito espécies, três das 

quais para além das seis atrás referidas. Todo este material é descrito em seguida. 

Analisa-se a zoogeografia das quinze espécies agora registadas para os Açores e discutese 

a provável origem da fauna de picnogonídeos Açorianos. 

INTRODUCTION 

Pycnogonids are a group of the arthropods 

with minimal dispersive ability 

(Bamber, 1998); the larvae are not planktonic, 

and there are only limited examples 

of adults swimming, and then not as a 

directional migratory movement (Arnaud 

& Bamber, 1987). A few genera, such as 

the deep-water (Bathypallenopsis) and the 

generally shallow-water Anoplodactylus, 

are known to live upon medusae, and 

thus obtain passive dispersion in the 

plankton (Lebour, 1916; Arnaud & 

Bamber, 1987; Bamber, 2002). Some 

species are known to have spread in fouling 

communities on ship’s hulls (Krapp & 

Sconfietti, 1983; Bamber, 1985). 

The Azores are a group of islands 

somewhat isolated in the north-east 

Atlantic, lying adjacent to the Mid- 

Atlantic Ridge some 1600 km west of 

Portugal and 1730 km southeast of 

Newfoundland. The main surface water 

currents reaching the archipelago bring 

waters from two directions: the Azores 

drift, a diffuse southerly arm of the Gulf 

Stream breaking off from the North

168 AÇOREANA 

2009, Sup. 6: 167-182 

Atlantic Drift supplies water from the 

Americas, while the somewhat less-significant 

western eddies of the Canary 

Current bring waters from Spain and 

North Africa; below these, the midwater 

current brings warm, hyperhaline water 

from the Mediterranean outflow (Morton 

et al., 1998; Gofas, 1990). For the passively-dispersing 

Pycnogonida, therefore, it is 

of some interest to determine the suite of 

species which has colonized this archipelago, 

and their provenance. 

The only previous specific study of 

the Pycnogonida of the Azores was that 

reported by Arnaud (1974), who recorded 

ten species from shallow waters, including 

the discovery of the currently 

Azorean-endemic Achelia anomala, as well 

as four species from waters of >500 m 

depth. Other previous records of pycnogonids 

in the Azores were noted by 

Loman (1912), Bouvier (1917), Arnaud 

(1978), Stock (1971; 1990) and Bamber 

(2002), although a number of these were 

from abyssal depths. Munilla & Sanchez 

(1988) reported on pycnogonids from the 

Canary Islands. 

The twelve species recorded previously 

in litt. from the Azores, from depths 

shallower than 100 m, are: 

Family Ammotheidae: 

Achelia anomala Arnaud, 1974 (currently 

endemic); 

A. echinata Hodge, 1864; 

Tanystylum orbiculare Wilson, 1878. 

Family Phoxichilidiidae: 

Anoplodactylus angulatus (Dohrn 1881); 

A. maritimus Hodgson, 1915; 

A. petiolatus (Kroyer, 1844); 

A. pygmaeus (Hodge, 1864). 

Family Callipallenidae: 

Callipallene emaciata (Dohrn, 1881) 

Family Endeidae: 

Endeis spinosa (Montagu, 1808); 

E. straughani Clark, 1970 (see Stock, 

1990; an Australian species, possibly 

a misidentification, or an 

immigrant via ships’-hull-fouling, 

see Bamber, 1979 [as E. picta]) 

Family Rhynchothoracidae: 

Rhynchothorax philopsammum Hedgpeth, 

1951; 

R. monnioti Arnaud 1974. 

No members of the speciose family 

Nymphonidae have yet been recorded. 

During the Third International 


Biology at Vila Franca do Campo, on the 

island of São Miguel, Azores, in July 2006, 

sampling of the littoral and sublittoral 

benthos around São Miguel was undertaken 

in order to characterize the smaller 

marine arthropod fauna of this region, 

including pycnogonids. In the event, 50 

specimens representing six species, one 

new to science, were collected. In addition, 

a previous collection from around 

the island was made available, from 112 

samples from which a further 3705 pycnogonid 

specimens were identified, representing 

eight species, three additional 

to the above six. All of this material is 

described below. 

MATERIAL AND METHODS 

The present material comes from two 

sources. During the Workshop at Vila 

Franca do Campo in July 2006, a number 

of littoral and infralittoral habitats on the 

island of São Miguel were sampled for 

pycnogonids, including crevice habitats, 

macroalgae and soft sediments. The principal 

sampling areas were the littoral sediments, 

rocks and algae below the Clube 

Naval building (the old Vila Franca do 

Campo abattoir); the sediments and algae 

within the flooded crater of the Ilhéu de 

Vila Franca; and the soft sediments off 

Vila Franca do Campo (ca N37° 43’ W 

25°25’), from 12 to 200 m depth. Offshore 

sediments were sampled using a 0.025 m 2

BAMBER & COSTA: PYCNOGONIDS FROM SÃO MIGUEL 169 

van Veen grab and various dredges. All 

samples were washed through a 0.5 mm 

mesh sieve, and specimens sorted alive. 

Some of these specimens were fixed in 

absolute ethanol to allow DNA analysis. 

Extensive material collected in 1996 

and 1997, from infralittoral rocky-substratum 

sites around São Miguel, was also 

analysed in detail. Replicate samples of 

algae (Stypocaulon scoparia, Halopteris filicina 

and Zonaria tournefortii) were collected 

by SCUBA diving at depths from between 

5 and 16 m. Details of the sampling and 

protocols are given by Costa & Ávila 

(2001). The sampling sites, anti-clockwise 

around the Island from the north-west, 

were Mosteiros, Ponta da Ferraria, Santa 

Clara , Pesqueiro, Emissário, Sinaga, 

Atalhada, Ribeira da Praia, Caloura, Porto 

de Vila Franca do Campo, Ilhéu de Vila 

Franca, Ribeira Quente, Faial da Terra, 

Nordeste, Porto Formoso, Ladeira de 

Velha, Ribeirinha, Lactoaçoreana, Cofaco 

and São Vicente. These sites variously 

represented exposed and sheltered 

shores, undisturbed, naturally disturbed 

(near shallow-water vent sites or stream 

mouths) and polluted shores. 

Voucher and type-material has been 

lodged in the collections of The Natural 

History Museum, London (NHM). The 

higher taxonomy is based on Arnaud & 

Bamber (1987). All measurements are 

axial, measured dorsally for the trunk 

and proboscis (trunk length being from 

the anterior margin of the cephalon to the 

posterior margin of the fourth lateral 

processes), laterally for oviger and legs. 

SYSTEMATICS 

Family Ammotheidae Dohrn, 1881 

Achelia echinata Hodge, 1864 

Figure 1A 

Material: 1 male, 2 females, 1 subadult, 

WVF011, northeastern side of Ilhéu de 

Vila Franca do Campo, 16 m, scuba dive 

collection by Gonçalo Calado, José Pedro 

Borges, Joana Xavier, Paola Rachello, 

Patrícia Madeira, 20 July 2006. 3 males, 4 

females, 2 subadults, WVF040, off Amora, 

Ponta Garça, São Miguel, Azores, N37° 

42’ 720” W25° 21’ 554”, 37.8 m depth, 

small dredge sample, 26 July 2006, coll. 

António de Frias Martins & Jerry 

Harasewych. 1 female, 1 subadult, littoral 

rock scrape, Vila Franca do Campo 

marina, 19 July 2006, coll. Kathe Jensen. 1 

male with eggs, Isl. 24.5, mid-lagoon 

algae, within the flooded crater of the 

Ilhéu de Vila Franca, 24 July 2006, coll. A. 

Salvador, R. Robbins & R.B. 

12 specimens, Mosteiros, 8 November 

1996; 139 specimens, Mosteiros, 17 June 

1997; 41 specimens, Ponta da Ferraria, 17 

June 1997; 53 specimens, Santa Clara, 7 

November 1996; 201 specimens, Santa 

Clara, 12 June 1997; 20 specimens, 

Pesqueiro, 7 November 1997; 21 

specimens, Emissário, 2 November 1996; 

237 specimens, Emissário, 12 June 1997; 

37 specimens, Sinaga, 30 May 1997; 1 

specimen, Atalhada, 25 October 1996; 246 

specimens, Atalhada, 30 May 1997; 32 

specimens, Ribeira da Praia, 25 October 

1996; 70 specimens, Ribeira da Praia, 16 

June 1997; 61 specimens, Caloura, 14 

October 1996; 119 specimens, Caloura, 26 

June 1997; 48 specimens, Porto de Vila 

Franca do Campo, 9 May 1997; 8 

specimens, Ilhéu de Vila Franca do 

Campo, 24 October 1996; 47 specimens, 

Ilhéu de Vila Franca do Campo, 16 June 

1997; 44 specimens, Ribeira Quente, 18 

June 1997; 5 specimens, Faial da Terra, 17 

October 1996; 59 specimens, Faial da 

Terra, 30 May 1997; 19 specimens, 

Nordeste, 21 November 1966; 143 

specimens, Nordeste, 2 May 1997; 36 

specimens, Porto Formoso, 30 October 

1996; 140 specimens, Porto Formoso, 19 

June 1997; 13 specimens, Ladeira da 

Velha, 19 June 1997; 16 specimens, 

Ribeirinha, 8 October 1996; 140

170 AÇOREANA 

2009, Sup. 6: 167-182 

specimens, Ribeirinha, 25 June 1997; 10 

specimens, Lactoaçoreana, 14 October 

1996; 76 specimens, Lactoaçoreana, 25 

June 1997; 5 specimens, Cofaco, 25 June 

1997; 13 specimens, São Vicente, 7 

October 1996; 18 specimens, São Vicente, 

6 May 1997; all coll. João Brum & A.C.C. 

Remarks: with a total of 2146 specimens 

from 110 samples listed above, Achelia 

echinata is easily the commonest shallowwater 

pycnogonid in the Azores (occurring, 

for example, at every sampling site 

in 1996 and 1997). In the light of the 

potential for cryptic sibling species in this 

taxon, specimens collected in alcohol 

were analysed for their 16s DNA to compare 

with specimens from England which 

had already been analysed as part of a 

larger project (Bamber et al., in prep.) and 

showed 100% agreement. All the 

Azorean material was collected in association 

with algae, on which, inter alia, this 

species is known to feed (Bamber & 

Davis, 1982). 

Ammothella longipes (Hodge, 1864) 

Figure 1B 

Ammothoa longipes Hodge, 1864 

Achelia longipes sens. auctt.; 

Achelia magnirostris (Dohrn, 1881) 

Material: 1 specimen, Atalhada, 30 May 

1997; 1 specimen, Ribeira da Praia, 16 



specimens, Ribeira Quente, 18 June 1997; 

31 specimens, Lactoaçoreana, 25 June 

1997; all coll. João Brum & A.C.C. 

Remarks: This species has been recorded 

from the Atlantic coasts of Europe 

(England being the type-locality), the 

Mediterranean and North Africa, as well 

as the Canaries (Sanchez & Munilla, 

1989), but had not been recorded previously 

from the Azores. It is readily distinguished 

from sympatric Achelia echinata 

by the fleshy, glabrous dorsodistal 

tubercles on the lateral processes, the lack 

of spinose tubercles on the lateral 

processes and coxae, and in having nine 

palp articles (A. echinata having eight). 

Family Phoxichilidiidae Sars, 1891 

Anoplodactylus amora sp. nov. 

Figure 2. 

Material: 1 male, holotype 

(NHM.2007.416), 1 male, 1 subadult 

female, paratypes (NHM.2007.417-418), 

VWF040, off Amora, Ponta Garça, São 

Miguel, Azores, N37° 42’ 720” W25° 21’ 

554”, 37.8 m depth, small dredge sample, 

26 July 2006, coll. António de Frias 

Martins & Jerry Harasewych; 1 subadult 

male, Vila Franca do Campo marina, São 

Miguel, Azores, littoral rock scraping, 19 

July 2006, coll. Kathe Jensen. 

2 males, paratypes (NHM.2007.419- 

420), Caloura, 26 June 1997; 1 male, 

paratype (NHM.2007.421), Ilhéu de Vila 

Franca do Campo, 16 June 1997; 1 

subadult male, 1 female, paratypes 

(NHM.2007.422-423), Faial da Terra, 30 

May 1997; all coll. João Brum & A.C.C. 

Description of male: habitus typical of the A. 

petiolatus form, trunk (Figure 2A, B) almost 

glabrous, with segment articulations 

between cephalon, trunk segment two and 

trunk segment three. Cephalon overhanging 

proboscis base to half proboscis length, 

lateral anterior margin with slight shoulders 

bearing short seta; dome-shaped ocular 

tubercle slightly taller than wide, without 

distal tubercle, with four pigmented eyes 

and laterodistal sensory pit; oviger attachment 

on mid-ventral surface of first lateral 

process. Lateral processes shorter than segment 

width, separated by less than their 

own diameter, with simple anterodistal seta 

and very small dorsodistal tubercle (little 

more than a swelling). Abdomen simple, 

sub-clavate, angled upward at about 45º, 

exceeding distal edge of coxa 1 of leg 4, sub-


FIGURE 1. A, Achelia echinata, entire, dorsal (modified after Sars, 1893); B, Ammothoella longipes, 

entire, dorsal.

172 AÇOREANA 

2009, Sup. 6: 167-182 

distally bearing pair of lateroventral spines 

and single mid-dorsal spine. 

Proboscis stout, parallel-sided, one-third 

as long as trunk. 

Chelifore (Figure 2C) scape of one article, 

slender, with sparse dorsal and distal 

setae; chela compact, fingers just shorter 

than palm, palm and moveable finger 

setose, cutting edges naked. Palp absent. 

Oviger (Figure 2D) of six articles, proximal 

article short, compact; article 2 (O2) 

three times as long as wide, with single dor- 

FIGURE 2. Anoplodactylus amora sp. nov., male holotype, A, lateral and B, dorsal; C, right chelifore; D, left 

oviger; E, third left leg; F, detail of distal articles of third leg; G, coxa 2 and femur of male A. petiolatus. 

Scale line = 1 mm for A and B, 0.4 mm for C and D, 0.8 mm for E, 0.3 mm for F, 0.5 mm for G.


sal and ventral setae; O3 longest, 2.3 times 

length of O2, slender, with sparse dorsal and 

ventral setae; O4 just shorter than O2, 

curved, with dorsodistal setae; O5 0.6 

times as long as O4, with numerous setae 

pointing proximally; O6 compact, acornshaped, 

half length of O5, with outer and 

ventral setae pointing proximally. 

Third leg (Figure 2E) elongate, not 

slender. Coxa 1 compact, with sparse distal 

setae, without dorsal tubercle; coxa 2 

almost three-times as long as coxa 1, 2.5 

times as long as wide, with pronounced 

ventrodistal genital spur; coxa 3 just more 

than half length of coxa 2. Femur stout, 

less than four times as long as wide, twice 

as long as coxa 2, setose, without conspicuous 

dorsodistal spur; cement-gland tube 

externally very short, arising at 0.6 the 

length of the femur, extending subcutaneously 

half way to proximal end of 

femur. Tibia 1 shorter than femur (0.9 

times femur length), setose as figured; 

tibia 2 five times as long as wide, as long 

as femur, more densely setose, longest 

dorsodistal seta not on a spur. Tarsus 

small, triangular, with numerous ventral 

and single dorsal setae. Propodus (Figure 

2F) with distinct heel, with two large 

proximal and five slender distal heel 

spines; sole with row of seven simple submarginal 

setae in distal two-thirds, seven 

short, recurved marginal spines and distal 

lamina for 30% of sole length. Main 

claw two-thirds as long as propodus, auxiliary 

claws small, lateral, slender, 0.15 

times length of main claw. 

Description of female: generally as male, 

but ovigerous legs absent, femur swollen; 

no ventral tubercles on proboscis. 

Measurements of holotype male (mm). – 

Trunk length: 1.91; trunk segment 2 

length: 0.29; width across 2nd lateral 

processes: 0.99; abdomen length: 0.42; 

proboscis length: 0.53. 

Lengths of oviger articles 1 to 6 

respectively: 0.13; 0.31; 0.73; 0.27; 0.17; 

0.09. 

Fourth leg, lengths of coxa 1: 0.24; 

coxa 2: 0.69; coxa 3: 0.38; femur: 1.40 

(width 0.36); tibia 1: 1.23; tibia 2: 1.40 

(width 0.28); tarsus: 0.12; propodus: 0.55; 

main claw: 0.37; auxiliary claw: 0.06. 

Etymology: named after the location on 

São Miguel off which the type-locality 

lies. 

Remarks: there are two previously 

described species of Anoplodactylus which 

have a subcutaneous proximal extension 

of the cement gland tube, viz. A. erectus 

Cole, 1904 from the Pacific (the Americas 

from Chile to British Columbia, Hawaii, 

Polynesia, Japan, Korea and Hong Kong) 

at depths of 0 to 90 m, and A. amoybios 

Bamber, 2004 from Atlantic Equatorial 

Africa (Gabon and Equatorial Guinea) at 

depths of 36 to 63 m (see Stock, 1955; 

Müller, 1990; Bamber, 2004). Both of these 

species also share the characters of a sixarticled 

ovigerous leg, the ocular tubercle 

overhanging the proboscis, small auxiliary 

claws and a propodal lamina. A. 

amoybios, the only other Atlantic species, 

is immediately distinct in having dorsodistal 

tubercles on coxa 1 of all legs. A. 

erectus normally has conspicuous dorsodistal 

tubercles on the lateral processes, 

unlike the present species. In addition, A. 

erectus is conspicuously more slender 

than A. amora sp. nov, femur and tibia 2 

length:width ratios being 5.3 and 7.7 

respectively (3.9 and 5 respectively in A. 

amora), and the width across trunk segment 

2 is four times the length of the segment 

(

174 AÇOREANA 

2009, Sup. 6: 167-182 

(Krøyer, 1844) from the Azores, at depths 

between 1 and 30 m; that species is readily 

distinguished from A. amora by its 

cement gland configuration (without subcutaneous 

extension, see Figure 2G). 

Anoplodactylus angulatus (Dohrn, 1881) 

Figure 3A to C 

Material: 1 female, WVF011, northeastern 

side of Ilhéu de Vila Franca do Campo, 16 

m, scuba dive collection by Gonçalo 

Calado, José Pedro Borges, Joana Xavier, 

Paola Rachello, Patrícia Madeira, 20 July 

2006.1 male, 3 females, 1 subadult, Isl. 

24.4, attached littoral algae, south wall 

within the flooded crater of the Ilhéu de 

Vila Franca, 24 July 2006, coll. A. 

Salvador, R. Robbins & R.B.; 2 males with 

eggs, 1 male, 4 females, 2 subadults, Isl. 

24.5, mid-lagoon algae, within the flooded 

crater of the Ilhéu de Vila Franca, 24 

July 2006, coll. A. Salvador, R. Robbins & 

R.B. 

1 specimen, Mosteiros, 8 November 


1997; 1 specimen, Santa Clara, 7 

November 1996 ; 3 specimens, Santa 



specimens, Emissário, 2 November 1996; 

24 specimens, Emissário, 12 June 1997; 4 

specimens, Sinaga, 30 May 1997; 25 


specimen, Ribeira da Praia, 25 October 


June 1997; 7 specimens, Caloura, 26 June 

1997; 6 specimens, Porto de Vila Franca 

do Campo, 9 May 1997; 2 specimens, 

Ribeira Quente, 18 June 1997; 10 

specimens, Nordeste, 2 May 1997; 1 

specimen, Porto Formoso, 30 October 

1996; 3 specimens, Porto Formoso, 19 June 

1997; 4 specimens, Ladeira da Velha, 19 

June 1997; 10 specimens, Ribeirinha, 25 

June 1997; 1 specimen, Cofaco, 25 June 

1997;2 specimens, São Vicente, 6 May 

1997; all coll. João Brum & A.C.C. 

Remarks: the commonest species of 

Anoplodactylus in the Azores, A. angulatus 

is also known from the Mediterranean, 

the Atlantic coasts of Northern Europe 

and North Africa (Morocco), as well as 

the Canary Islands. It is an algal-associated 

species without a conspicuous overhang 

of the anterior of the cephalon, and 

distinguished from sympatric specimens 

of A. virescens (also occurring in algae) by 

the marked angular distal corners of the 

proboscis (Figure 3B). 

Anoplodactylus maritimus Hodgson, 1915 

Figure 3D 

Incl. A. parvus Giltay, 1934; non-A. parvum 

(Hilton, 1942) = A. hokkaidoensis (Utinomi, 

1954). 

Material: 1 specimen, Emissário, 12 June 

1997; 2 specimens, Caloura, 26 June 1997; 

15 specimens, Porto de Vila Franca do 

Campo, 9 May 1997; 1 specimen, Ilhéu de 

Vila Franca do Campo, 24 October 1996; 2 


Campo, 16 June 1997; 10 specimens, 

Lactoaçoreana, 25 June 1997; all coll. João 

Brum & A.C.C. 

Remarks: Anoplodactylus maritimus has 

been recorded previously throughout 

Macaronesia, other than which it is a 

species of the Atlantic coast of the 

Americas (see discussion below) but has 

not been recorded from the Atlantic 

coasts of mainland Europe, North Africa 

or the Mediterranean. 

Anoplodactylus pygmaeus (Hodge, 1864) 

Figure 3E, F 

Material: 1 female, Isl. 24.4, attached lowlittoral 

algae, south wall within the flooded 

crater of the Ilhéu de Vila Franca;1 

female, Isl. 24.5, mid-lagoon algae, within 

the flooded crater of the Ilhéu de Vila 

Franca; both 24 July 2006, coll. A. 

Salvador, R. Robbins & R.B.


FIGURE 3. A to C, Anoplodactylus angulatus: A, entire, dorsal; B, proboscis, ventral; C, distal leg articles 

(lam = lamina); D, A. maritimus, distal leg articles; E and F, A. pygmaeus: E, body, lateral; F, third 

leg, entire; G to H, A. virescens: G, proboscis, ventral; H, distal leg articles.

176 AÇOREANA 

2009, Sup. 6: 167-182 

1 specimen, Pesqueiro, 7 November 

1997; 4 specimens, Porto de Vila Franca 

do Campo, 9 May 1997; 1 specimen, Faial 

da Terra, 30 May 1997; 1 specimen, 

Ribeirinha, 25 June 1997; all coll. João 

Brum & A.C.C. 

Remarks: Anoplodactylus pygmaeus is a 

small species (as its name suggests), similar 

to A. petiolatus but with a less-marked 

anterior cephalic overhang and with distinct 

distal spines on the dorsodistal 

tubercles of the lateral processes; auxiliary 

claws are absent. It has been recorded 

throughout the North Atlantic and 

Mediterranean, including the Canaries 

and the Azores, in shallow water (the 

record from 3850 m depth on the Cape 

Verde Slope by Bamber & Thurston, 1993, 

is presumed to be a result of samplinggear 

contamination from the fouling community 

on the ship’s hull). 

Anoplodactylus virescens (Hodge, 1864) 

Figure 3G, H 

Material: 1 female, WVF040, off Amora, 

Ponta Garça, São Miguel, Azores, N37° 

42’ 720” W25° 21’ 554”, 37.8 m depth, 

small dredge sample, 26 July 2006, coll. 

António de Frias Martins & Jerry 

Harasewych. 

Remarks: while isolated females of this 

genus can be difficult to identify to 

species, Anoplodactylus virescens, with its 

trunk segmentation only between somites 

1, 2 and 3, its presence of small auxiliary 

claws, its lack of propodal lamina, of dorsodistal 

lateral-process tubercles, of ventral 

proboscis tubercles and of chela teeth, 

is only confusable with A. robustus 

(Dohrn, 1881), but the latter species has 

angulate corners to its proboscis, unlike 

the present species. A. virescens is known 

from the north-east Atlantic from the 

United Kingdom to Morocco and 

throughout the Mediterranean. This is the 

first record of this species for the Azores. 

Family Callipallenidae Hilton, 1942 

Callipallene emaciata (Dohrn, 1881) 

Figure 4A 

Material: 1 male, 3 females, 1 juvenile, Isl. 

24.4, attached low-littoral algae, south 

wall within the flooded crater of the Ilhéu 

de Vila Franca; 5 females, 1 male, Isl. 24.5, 

mid-lagoon algae, within the flooded 

crater of the Ilhéu de Vila Franca; both 24 


R.B. 

4 specimens, Mosteiros, 8 November 


1997; 12 specimens, Ponta da Ferraria, 17 

June 1997; 24 specimens, Santa Clara, 7 




specimens, Emissario, 2 November 1996; 

179 specimens, Emissario, 12 June 1997; 

26 specimens, Sinaga, 30 May 1997; 1 

specimen, Atalhada, 25 October 1996; 222 


specimens, Ribeira da Praia, 25 October 


June 1997; 18 specimens, Caloura, 14 





Campo, 16 June 1997; 13 specimens, 

Ribeira Quente, 18 June 1997; 4 

specimens, Faial da Terra, 17 October 

1996; 21 specimens, Faial da Terra, 30 May 

1997; 4 specimens, Nordeste, 21 

November 1966; 51 specimens, Nordeste, 

2 May 1997; 25 specimens, Porto Formoso, 

30 October 1996; 69 specimens, Porto 

Formoso, 19 June 1997; 5 specimens, 

Ladeira da Velha, 19 June 1997; 29 

specimens, Ribeirinha, 8 October 1996; 67 

specimens, Ribeirinha, 25 June 1997; 14 

specimens, Lactoaçoreana, 14 October 

1996; 26 specimens, Lactoaçoreana, 25 

June 1997; 4 specimens, Cofaco, 25 June


1997; 17 specimens, São Vicente, 7 

October 1996; 1 specimen, São Vicente, 6 

May 1997; all coll. João Brum & A.C.C. 

Remarks: the European shallow-water 

species of the genus Callipallene are small, 

cryptic species normally associated with 

epizoan turfs of bryozoans and hydroids. 

The distinction of these taxa was analysed 

by Stock (1952), since when his subspecies 

have been raised to full specific rank (C. 

brevirostris (Johnston, 1837), C. emaciata, C. 

FIGURE 4. A to B, Callipallene emaciata: A, entire, dorsal; B, distal; leg articles; C, Endeis spinosa, 

entire, dorsal (after Bamber, 1983).

178 AÇOREANA 

2009, Sup. 6: 167-182 

tiberi (Dohrn, 1881), C. phantoma (Dohrn, 

1881)). While C. brevirostris is the commonest 

species on the Atlantic coasts of 

North Europe, the present species is the 

only Callipallene recorded from the shallow 

waters of the Azores (the deeperwater 

C. producta Sars, 1888 has been 

recorded off the Azores (Bouvier, 1917; 

Arnaud, 1974) at depths >800 m), and is 

more commonly associated with algae 

than are the other species listed above. 

All the Azorean specimens were collected 

from algae, the species being the second 

commonest recorded (occurring, for 

example, at every sampling site in 1996 

and 1997). 

Family Endeidae Norman, 1908 

Endeis spinosa (Montagu, 1808) 

Figure 4B 

Material: 3 specimens, Mosteiros, 8 


Clara, 12 June 1997; 1 specimen, Sinaga, 

30 May 1997; 5 specimens, Atalhada, 30 

May 1997; 9 specimens, Ribeira da Praia, 

16 June 1997; 1 specimen, Caloura, 14 


June 1997; 1 specimen, Porto de Vila 



Campo, 16 June 1997; 1 specimen, Ribeira 

Quente, 18 June 1997; 2 specimens, Faial 

da Terra, 30 May 1997; 1 specimen, 

Nordeste, 2 May 1997; 2 specimens, Porto 

Formoso, 19 June 1997; 1 specimen, 

Lactoaçoreana, 14 October 1996; 1 

specimen, Lactoaçoreana, 25 June 1997; 1 

specimen, São Vicente, 6 May 1997; all 

coll. João Brum & A.C.C. 

Remarks: a widespread North Atlantic 

species, where its only native congener is 

Endeis charybdaea (Dohrn, 1881) on 

European coasts. Stock’s (1990) record of 

E. meridionalis (Böhm, 1879) from Cape 

Verde is highly suspect. E. mollis 

(Carpenter, 1904) may be a Lessepsian 

immigrant to the Mediterranean. While 

the larval biology of this species is 

unknown, there is evidence of its waterborne 

transport, perhaps by association 

with hydromedusae. 

DISCUSSION 

There are now fifteen species of shallow-water 

pycnogonids recorded from 

the Azores: Ammothoella longipes, 

Anoplodactylus virescens and A. amora sp. 

nov. are newly recorded for the islands 

from the present study. Achelia echinata, 

Anoplodactylus angulatus and Callipallene 

emaciata are common. 

Analysis of the provenance of these 

species is somewhat challenged by the 

presence of species which have in the past 

been misidentified, early records worldwide 

depending on what was at the time 

a limited literature, and a limited understanding 

of specific characters. 

For example, Achelia echinata, a species 

with a type locality in the United 

Kingdom, has been recorded in the literature 

not only throughout the north-east 

Atlantic and Mediterranean (an accepted 

distribution), but also from China and 

Japan (as subspecies), from California 

and Alaska. Recent molecular work 

(Bamber et al. in prep.) has shown not 

only that the material from the Azores is 

fully conspecific with material from the 

United Kingdom, but also that the 

Chinese-Japanese “subspecies” is in fact a 

distinct species. The North- and East- 

Pacific material is highly suspect, and is 

not accepted herein. 

Rhynchothorax anophtalmus Arnaud, 

1972, recorded as such from the Azores by 

Arnaud (1974), was synonymized with R. 

philopsammum Hedgpeth, 1951 by Arnaud 

& Krapp (1990), when comparing specimens 

from the Mediterranean and from 

California: R. philopsammum is a 

Californian species, also recorded from


the Subantarctic, Pacific Mexico, 

Magellanic Chile, a number of Pacific 

Islands and Belize. Other than the Pacific 

material , the species is recorded (as R. 

anophtalmus) from Azores and the 

Mediterranean. Rhynchothorax is an interstitial 

genus which would not be expected 

to be transported by ships’-hull fouling or 

by floating weed. With this disparity in 

zoogeography, closer examination may 

yet show these two taxa to be distinct. 

Records of R. monnioti (type-locality the 

Azores) from Brazil and Trinidad 

(Arnaud & Krapp, 1990; Child, 1988) are 

dubious, and not accepted herein. 

In the light of the above, the species 

currently known from the Azores show a 

number of zoogeographies: 

- two currently endemic species – Achelia 

anomala (not recorded since the original 

discovery and description: 

Arnaud, 1974), and Anoplodactylus 

amora (recorded herein for the first 

time); Morton & Britton (2000) point 

out that endemism is low in these 

islands owing to their “youthfulness” 

(earliest emergence about 8 My ago), 

and such species are likely to be found 

elsewhere in the future; 

- two species from Macaronesia and the 

Mediterranean - Rhynchothorax anophtalmus 

from the Mediterranean; R. 

monnioti from the Mediterranean and 

the Canary Islands; 

- three species from the North-east 

Atlantic, the Mediterranean and 

North Africa (Morocco) – Ammothoella 

longipes, Anoplodactylus angulatus, 

Anoplodactylus virescens (the first two 

also recorded elsewhere in the 

Macaronesian system); 

- five species found in the Mediterranean 

and throughout the 

North Atlantic, including the 

Americas – Achelia echinata, 

Anoplodactylus petiolatus, Anoplodactylus 

pygmaeus, Callipallene emaciata and 

Endeis spinosa: closer study may yet 

distinguish cryptic species within 

these data; 

- a species found throughout the southern 

North Atlantic – Tanystylum orbiculare 

(distribution as the previous group 4, 

but not including the European 

Atlantic coasts); 

- a species from Macaronesia and the 

West Atlantic only – Anoplodactylus 

maritimus, recorded from Madeira, 

Cape Verde Islands, the Canaries and 

the Americas from Chesapeake Bay 

through the Gulf of Mexico and the 

Caribbean to Brazil; 

- an immigrant species – Endeis straughani 

(if a valid record), a species native 

to Australia but since recorded in 

ship’s-hull-fouling from Ghana 

(Bamber, 1979, as E. picta; Staples, 

1982). 

Apart from the presently endemic 

species and the anthropogenic immigrant, 

all but one of these species are also 

found in the Mediterranean, and of these 

all but the two Rhynchothorax species are 

recorded from Morocco. Five of these 

twelve species have not been found in the 

Western Atlantic. These data give a 

strong indication that the pycnogonid 

fauna of the Azores is largely of an easterly 

origin, from the Mediterranean, North 

Africa or Atlantic Europe. 

The notable exception is 

Anoplodactylus maritimus, recorded from 

the Azores, the Canary Islands, Cape 

Verde and Madeira (see Fage & Stock, 

1966; Stock, 1990; Child, 1992), and found 

in nine of the samples from 1996 and 

1997. This species is also found in the 

Americas (see above), including in the 

Sargasso Sea, and is commonly recorded 

amongst floating Sargassum (Bourdouillon, 

1955; Stock, 1954; Stock, 

1957; McCloskey, 1973; Stock,1992; Child, 

1992); indeed, Hodgson (1915) found it in 

floating Sargassum south of the Azores. In

180 AÇOREANA 

2009, Sup. 6: 167-182 

addition, Stock (1994) recorded “>500 

specimens” of A. maritimus associated 

with surface-floating hydroids in the 

mid-Atlantic, at 24.75°N 44 to 53°W. 

Drift dispersal in algae as a viable 

means of passive migration by pycnogonids 

was analyzed by Bamber (1998). 

Perhaps significantly, other species 

recorded amongst Sargassum are Endeis 

spinosa, Anoplodactylus petiolatus and 

Tanystylum orbiculare (see Timmermann, 

1932; Hedgpeth, 1948), and the first two 

of these have been recorded on Sargassum 

in the vicinity of the Azores (Hedgpeth, 

1948: Figure 7). 

Finally, it is notable that the most 

diverse genus recorded, Anoplodactylus, 

includes species which are known to live 

upon medusae, and thus obtain passive 

dispersion in the plankton (see 

Introduction, above); the comparatively 

widespread distribution of Anoplodactylus 

species is commonly attributed to this 

process (see Bamber, 1998 for discussion). 

Thus, the proximity and known distributions 

of most species would favour colonization 

by species from the east 

(Mediterranean, North Africa, Atlantic 

Southern Europe), and as far as we know 

this can be the only source of five of the 

species discussed above; Morton & 

Britton (2000) found that most components 

of the Azorean marine fauna show 

affiliation with the Mediterranean and 

southern Europe. However, the incidence 

of species recorded in the Americas in 

floating algae gives a mechanism for 

transport, which would argue for colonization 

from the west. In reality, pycnogonid 

colonization of the shallow water 

habitats of the Azores may be attributed 

to both processes. However, the mechanism 

of immigration from the east 

remains purely speculative. 

For the species occurring in the 1996 

and 1997 data in sufficient numbers for 

interpretation, no trends were detected in 

relation to disturbance or exposure, 

although Anoplodactylus amora only 

occurred at non-polluted sites. 


We are indebted to Roni Robbins for 

assistance in the field collecting and sample 

analysis in 2006, to Andreia Salvador 

for sampling assistance in Ilhéu de Vila 

Franca, to João Brum for assistance in the 

diver-collections in 1996 and 1997, to 

Brian Morton for discussions on the origins 

of the Azorean fauna, particularly to 

António de Frias Martins for the organisation 

of, and inviting us to the 

Workshop, and to the other participants 

at the workshop for samples and for 

entertainment. 


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THE TANAIDACEANS (ARTHROPODA: PERACARIDA: TANAIDACEA) OF SÃO 

MIGUEL, AZORES, WITH DESCRIPTION OF TWO NEW SPECIES, AND A NEW 

RECORD FROM TENERIFE 

Roger N. Bamber 1 & Ana Cristina Costa 2 

1 

The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. e-mail: r.bamber@artoo.co.uk 

2 


Azores, Portugal 

ABSTRACT 

During the Third International Workshop of Malacology and Marine Biology in São 

Miguel, Azores, in July 2006, sampling of the littoral and sublittoral benthos was undertaken 

in order to characterize the smaller marine arthropod fauna of this region, including 

tanaidaceans. In the event, 338 tanaidacean specimens representing three species were 

collected; two of these species, Leptochelia caldera and Paratanais martinsi, are new to science. 

In addition, previous tanaidacean material collected around São Miguel in 1996 and 

1997 was analyzed, from which a further 272 specimens (three species, one not found in 

2006) were identified. No apseudomorph tanaidaceans were found. All of the material is 

described, and an attempt is made to investigate the provenance of the Azorean 

tanaidacean fauna, although the general lack of data from the North-east Atlantic precludes 

any reasonable interpretation other than some possible links with the 

Mediterranean. However, recent material of Zeuxo exsargasso collected from Tenerife, in 

the Canary Islands, does suggest Macaronesian links with the West Atlantic. 

RESUMO 

Durante o 3º Workshop Internacional de Malacologia e Biologia Marinha em São 

Miguel, Açores, em Julho de 2006, fizeram-se amostragens do bentos litoral e sublitoral de 

modo a caracterizar a fauna de pequenos artrópodes marinhos dessa região, incluindo os 

tanaidáceos. Assim, recolheream-se 338 espécimens de tanaidáceos representando três 

espécies; duas dessas espécies, Leptochelia caldera e Paratanais martinsi, são novas para a 

ciência. Para além disso, analisou-se material recolhido previamente à volta de São Miguel 

em 1996 e 1997, do qual foram identificados 272 especimens ( três espécies, uma das quais 

não encontrada em 2006). Não se encontraram tanaidáceos apseudomorfos. Todo o 

material foi descrito, e abordou-se a questão da proveniência da fauna tanaidácea 

Açoreana, embora a ausência generalizada de dados do Nordeste Atlântico impeça 

qualquer interpretação razoável para além de algumas possíveis ligações com o 

Mediterrâneo. No entanto, material recente de Zeuxo exsargasso recolhido em Tenerife, 

Ilhas Canárias, sugere ligações Macaronésicas com o Atlântico Oeste. 

INTRODUCTION 









Drift, a diffuse southerly arm of the Gulf 

Stream breaking off from the North 


Americas, while the somewhat less-significant 

western eddies of the Canary 



current brings warm, hyperhaline water

184 AÇOREANA 

2009, Sup. 6: 183-200 

from the Mediterranean outflow (Gofas, 

1990; Morton et al., 1998). This hydrography 

clearly has implications for the colonization 

of the islands by benthic marine 

species. 

Tanaidaceans are a group of the 

arthropods with minimal dispersive ability; 

the larvae are not planktonic, and 

there are only limited examples of adults 

swimming (Bamber, 1998). Some species 

are known to have spread in fouling communities 

on ship’s hulls (Bamber, 1977), 

and others are known to live in floating 

algae (Sieg, 1980) or ectoparasitically on 

turtle tests and on manatees (see Morales- 

Vela et al., 2008). It is therefore of some 

interest to determine the suite of species 

which has colonized the Azores archipelago, 

and their provenance. 

Previous records of tanaidaceans from 

the Azores are very sparse, and largely 

incidental, although Dollfus (1897) 

reported on the tanaidacean material collected 

around the Azores during the 

cruise of the Hirondelle in 1887 and 1888. 

All but two of the species recorded in litt. 

are from deep-water (>100 m; mostly 

>500 m). Dollfus (1897) reported 

Leptochelia savignyi Krøyer, 1842 from 

Horta; despite the controversy regarding 

earlier records of sibling species of this 

taxon, Dollfus’ records appear to be substantiated 

(see below); L. savignyi was 

described originally from Madeira, so its 

occurrence in the Azores is not surprising. 

Dollfus (1897) also recorded Tanais 

dulongii (Audouin, 1826) (as Tanais 

cavolinii Milne-Edwards, 1828) from Baïe 

de Fayal, and described as new 

T. grimaldii from Horta, distinguishing the 

two on the shape of the cephalon (his new 

species having a shorter cephalon) and 

the number of uropod articles; there is 

doubt whether his T. dulongii specimens 

were in fact of that species (see below); 

Morton et al. (1998) mention the occurrence 

of Tanais in littoral algal mats. 

Finally, the record of “Paratanais atlanticus” 

of Dollfus (1897) is incertae sedis, but 

not attributable to Paratanais (see below). 

The nine confirmed species recorded 

from around the Azores are: 

Suborder Apseudomorpha 

Leviapseudes leptodactylus (Beddard, 

1886), Azores 1830 m. 

Suborder Tanaidomorpha 

Superfamily Tanaoidea 

Tanais grimaldii Dollfus, 1897, Azores, 

littoral, 5-6 m. 

Superfamily Paratanaoidea 

Siphonolabrum mirabile Lang, 1872, 

Azores 3500-4165 m. 

Agathotanais hanseni Lang, 1971, E. 

Pacific & Azores, 2861-4165 m 

Leptognathia abyssorum (Dollfus, 1897), 

Azores, 1287 m. 

Paratyphlotanais richardi (Dollfus, 1897), 

W Ireland, Azores, 699-1287 m. 

Typhlotanais spiniventris Dollfus, 1897, 

Azores, 130-1287 m. 

Mesotanais dubius Dollfus, 1897, 

Azores, 1287 m. 

Leptochelia savignyi Krøyer, 1842, 

Azores, 5 to 6 m. 



Biology at Vila Franca do Campo, São 

Miguel, in July 2006, sampling of the littoral 

and sublittoral benthos was undertaken 

in order to characterize the smaller 

marine arthropod fauna of this region, 

including tanaidaceans. In the event, 338 

specimens representing three species 

were collected; two of these species are 

new to science. In addition, a previous 

collection from around the island was 

made available, from which a further 272 

specimens (three species, one additional 

to the above three) were identified. No 

apseudomorph tanaidaceans were found.

BAMBER & COSTA: TANAIDACEANS FROM SÃO MIGUEL 185 

In addition, recently collected material 

from Tenerife, in the Canary Islands, 

kindly supplied to us by Brian Morton, 

revealed a new record of a tanaidacean, 

relevant to the origins of the 

Macaronesian fauna. All of the material is 

described below. 


The present Azores material comes 

from two sources. During the Workshop 

at Vila Franca do Campo in July 2006, a 

number of littoral and infralittoral habitats 

on the island of São Miguel were sampled 

for tanaidaceans, including crevice 

habitats, macroalgae and soft sediments. 

The principal sampling areas were the littoral 

sediments, rocks and algae below 

the Clube Naval building (the old Vila 

Franca do Campo abattoir); the sediments 

and algae within the caldera of the Ilhéu 

de Vila Franca; and the soft sediments off 

Vila Franca do Campo (ca N37º 43’ W25º 

25’), from 12 to 200 m depth. Offshore 

sediments were sampled using a 0.025 m 2 

van Veen grab and various dredges. All 

samples were washed through a 0.5 mm 

mesh sieve, and specimens sorted alive. 

Some of these specimens were fixed in 

absolute ethanol to allow DNA analysis. 

Extensive material collected in 1996 

and 1997, from 11 (1996) and 20 (1997) 

infralittoral rocky-substratum sites 

around São Miguel, was also analysed in 

detail. Samples of algae (Stypocaulon scoparia, 

Halopteris filicina and Zonaria tournefortii) 

were collected by SCUBA diving at 

depths from between 5 and 16 m. Details 

of the sampling and protocols are given 

by Costa & Ávila (2001). The sampling 

sites, anti-clockwise around the Island 

from the north-west, were Mosteiros, 

Ponta da Ferraria, Santa Clara, Pesqueiro, 

Emissário, Sinaga, Atalhada, Ribeira da 

Praia, Caloura, Porto de Vila Franca do 

Campo, Ilhéu de Vila Franca, Ribeira 

Quente, Faial da Terra, Nordeste, Porto 

Formoso, Ladeira de Velha, Ribeirinha, 

Lactoaçoreana, Cofaco and São Vicente. 

These sites variously represented exposed 

and sheltered shores, undisturbed, naturally 

disturbed (near shallow-water vent 

sites or stream mouths) and polluted 

shores. 

Finally, a collection of littoral algal 

turf was made from a rocky intertidal 

platform at Playa de Fanabe, Costa Adeje, 

Tenerife, Canary Islands, in June 2007, 

from which a further species not found in 

the São Miguel material was extracted. 

Terminology used herein recognizes 

the first pair of antennae as antennules, 

the second pair as the antennae. The first 

maxilla is termed the maxillule. The first 

pair of pereopods (of six) is the pair 

immediately posterior to the chelipeds. 

Serially repetitive body-parts, such as the 

pereonites and subdivisions of antennal 

flagella are segments, others (such as the 

parts of the limb) are articles. Total length 

is measured axially from the tip of the 

rostrum to the posterior edge of the pleotelson; 

measurements were made dorsally 

on the body and antennules, and laterally 

on the pereopods and antennae. The 

term ‘spines’ is used in the traditional 

sense to distinguish between rigid ‘thornlike’ 

structures and the more flexible 

‘hair-like’ setae (in keeping with their etymology 

and all historic literature); nonarticulating 

spine-shaped extensions of 

the cuticle are considered to be apophyses; 

comb–rows of fine setules, occasionally 

present on maxillae and pereopod 

articles, inter alia, are referred to as 

microtrichia. 

Voucher and type-material has been 

lodged in the collections of The Natural 

History Museum, London (NHM). The 

higher taxonomy is based on Guţu & Sieg 

(1999).

186 AÇOREANA 

2009, Sup. 6: 183-200 

SYSTEMATICS 

SÃO MIGUEL MATERIAL 

Suborder TANAIDOMORPHA Sieg, 1980 

Superfamily Tanaoidea Dana, 1849 

Family Tanaidae Dana, 1849 

Subfamily Tanainae Dana, 1849 

Genus Tanais Latreille, 1831 

Tanais grimaldii Dollfus, 1897 

Figure 1A, B 

Sieg, 1980, pp. 84-91; Figure 31, 22. 

Material: 1 female with oostegites, 1 

female with empty brood pouch, 2 

subadult females, 2 neuters, 1 manca 

(Registration N os NHM.2007.764-771), Isl. 

24.2, drift algae within the flooded crater 

of the Ilhéu de Vila Franca, 24 July 2006, 

coll. A. Salvador, R. Robbins & R.B.; 12 

females with oostegites, 4 brooding 

females, 6 males, 148 neuters, 24 mancae 

(NHM.2007.772-781), Isl. 24.4, attached 

low-littoral algae, south wall of the flooded 



R.B.; 4 females with oostegites, 1 brooding 

female, 4 males, 31 subadult females, 

42 neuters, 28 mancae, Isl. 24.5, midlagoon 

algae, within the flooded crater of 

the Ilhéu de Vila Franca, 24 July 2006, coll. 

A. Salvador, R. Robbins & R.B.; 2 neuters, 

2 mancae (NHM.2007.782-785), WVF011, 

northeastern side of Ilhéu de Vila Franca 

do Campo, 16 m, scuba dive collection by 

Gonçalo Calado, José Pedro Borges, Joana 

Xavier, Paola Rachello, Patrícia Madeira, 

20 July 2006. 

1 brooding female, 1 neuter, 1 manca 

II, Mosteiros, 17 June 1997; 1 neuter, 1 

manca II, Ponta da Ferraria, 17 June 1997; 

3 females (1 brooding), 2 males, 1 manca, 

Emissário, 2 November 1996; 1 manca, 

Atalhada, 30 May 1997; 1 female, 4 

neuters, 15 mancae, Caloura, 26 June 

1997; 5 neuters, 1 manca, Ribeira Quente, 

November 1996; 1 manca, Ribiera Quente, 

18 June 1997; 1 manca, Nordeste, 21 

November 1996; 1 male, 1 brooding 

female, 1 neuter, 2 mancae. Nordeste, 2 

May 1997; 1 female, Porto Formoso, 30 

October 1996; 4 males, 1 female with oostegites, 

3 neuters, 2 mancae, Porto 

Formoso, 19 June 1997; 2 males, 3 mancae, 

Ladeira da Velha, 19 June 1997; 4 females 

(1 brooding, 2 with oostegites), 1 male, 

4neuters, 1 manca, Ribeirinha, 8 October 

1996; 2 females with oostegites, 7 neuters, 

5 mancae, Ribeirinha, 25 June 1997; 4 

females (1 brooding), 8 neuters, 9 mancae, 

Lactoaçoreana, 25 June 1997; 1 male, 1 

female, Cofaco, 7 October 1996; 4 females 

(1brooding), 7 males, 6 neuters, 1 manca, 

São Vicente, 7 October 1996; 3 females, 3 

neuters, 6 mancae, São Vicente, 6 May 

1997. All coll. A.C.C. & João Brum. 

Remarks: Tanais grimaldii is one of the only 

two littoral tanaidacean species recorded 

previously from the Azores: the typelocality 

is Horta (Faial), at 5 to 6 m depth 

(Dollfus, 1897). T. grimaldii is distinguished 

from its only northeast Atlantic 

congener, T. dulongii (Audouin, 1826) by 

the conformation of the laciniae mobili of 

the mandibles (see Sieg, 1980, Figure 31), 

but also adults of the present species have 

one more article in the uropod (basis plus 

three-segmented endopod: Figure 1B) 

than does T. dulongii (basis plus two-segmented 

endopod). Dollfus (1897) also 

recorded Tanais dulongii (Audouin, 1826) 

(as Tanais cavolinii Milne-Edwards, 1828) 

from Baie de Fayal, distinguishing it and 

T. grimaldii on the shape of the cephalon 

(his new species having a shorter 

cephalon) and the number of uropod articles; 

it is unclear whether his specimens 

of putative T. dulongii were fully mature, 

as only adult T. grimaldii have four segments 

to the uropod endopod (T. dulongii 

can be distinguished from immature 

T. grimaldii with three-segmented


FIGURE 1. A, B, Tanais grimaldii Dollfus, 1897: A, dorsal; B, uropod; C to E, Leptognathia breviremis 

(Lilljeborg, 1864): C, dorsal; D, uropod; E, cheliped. (A redrawn after Sars, 1886; B, Azores specimen; 

C to E modified from Sars, 1899). 

endopods, as the penultimate segment is 

twice as long as the ultimate segment in 

the former, only slightly longer in the latter). 

Dollfus did not examine the 

mandibular structure, and indeed would 

have been unaware of its significance. 

It is likely that all previous records of 

Tanais dulongii from the Azores in fact 

refer to T. grimaldii. The present species is 

also recorded from the Italian 

Mediterranean (Gulf of Naples, Ischia 

and Linosa); there are unconfirmed 

records from the western Mediterranean 

and Casablanca (Sieg, 1980). 

All the material reported above was 

collected from algae at less than 17 m 

depth. Brooding females and mancae 

were present in all the months sampled 

(May, June, July, October and November). 

Superfamily Paratanaoidea Lang, 1949 

Family Anarthruridae Lang, 1971 

Subfamily Leptognathiinae Sieg, 1976 

Genus Leptognathia Sars, 1882

188 AÇOREANA 

2009, Sup. 6: 183-200 

cf. Leptognathia breviremis (Lilljeborg, 

1864) 

Figure 1C to E 

Material: 1 headless specimen, Ponta da 

Ferraria, 17 June 1997, coll. A.C.C. & João 

Brum. 

This damaged specimen is almost 

unidentifiable; the cheliped and uropods are 

appropriate to L. breviremis. This species is 

known from the eastern North Atlantic, but 

has been recorded (dubiously) from the 

North Pacific. A figure of L. breviremis (modified 

from Sars, 1899) is given to aid possible 

recognition of this taxon in the Azores in 

future (Figure 1, C to E). 

Family Leptocheliidae Lang, 1973 

Genus Leptochelia Dana, 1849 

Leptochelia caldera sp. nov. 

Figures 2, 3 

Material: 1 female with oostegites, holotype 

(NHM.2007.424), 1 male, allotype 

(NHM.2007.425), 13 females, paratypes 

(NHM.2007.426-435), Isl. 24.4, attached 

low-littoral algae, south wall of the flooded 



R.B.; 2 females, paratypes 

(NHM.2007.436-437), Isl. 24.5, midlagoon 

algae, within the flooded crater of 

the Ilhéu de Vila Franca, 24 July 2006, coll. 

A. Salvador, R. Robbins & R.B.. 

2 females, Pesqueiro, 7 November 

1997; 3 males, 32 females (3 brooding), 5 

neuters, Emissário, 2 November 1996; 3 

females, 1 male, Emissário, 12 June 1997; 1 

manca, Sinaga, 30 May 1997; 4 males, 40 

females (2 brooding), 9 neuters, 2 mancae, 

Atalhada, 25 October 1996; 1 male, 14 

females, 7 neuters, Atalhada, 30 May 

1997; 1 female, Caloura, 26 June 1997; 1 

neuter, Faial da Terra, 12 June 1997; 1 

female, 1 neuter, Nordeste, 12 May 1997; 1 

male, 1 neuter, Ribeirinha, 8 October 1996; 

all coll. A.C.C. & João Brum. 

Description of female: body (Figure 2A) 

slender, holotype 3.6 mm long, 7.3 times 

as long as wide. Cephalothorax subrectangular, 

1.6 times as long as wide, as 

long as pereonites 2 and 3 together, with 

slight rostrum, eyelobes rounded, eyes 

present and black, single setae at posterior 

of eyelobes and posterolaterally. Six 

free pereonites; pereonite 1 shortest, pereonites 

2 and 3 subequal, 1.3 times as long 

as pereonite 1; pereonites 4 and 5 subequal 

(pereonite 4 longest) and 1.25 times 

as long as pereonite 2; pereonite 6 just 

longer than pereonite 1 (all pereonites 

respectively 1.9, 1.4, 1.3, 1.1, 1.0 and 1.4 

times as wide as long). Pleon of five free 

subequal pleonites bearing pleopods; 

each pleonite about 3.5 times as wide as 

long, with paired lateral setae. Pleotelson 

(Figure 2M) semicircular, 0.16 times as 

long as pleon, 2.8 times as wide as long, 

with paired lateral setae, paired posterolateral 

setae on each side and two distal 

setae. 

Antennule (Figure 2D) of four tapering 

articles, proximal article 3.6 times as 

long as wide, 1.5 times as long as distal 

three articles together, with two long 

outer and single short dorsal and inner 

setae; second article twice as long as wide, 

distal outer seta shorter than article; third 

article 1.3 times as long as second and 

with one aesthetasc; fourth article minute, 

eccentric, with five distal setae. 

Antenna (Figure 2G) of six articles, 

proximal article compact, naked; second 

article as long as wide, with single ventrodistal 

and dorsodistal slender spines; 

third article 1.3 times as long as wide, 

with dorsodistal slender spine; fourth 

article longest, three times as long as 

wide; fifth article half as long as fourth; 

sixth article minute. 

Labrum (Figure 2H) rounded, setose, 

typical of genus. Left mandible (Figure 

2I) with crenulate lacinia mobilis wider 

than pars incisiva, proximal crenulation


FIGURE 2. Leptochelia caldera sp. nov., A, holotype female, dorsal; B, allotype male, dorsal; C, allotype 

male, lateral; D, female antennule; E, male antennule; F, male antenna; G, female antenna; H, 

labrum, lateral; I, left mandible; J, maxillule and maxilla; K, maxilliped; L, epignath; M, pleotelson 

and left uropod. Scale line = 1 mm for A, B and C, 0.2 mm for D to G and M, 0.1 mm for H to L.

190 AÇOREANA 

2009, Sup. 6: 183-200 

on pars incisiva, pars molaris with strong 

rugosity; right mandible similar but without 

lacinia mobilis. Labium typical of 

genus, distally finely setose, without 

palp. Maxillule (Figure 2J) with seven 

long and two short distal spines and 

setose margins; palp of two articles with 

two distal setae; maxilla simple, ovoid. 

Maxilliped (Figure 2K) palp first article 

naked, second article with one outer and 

three inner setae, and distal seta exceeding 

distal margin of third palp article; 

third and fourth articles with filtering 

rows of six and seven setae respectively, 

third article with two further inner distal 

setae, fourth article with submarginal 

outer seta; basis with three long setae 

extending to third palp article; endites 

distally with single outer seta and one 

inner rounded and two robust spatulate 

spines. Epignath (Figure 2L) elongate, 

arcuate, with setose margin distally and 

proximally. 

Cheliped (Figure 3A) with rounded, 

compact basis 1.9 times as long as wide, 

with inner distal seta; merus subtriangular 

with three ventral setae; carpus 1.9 

times as long as wide, with three midventral 

setae; propodus typical for the genus, 

fixed finger with three ventral and three 

inner setae, cutting edge crenulate, setal 

row at base of dactylus of three setae; 

dactylus with proximal seta. 

Pereopod 1 (Figure 3C) longer than 

other pereopods, coxa with seta and 

rounded apophyses; basis slender, 4.1 

times as long as wide, with dorsoproximal 

seta; ischium compact with one seta; 

merus just shorter than carpus; carpus 

with three distal setae, longest of which is 

0.4 times length of propodus; propodus 

as long as carpus and merus together, 

with three dosrodistal setae on slight 

mound, one ventrodistal seta; dactylus 

slender, extending into shorter slender 

unguis, the two together as long as propodus. 

Pereopod 2 (Figure 3D) more compact 

than pereopod 1; basis 3 times as 

long as wide; ischium with 2 setae, longer 

seta longer than ischium width; merus 

longer than carpus, merus with strong 

ventrodistal spine, carpus with shorter 

ventrodistal spine; propodus with paired 

dorsodistal setae and ventrodistal spine; 

merus, carpus and propodus with ventral 

microtrichia; dactylus and short unguis 

together 0.6 times as long as propodus; 

dactylus (Figure 3E) with collar of fine 

setules at half length and dorsodistal 

setule. Pereopod 3 (Figure 3F) similar to 

pereopod 2, including longer seta on 

ischium and dactylus setulation, but carpus 

longer than merus and with short 

outer distal seta. 

Pereopod 4 (Figure 3G) basis stout, 2.4 

times as long as wide; ischium with two 

short setae; merus shorter than carpus, 

merus with two short, ventrodistal 

spines, carpus with outer, ventral and 

inner curved distal spines; propodus 

longer than carpus, with two ventrodistal 

short spines, four dorsodistal setae, one 

as long as dactylus; dactylus and unguis 

partially fused into a claw, curved. 

Pereopods 5 (Figure 3H) and 6 as pereopod 

4, but with pereopod 5 propodus 

with fewer distal setae. 

Pleopods all alike, typical for the 

genus, with single dorsal plumose seta on 

basis. 

Uropod (Figure 2M) biramous, basis 

naked; exopod of one segment, 0.7 times 

as long as proximal endopod segment, 

outer distal seta longer than inner distal 

seta; endopod of six segments, distal segments 

slender. 

Description of male: typical primary male, 

half length of female (allotype length 

1.7 mm), body (Figure 2B, C) more compact, 

cephalon just longer than pereonites 

1 to 3 together, with large eyelobes bearing 

conspicuous black eyes; pereonite 1 

shortest, pereonites 2 to 6 progressively


FIGURE 3. Leptochelia caldera sp. nov., A, female cheliped; B, male cheliped; C, pereopod 1; D, pereopod 

2; E, detail of distal articles of pereopod 2; F to H, pereopods 3, 4 and 5 respectively. Scale 

line = 0.2 mm for A to D and F to H, 0.1 mm for E.

192 AÇOREANA 

2009, Sup. 6: 183-200 

longer, pereonite 4 1.7 times as long as 

pereonite 1. Five free pleonites, subequal 

in length, pleotelson just longer than 

pleonite 5. Sexual dimorphism as follows. 

Antennule (Figure 2E) elongate, first 

peduncle article arcuate, 4 times as long 

as wide; second article 0.6 times as long as 

first with ventrodistal penicillate setae 

and 2 midventral simple setae; third article 

0.6 times as long as second, with dorsodistal 

seta; flagellum of 7 segments, 

first with 4 proximal and 5 distal aesthetascs, 

segments 2 to 5 with 4, 4, 3 and 3 

distal aesthetascs respectively; segments 6 

and 7 more slender than others. Antenna 

(Figure 2F) similar to that of female but 

more compact. Mouthparts atrophied. 

Cheliped (Figure 3B) larger than that 

of female; carpus slender, 3.6 times as 

long as wide, with ventrodistal invagination 

to accommodate propodus on reflexion; 

propodus fixed finger shorter than 

palm, with two inner tooth-like apophyses 

on cutting edge; moveable finger with 

spinules along cutting edge. 

Pleopods more setose than those of 

female. 

Etymology: caldera (from the Spanish 

cauldron) is a volcanic crater, the 2006 

type-material of this species having been 

collected from the sea-water-flooded 

crater of Ilhéu de Vila Franca. 

Remarks: there are four recorded species 

of Leptochelia with only 3 maxilliped 

basis setae, none of them occurring in the 

North Atlantic or Mediterranean, viz. 

L. itoi Ishimaru, 1982 (from Japan), 

L. lusei Bamber & Bird, 1997 (from Hong 

Kong), L. nobbi Bamber, 2005 (from 

Western Australia) and a species from 

Queensland, Australia (Bamber, in 

press). Of these four, only L. nobbi has a 

proximal antennule article more than 3 

times as long as wide, but that species 

has a compact basis to pereopod 1 (only 

three times as long as wide) and a uropod 

exopod only half as long as the 

proximal endopod segment length. 

Despite its much more compact proximal 

antennule peduncle article (2.5 times 

as long as wide), L. itoi shows most similarity 

to L. caldera sp. nov., but the cheliped 

basis of the Japanese species is 

more compact (1.5 times as long as 

wide), the distal seta of antennule 

peduncle article 2 is as long as the article 

(shorter in L. caldera), and the dactylus 

plus claw of the first pereopod is much 

longer than the propodus (1.33 times as 

long, compared with subequal in length 

in L. caldera). 

The number of maxilliped basis setae 

in L. neapolitana Sars, 1882 is not known, 

but that species again differs from 

L. caldera in that the proximal antennule 

peduncle article is shorter (three times as 

long as wide), the uropod exopod is less 

than half the length of the proximal 

endopod segment, the dactylus plus 

claw of the first pereopod is much longer 

than the propodus (1.36 times as long) 

and the cheliped basis more compact 

(1.35 times as long as wide). 

The male of the present species is 

generally very similar to the male of 

L. dubia sensu Sars, 1886 (non Krøyer, 

1842), but that species has longer anterior 

pereonites, and is without the degree 

of ventrodistal invagination on the cheliped 

carpus shown by L. caldera (and the 

female of Sars’ species has five maxilliped 

basis setae). The male of L. neapolitana 

has a similar cheliped to the present 

species, but has fewer antennule flagellum 

segments and more attenuate pereonites. 

None of the species described previously 

has the peculiar collar of setules at 

half length on the dactylus of pereopods 

two and three shown by Leptochelia 

caldera.


Dollfus (1897) recorded a male and a 

female of Leptochelia savignyi Krøyer, 

1842 from Horta at 5 to 6 m depth. This 

taxon has been the subject of some confusion 

over the last 150 years (including 

erroneous synonymy with L. dubia 

Krøyer, 1842), and it is now apparent 

that numerous species of Leptochelia 

await distinction based on morphological 

features of both genders which were 

not examined in any detail before 

Ishimaru (1985) (see Bamber, 2005 for 

discussion). From his subsequent report 

on Mediterranean and Atlantic species 

(Dollfus, 1898), it is apparent that 

Dollfus distinguished L. savignyi correctly 

on the basis of four longer articles in 

the antennule of the female (inter alia). 

There is debate whether the extra antennular 

article is an intermediate feature of 

a female changing into a male in a genus 

known to show progynous hermaphroditism 

(e.g. Smith, 1906); while this trend 

has been observed in some taxa of 

Leptochelia (R. Heard, pers. comm.), the 

male of L. savignyi sensu Sars, 1886 is a 

primary male, while Larsen & Rayment 

(2002) found this antennular structure a 

consistent feature of females of their new 

species L. elongata, including an ovigerous 

paratype. Dollfus (1898) reported a 

number of collections including females 

attributed to L. savignyi from the 

Mediterranean and the eastern Atlantic, 

implying a frequency of this antennular 

morphology unlikely to be shown only 

by transitional hermaphrodite specimens. 

His records from the Azores are 

thus accepted as valid, and L. savignyi 

sensu Sars, 1886 (see Figure 6A, B) is 

accepted as the same as L. savignyi 

Krøyer, 1842. 

Krøyer (1842) named a third species 

of Leptochelia, “Tanais” edwardsii, also 

from Madeira, but based only on the 

male. While it is possible that this taxon 

may never be confirmed (it is assumed to 

be the male of L. savignyi), from his figure 

(Krøyer, 1842: pl.2: Figures 13-19) it 

does not have the same antennular or 

pleotelson proportions as L. caldera. 

Family Paratanaidae Lang, 1949 

Sufamily Paratanaidinae Lang, 1949 

Genus Paratanais Dana, 1852 

Paratanais martinsi sp. nov. 

Figures 4, 5 

?Paratanais euelpis Monod, 1925, non 

Paratanais euelpis Barnard, 1920. 

Non- Paratanais atlanticus Dollfus, 

1897 (incertae sedis) 

Material: 1 female with brood pouch (in 

tube), holotype (NHM.2007.438), 1 female 

dissected, 2 females, 1 manca, 1 headless 

female, paratypes,(NHM.2007.439-440), 

WVF040, off Amora, Ponta Garça, São 

Miguel, Azores, 37º42’720”N 

25º21’554”W, 37.8 m depth, small dredge 

sample, 26 July 2006, coll. António de 

Frias Martins & Jerry Harasewych. 

Description of female: body (Figure 4A) 

elongate, slender, 4.2 mm long, eight 

times as long as wide, colour translucent 

white, eyes black. Cephalothorax subrectangular, 

1.2 times as long as wide, with 

slight rounded rostrum, single mid-lateral 

setae; eyes present, pigmented. Six free 

cylindrical pereonites; pereonite 1 shortest, 

with single anterolateral setae; pereonites 

2 and 3 subequal, twice as long as 

pereonite 1, also with single anterolateral 

setae; pereonites 4 and 5 subequal (pereonite 

4 longest), 1.2 times as long as pereonite 

2, pereonite 6 just shorter than pereonite 

2 (all pereonites respectively 2, 1.1, 

1.0, 0.9, 0.9 and 1.2 times as wide as long). 

Pleon of five free subequal pleonites bearing 

pleopods; pleonites 4.5 times as wide 

as long; pleonites 1 to 4 with one 

plumose, articulating lateral seta on each 

side, pleonite 5 with simple lateral seta.

194 AÇOREANA 

2009, Sup. 6: 183-200 

Pleotelson semicircular, short, 1.9 times as 

wide as long, distally with paired dorsal 

and paired terminal setae. 

Antennule (Figure 4B) of four articles, 

proximal article 2,3 times as long as wide, 

second article 1.3 times as long as wide, 

about one-third length of first, with dorsal 

seta longer than article length; third 

article nearly two-thirds length of second; 

distal article slender, longer than second 

and third articles together, with five distal 

setae and single aesthetasc. 

Antenna (Figure 4C) of six articles, 

proximal article compact, naked; second 

article with long ventrodistal and short 

dorsodistal apophyses each bearing seta; 

third article as long as wide, naked, with 

dorsodistal spine; fourth article just 

longer than second, with two distal simple 

setae; fifth article half as long as 

fourth with two distal setae; sixth article 

minute with five longer and one shorter 

distal setae. 

Labrum (Figure 4D) apically rounded, 

setose. Left mandible (Figure 4E) 

with crenulate pars incisiva and wide, 

crenulate lacinia mobilis; pars molaris 

robust with elaborate marginal “teeth”. 

Right mandible (Figure 4F) without 

lacinia mobilis, pars molaris less elaborate. 

Labium (Figure 4G) simple, finely 

setose, with fine outer marginal spinule, 

without palp. Maxillule (Figure 4H) 

with seven longer and two shorter distal 

spines, palp slender with two long distal 

setae; maxilla ovoid, naked. Maxilliped 

(Figure 4I) endites characteristic of 

genus, wide with denticulate outer margin, 

two inner distal ovate tubercles and 

single inner seta; palp first article naked, 

second article inner margin with two 

simple setae and shorter distal spine, 

outer margin with distal seta; third article 

with three inner bidenticulate spines, 

adjacent shorter simple spine; fourth 

article with four inner bidenticulate 

spines, single inner submarginal and 

outer simple setae; single inner spine on 

basis exceeding distal margin of endites. 

Epignath (Figure 4J) ribbon-like, 

glabrous but with two fine distal setae. 

Cheliped (Figure 5A) compact, carpus 

1.2 times as long as wide; propodus 

wider than long, fixed finger short with 

lamellate apophyses on cutting edge, terminal 

spine indistinct; dactylus with 

dorsoproximal simple seta. 

Pereopod 1 (figue 5B) longer than 

others, coxa simple with seta; basis slender, 

arcuate, five times as long as wide; 

ischium compact with single seta; merus 

1.5 times as long as carpus; propodus 1.2 

times as long as merus, with one ventral 

and three dorsal distal setae; dactylus 

with distinct, slender claw, both together 

as long as propodus. Pereopod 2 (Figure 

5C) similar to pereopod 1, but more compact, 

basis 3.1 times as long as wide, 

ischium with tow setae, merus shorter 

than carpus with ventral microtrichia 

and ventrodistal spine, carpus with ventral 

microtrichia, two ventrodistal and 

one larger inner distal spines. Pereopod 

3 (Figure 5D) similar to pereopod 2. 

Pereopod 4 (Figure 5E) basis robust, 2.2 

times as long as wide; merus 0.8 times as 

long as carpus, each with spination as 

pereopod 3; propodus longer than carpus 

with mid-dorsal penicillate seta, 

dorsodistal slender spine and ventrodistal 

stout spine; dactylus and claw forming 

unguis, curved, two-thirds as long as 

propodus. Pereopod 5 (Figure 5F) as 

pereopod 4. Pereopod 6 (Figure 5G) as 

pereopod 4, but propodus more compact 

with two ventrodistal spines and three 

dorsodistal setae adjacent to slender 

spine. 

Pleopods (Figure 4K) all alike, with 

naked basis, endopod with single inner 

plumose seta; exopod without setae on 

inner margin. 

Uropod (Figure 5H) basis naked, 

endopod of two segments, exopod of one


FIGURE 4. Paratanais martinsi sp. nov., A, holotype, dorsal; B, antennule; C, antenna; D, labrum; 

E, left mandible; F, right mandible; G, labium; H, maxillule and maxilla; I, maxilliped; J, epignath: 

K, pleopod (plumose nature of all setae not shown). Scale line = 1 mm for A, 0.2 mm for B and C, 

0.1 for D to J.

196 AÇOREANA 

2009, Sup. 6: 183-200 

segment, shorter than proximal segment 

of endopod. 

Male unknown. 

Etymology: named after António de Frias 

Martins, in gratitude for his assistance in 

attending the workshop, and exemplary 

hospitality. 

Remarks: the genus Paratanais has been 

capably diagnosed by Lang (1973). The 

only species from the North Atlantic 

attributed previously to Paratanais are all 

incertae sedis. Bate & Westwood (1868) 

described “Paratanais” rigidus from one 

specimen collected from Laminaria holdfasts 

off Glasgow, western Scotland: while 

their four-articled antennule is appropriate, 

they carefully describe the uropod 

rami as both being of one segment, and an 

elongate, slender chela; their species is 

not a member of the genus Paratanais, but 

with the inadequate description and figures 

must remain incertae sedis. P. limicola 

FIGURE 5. Paratanais martinsi sp. nov., A, cheliped; B to G, pereopods 1 to 6 respectively; H, uropod. 

Scale line = 0.2 mm for A to G, 0.15 mm for H.


Harger, 1878 (see Harger, 1880, for 

description and figures) has a three-articled 

antennule, and the uropod has two 

segments in the exopod and five in the 

endopod: Harger himself (1880) moved 

this species, apparently correctly, to 

Leptochelia. Dollfus (1897) described 

“Paratanais” atlanticus from 130 m depth 

off the Azores, based on a male and two 

females; although the description is 

somewhat cursory, and the figures inadequate, 

his species clearly had a three-articled 

antennule in the female, and the uropod 

had a two-segmented exopod and a 

three-segmented endopod; Dollfus’ 

species thus cannot be a member of the 

genus Paratanais. Finally, Monod (1925) 

suspected a specimen from 110 m depth 

off Morocco to be P. euelpis Barnard, 1920 

(a species adequately refigured by Lang, 

1973), but without full confidence (and 

without description or figure). It is possible 

that his specimen, if indeed of this 

genus, was of the present species. 

In having the setose apophyses on the 

second article of the antenna, more characteristic 

of species of Leptochelia, 

Paratanais martinsi sp. nov. is similar only 

to P. gaspodei Bamber, 2005, with which it 

also shares the elongate, slender uropod 

segments, and the short chela. The present 

species differs from P. gaspodei in a 

number of characters, including being 

generally more slender (pleonites 3, 4 and 

5 as long as or longer than wide, all wider 

than long in P. gaspodei), with more slender 

articles in the antennule and antenna 

(proximal peduncle article of antennule 

less than twice as long as wide in 

P. gaspodei), in the inner spines on the 

maxilliped basis exceeding the distal margin 

of the endites (not reaching the margin 

in P. gaspodei), and with the merus of 

pereopod 1 being 1.5 times as long as the 

carpus (subequal in length in P. gaspodei). 

With regard to the possible Moroccan 

record of Monod (1925), P. euelpis also has 

a shorter merus to pereopod 1, and is 

without the apophyses on the second article 

of the antenna, as well as differences 

in mouthpart setation. 

TENERIFE MATERIAL 

Superfamily Tanaoidea Dana, 1849 

Family Tanaidae Dana, 1849 

Subfamily Pancolinae Sieg, 1980 

Genus Zeuxo Templeton, 1840 

Zeuxo (Parazeuxo) exsargasso Sieg, 1980 

Figure 6 C, D 

Zeuxo (Parazeuxo) exsargasso 

1980, 217-221, figure 61. 

Sieg, 

Material: 3 males, 3 females with 

oostegites, 1 brooding female 

(NHM.2007.757-763), 1 female with 

oostegites (dissected), rocky intertidal 

platform covered in algal turf, Playa de 

Fanabe, Costa Adeje, Tenerife, 27 June 

2007. Coll. B Morton. 

Remarks: Zeuxo exsargasso was only 

known from the type collection from 

floating Sargassum natans, 20 miles southeast 

of Bermuda (Sieg, 1980). Its presence 

in the Canary Islands implies the possibility 

of transport by drift from America via 

the Gulf Stream and the Azores and 

Canary Currents. The only other recorded 

species of Zeuxo in north-eastern 

Atlantic waters is the only-distantly-related 

Zeuxo (Zeuxo) holdichi Bamber, 1990, 

known from the Atlantic shores of France 

and Portugal and the English Channel 

(Bamber, 1990; & unpubl. data). Zeuxo 

species are distinguished from Tanais 

species in their having five dorsallydemarcated 

pleonites (Tanais has only 

four), and no dorsal rows of plumose 

setae on the pleon (Tanais has conspicuous 

rows on pleonites 1 and 2); the uropod 

of Z. exsargasso has five segments 

(Figure 6D).

198 AÇOREANA 

2009, Sup. 6: 183-200 

DISCUSSION 

There are now five species of shallowwater 

tanaidacean recorded from the 

Azores, Tanais grimaldii, Leptochelia savignyi 

sensu stricto, Leptochelia caldera, 

Paratanais martinsi and the unconfirmed 

Leptognathia from Ponta da Ferraria. 

The present data demonstrate that 

Azorean tanaidacean fauna inhabits littoral 

to infralittoral algae, is generally 

sparse, but may show a relatively high 

degree of endemism. For the only species 

occurring in the 1996 and 1997 data in sufficient 

numbers for interpretation, Tanais 

grimaldii, no trends were detected in relation 

to disturbance or exposure. 

Unfortunately, these taxa give little 

information on the origins of the Azorean 

fauna. The Leptognathia specimen tells 

nothing. The two species described as 

new above are as yet unknown from elsewhere: 

Leptochelia is a worldwide genus, 

while Paratanais is predominantly southern 

hemisphere in distribution (Australia, 

Subantarctica, South Africa) but also 

found in the Indo-West Pacific, the Kurile 

Islands and California. 

T. grimaldii has been recorded from 

the Mediterranean (Adriatic) by Sars 

(1886, as T. cavolinii), his figure clearly 

showing the appropriate uropod structure. 

Sieg (1980) attributes the record of 

T. chevreuxi from the Moroccan coast by 

Monod (1925) to T. grimaldii, but regards 

it as doubtful, as he did the record from 

the Bay of Naples by Smith (1906); neither 

author gives a description or figure, and 

the decision of Sieg (loc. cit.) is based on 

their attributing the authority for their 

name to Sars, 1886. Both of these records 

are surely incertae sedis. There are no pub- 

FIGURE 6. A and B, Leptochelia savignyi: A, dorsal; B, antennule (redrawn after Sars, 1886). C and 

D, Zeuxo (Parazeuxo) exsargasso (Tenerife specimen): A, dorsal; B, pleotelson and uropods, dorsal.


lished records of Tanais species from the 

Canary Islands or from Madeira. 

While there has been confusion over 

the years regarding Leptochelia savignyi 

(see above), the type locality is Madeira, 

and the only other valid records (other 

than those of Dollfus, 1897; 1898) are 

those of Sars (1886) from the 

Mediterranean, from the Ligurian Sea 

and off Sicily. 

Thus there are a few indications that 

the Azorean tanaidacean fauna may have 

links with the Mediterranean, but not 

elsewhere. Morton & Britton (2000) 

found that most components of the 

Azorean marine fauna show affiliation 

with the Mediterranean and southern 

Europe. However, the discovery of Zeuxo 

exsargasso in Tenerife strongly implies colonization 

from the western Atlantic to 

Macaronesia via the Gulf Stream, the 

Azores Current and the Canary Current 

(see Timmermann, 1932, for a discussion 

on faunistic transport in Sargassum). 

It is undoubtedly the case that the 

tanaidacean fauna of Macaronesia is very 

understudied, and that of the 

Mediterranean and Atlantic coasts of 

Europe and North Africa is little better 

known, still relying heavily on 19th century 

information. At the same time the 

speciation of such taxa as Leptochelia 

around the North-east Atlantic and 

Mediterranean needs proper study. 


We are indebted to Roni Robbins for 

assistance in the field collecting and sample 

analysis in 2006, to Andreia Salvador 


Franca, to João Brum for assistance in the 

diver collections in 1996 and 1997, to 

Brian Morton for collecting the Tenerife 

material, to António de Frias Martins for 

the organisation of, and inviting us to, the 

2006 Workshop, and to the other participants 

at the workshop for samples and 

for entertainment. 

LITERARURE CITED 

BAMBER, R.N., 1977. On mobile littoral 

environments. Porcupine Newsletter, 

1(4): 62-63. 

BAMBER, R.N., 1990. A new species of 

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THE SOFT-SEDIMENT INFAUNA OFF SÃO MIGUEL, AZORES, AND A 

COMPARISON WITH OTHER AZOREAN INVERTEBRATE HABITATS 

Roger N. Bamber & Roni Robbins 

The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. e-mail: roger.bamber@artoo.co.uk 

ABSTRACT 

During the Third International Workshop of Malacology and Marine Biology in the 

Azores in July 2006, sampling of the littoral and sublittoral soft-sediment benthos around 

Vila Franca do Campo, São Miguel, was undertaken in order to characterize the benthic 

infaunal communities, and to compare the faunal density and diversity with the communities 

associated with algae. 

The sedimentary infauna was particularly sparse, with the average number of species 

and individuals per sample being 4.4 and 25 respectively. Although density and diversity 

were greatest between 20 and 40 m depth, there was no community trend with depth 

down to 250 m. The dominant taxa were actively mobile species. By contrast, the fauna 

of algal habitats was far denser and more species rich: these samples had an average of 17 

species and 360 individuals per sample. 

These results indicate an unstructured, impoverished sedimentary infauna, dominated 

by errant taxa which can tolerate unstable sediments, and not well-characterized as a 

community. The impoverishment of sedimentary benthos of this part of the Azores is 

attributed to sediment instability. 

SUMÁRIO 

Durante o 3º Workshop Internacional de Malacologia e Biologia Marinha nos Açores 

em Julho de 2006, foram feitas amostragens do bentos do sedimento móvel do litoral e 

sublitoral perto de Vila Franca do Campo, São Miguel, com vista a caracterizar as 

comunidades bentónicas da infauna e comparar a densidade e diversidade da infauna com 

as comunidades associadas às algas. 

A infauna sedimentar era particularmente esparsa, sendo o número médio de espécies 

e indivíduos por amostragem 4,4 e 25, respectivamente. Embora a densidade e a 

diversidade fossem as mais elevadas entre os 20 e os 40 m de profundidade, não se notou 

tendência na comunidade com a profundidade até aos 250 m. Os taxa dominantes eram 

espécies activamente móveis. Em contraste, a fauna dos habitats algais era de longe mais 

densa e mais rica em espécies: ali se registou a média de 17 espécies e 360 indivíduos por 

amostra. 

Estes resultados indicam uma infauna sedimentar não estruturada, empobrecida, 

dominada por taxa errantes que podem tolerar sedimentos instáveis, e não bem 

caracterizada coo comunidade. O empobrecimento do bentos sedimentar desta parte dos 

Açores atribui-se à instabilidade do sedimento. 

INTRODUCTION 









Drift, a diffuse southerly arm of the Gulf 

Stream breaking off from the North 


Americas, while the somewhat less-sig-

202 AÇOREANA 

2009, Sup. 6: 201-210 

nificant western eddies of the Canary 



current brings warm, hyperhaline water 

from the Mediterranean outflow (Gofas, 

1990; Morton et al., 1998; Morton & 

Britton, 2000). This hydrography clearly 

has implications for the colonization of 

the islands by benthic marine species. 

While there have been a number of 

previous studies on the fauna associated 

with the rocky shores, or with littoral and 

infralittoral algae (e.g. Hawkins et al., 

1990; Bullock et al., 1990; Bullock, 1995; 

taxa reviewed by Morton & Britton, 2000), 

and which have normally been taxon specific 

(see papers by Chapman et auctt. in 

Morton, 1990), there have been few published 

studies of the soft-sediment communities, 

largely owing to their recognized 

sparseness. Morton (1990) points 

out that there was at that time no information 

on the soft shores of the Azores, as 

virtually all are high energy beaches, 

“superficially devoid of significant life” 

(see also Morton et al., 1998). Two surveys 

had been conducted of the soft-sediment 

fauna of the sheltered habitat within the 

flooded crater of Ilhéu de Vila Franca, one 

in 1988 and one in 1995 (Morton, 1990; 

Wells, 1995 respectively). 



Biology in the Azores in July 2006, sampling 

of the littoral and sublittoral softsediment 

benthos off Vila Franca do 

Campo was undertaken in order to gain 

some insight into the communities present. 

Some sites were sampled within the 

flooded crater of the Ilhéu de Vila Franca 

to compare with the findings of the 1988 

and 1991 studies. Comparisons were 

made between the invertebrate fauna of 

quantitative (grab plus shore collected) 

and qualitative (dredge) benthic samples, 

as well as the fauna associated with 

infralittoral algae. 

METHODS 

Quantitative samples were collected 

using a 0.025 m 2 Peterson grab, taking 

two replicates per sampling station. 

Littoral and infralittoral sands were sampled 

quantitatively using a plastic scoop, 

also to 0.05 m 2 . The samples were sieved 

across a 0.5 mm mesh and sorted live. 

Qualitative samples were collected 

using a variety of dredges, towed at 1 to 2 

knots for between 6 and 10 minutes. 

Crustaceans, pycnogonids, polychaetes, 

sipunculans, phoronids and echinoderms 

from these samples were retained 

serendipitously. 

The depths of the grab and dredge 

samples ranged between 12 and 250 m. 

Position fixing was by GPS. 

Qualitative algal samples were collected 

either from littoral rocks (sample 

Island 24.10), or from sublittoral sites in 

the crater of Ilhéu de Vila Franca. 

Material as collected from the dredge 

samples was also analyzed from one 

SCUBA collection on algae, sponges, etc., 

at 16 m depth. 

All taxa were identified to species 

where possible. Nomenclature and 

authorities are as in Costello et al. (2001). 

Sample numbers are prefixed “Island” for 

those collected within the Ilhéu de Vila 

Franca, otherwise “WVF”. 

RESULTS 

The sampling sites analyzed are listed 

in Table 1, with depths, sampling gear 

and univariate community statistics. 

Quantitative samples 

The twelve grab and shore-collected 

quantitative samples were analyzed 

together. The complete faunal data from 

these samples is given in Appendix 1. It 

is immediately apparent that the benthic 

infauna is indeed sparse: only one of the

BAMBER & ROBBINS: THE SOFT-SEDIMENT INFAUNA OFF SÃO MIGUEL 203 

TABLE 1. List of samples, with depth, sampling gear and univariate ‘community’ statistics. 

Sample Date depth, 

m 

sampling 

gear 

species individuals species 

richness 

Shannon- 

Weiner H' 

Evenness 

WVF005 19/7/06 42 dredge 7 15 2.216 2.28 0.8121 

WVF006 19/7/06 40 dredge 10 12 3.622 3.252 0.9788 

WVF007 19/7/06 178 dredge 10 26 2.762 2.887 0.8689 

WVF008 19/7/06 148 dredge 4 6 1.674 1.792 0.8962 

WVF011 20/7/06 16 scuba 26 196 4.574 3.861 0.8315 

WVF015 21/7/06 46 dredge 11 17 3.53 3.293 0.9518 

WVF016 21/7/06 19 dredge 8 24 2.203 2.772 0.924 

WVF019 21/7/06 23 grab 16 91 3.325 3.033 0.7581 

WVF020 21/7/06 50 grab 7 14 2.274 2.407 0.8573 

WVF021 21/7/06 118 grab 2 2 

Island24.1 24/7/06 0 scoop 1 17 

Island24.2 24/7/06 drift weed 16 129 3.087 2.845 0.7113 

Island24.3 24/7/06 0.5 scoop 0 

Island24.4 24/7/06 0.3 weed 25 719 3.505 3.328 0.7259 

Island24.5 24/7/06 0.5 weed 27 606 3.913 3.456 0.7352 

Island24.6 24/7/06 0.3 scoop 0 

Island24.7 24/7/06 0.5 scoop 0 

Island24.9 24/7/06 0.3 scoop 0 

Island24.10 24/7/06 0 weed scrape 3 7 

WVF034 25/7/06 16.7 grab 5 12 1.61 1.781 0.7669 

WVF035 25/7/06 23 grab 8 46 1.828 2.519 0.8398 

WVF036 25/7/06 36 grab 8 105 1.504 1.668 0.556 

WVF039 25/7/06 12 grab 1 1 

WVF040 26/7/06 38 dredge 5 20 1.335 1.882 0.8106 

WVF041 25/7/06 250 dredge 6 12 2.012 2.355 0.9112 

littoral/infralittoral samples contained 

any macrofauna, that on the sand bar in 

Ilhéu de Vila Franca (Island24.1) containing 

17 individuals of the errant isopod 

Eurydice affinis. Similarly, the grab samples 

at 12 m (WVF039) contained only a 

single specimen of the bivalve Ervilia castanea. 

Figure 1 shows the distribution 

with depth of faunal density (numbers 

per 0.05 m2), number of species and 

Shannon-Weiner diversity. The greatest 

faunal density and number of species 

occur between 20 and 40 m depth, but values 

are consistently low, with the average 

number of species and individuals per 

sample being 4.4 and 25 respectively. 

The community diversity is generally 

low, Shannon-Weiner index values ranging 

between 1 and 3, and also peaks at 

around 20 to 40 m depth, although, owing 

to the proportionately higher number of 

species (7) in a sparse fauna (n=14) at station 

WVF020, diversity remains in the 

upper range to 50 m. 

Dominant taxa were actively mobile 

species, the mysid Gastrosaccus normani, 

the amphipods Microdeutopus versiculatus 

and Harpinia laevis, the polychaete 

Armandia polyophthalma and the predatory 

polychaete Glycera capitata. The most 

numerous sessile species was the tubicolous 

polychaete Myriochele oculata,

204 AÇOREANA 

2009, Sup. 6: 201-210 

FIGURE 1. Univariate community parameters by depth for the sedimentary infauna, quantitative 

samples only. 

although that species was constrained to 

two stations. Indeed, of the 28 species 

recorded, 21 are actively mobile species 

less reliant on sedimentary-habitat stability. 

Qualitative samples 

The complete faunal data from the 

dredge samples is given in Appendix 2. 

These samples are not directly comparable 

as no molluscs were analyzed (they 

having been removed for other studies), 

and the univariate community statistics 

are only indicative, as the samples were 

not quantitative (although the Shannon- 

Weiner diversity index is relatively sample-size-independent). 

The fauna was again found to be 

sparse and inconsistent, 29 of the 42 

species recorded occurring in only one 

sample. Although no dredges shallower 

than 19 m were analyzed, there was an 

indication that the numbers of species 

and individuals were highest around 19 

to 50 m depth, similar to the results from 

the grab sample results, but the sample at 

178 m (WVF007) had high density and 

species number. Average numbers of 

species and individuals were 7.5 and 16.5 

respectively. 

The dominant taxa were again actively 

mobile species, including amphipods, 

decapods, errant polychaetes, while the 

only obligately sessile taxa were tubicolous 

species, the tanaidacean Paratanais 

martinsi, the spionid polychaete Spio 

armata and the phoronid Phoronis muelleri, 

each occurring only once. 

Overall, although extra species were 

found from the dredge samples (which 

had taken a larger volume per sample), 

the fauna was relatively similar to that 

found in the grab samples. 

Algal and other samples 

The non-sedimentary samples were 

analyzed in order to compare faunal densities 

and species complements of the 

non-infaunal community. 

Appendix 3 lists the complete faunal 

data from these samples, which comprise 

a scuba-collection (WVF011), floating 

(Island 24.2), submerged (Island 24.4, 

island 24.5) or littorally attached


(Island 24.10) algae in the Ilhéu de Vila 

Franca lagoon. 

It is immediately apparent that the 

fauna of these habitats was far denser and 

more species rich. Even though the littoral 

rock-pool algal sample was impoverished, 

the algal samples had an average of 17 

species and 360 individuals per sample. 

The dominant species were the crevicial 

or algal-associated polychaete 

Platynereis dumerilii, and peracarid crustaceans, 

notably algal-associated 

amphipods (Hyale spp., Erichthonius spp., 

Microdeutopus versiculatus, Corophium acutum, 

Caprella acanthifera), isopods 

(Dynamene bidentata, Cymodoce truncata, 

Paranthura costana) and tanaidaceans 

(Tanais grimaldii), together with algalassociated 

pycnogonids (Anoplodactylus 

pygmaeus, A. angulatus) and the only 

cumacean recorded during the 2006 surveys, 

Cumella limicola. However, other 

dominant species (the polychaete Fabricia 

stellata, the amphipod Dexamine spinosa) 

are taxa which commonly occur in sediments, 

but which were not recorded away 

from the algae. 

Most of the species recorded from the 

scuba sample, collected from 16 m depth 

to the north-east off Ilhéu de Vila Franca, 

were the same as those dominant taxa 

from the algal samples, confirming this 

dense and diverse community is not 

restricted to the sheltered waters of the 

crater lagoon. 

DISCUSSION 

The data from both sets of sedimentary 

infaunal samples were subjected to 

multivariate analysis simply on presence 

or absence of species, owing to the qualitative 

nature of the dredge samples. The 

data set was reduced to those 30 taxa 

occurring as more than one individual in 

the survey, and to those samples with 

some fauna. 

The dendrogram of sample similarity 

(Figure 2) shows relatively poor clustering 

owing to the sparse and patchy nature 

of the fauna. Sample WVF040 is quite distinct, 

the three species recorded there 

(two pycnogonids and a tanaidacean) not 

being found elsewhere; this sample was 

not analyzed further. Similar reasons 

account for the isolation of samples 

WVF007, WVF008 and Island24.1. The 

remaining six grab samples and five 

dredge samples show that there is, in fact, 

no particular trend with depth in the 

community, dredge sample WVF041, 

taken at 250 m, falling comfortably within 

the subcluster of four grab samples taken 

at depths between 16.7 and 36 m. This 

lack of a trend is confirmed by the nonparametric 

ordination by multidimensional 

scaling (MDS) (Figure 3). 

Although the stations WVF007 and 

WVF008, both deeper than 140 m, are isolated, 

the species which distinguish them, 

Onuphis eremita and Ebalia tuberosa, are 

not species confined to deeper water. 

These results indicate an unstructured, 

impoverished sedimentary infauna, 

dominated by errant taxa which can 

tolerate unstable sediments, and not wellcharacterized 

as a community, thus not 

showing any particular community 

trends with depth. This fauna is most 

impoverished in shallow depths (

206 AÇOREANA 

2009, Sup. 6: 201-210 

FIGURE 2. Dendrogram by Bray-Curtis similarity (%) for sedimentary benthic infaunal samples 

(presence-absence data). 

which would most readily recruit. It was 

notable that the densest algal-associated 

communities occurred sublittorally, most 

littoral algae of the adjacent São Miguel 

shoreline supporting little of no fauna. 


We are indebted to Andreia Salvador 


Franca, to António de Frias Martins for 

FIGURE 3. MDS ordination based on the similarity 

data from figure 2, with circles of diameter 

proportional to sample depth. 

the organisation of, and inviting us to the 

Workshop, and to the other participants 

at the workshop for samples, assistance 

with the boat-work, discussion and entertainment. 


BULLOCK, R.C., 1995. The distribution 

of the molluscan fauna associated 

with the intertidal coralline algal turf 

of a partially submerged volcanic 

crater, the Ilhéu de Vila Franca, São 

Miguel, Azores. In: MARTINS, A.M.F. 


Azores (Proceedings of the Second 


and Marine Biology, São Miguel, 1991). 


BULLOCK, R.C., R.D. TURNER & R.A. 

FRALICK, 1990. Species richness and 

diversity of algal-associated micromolluscan 

communities from São 

Miguel, Azores. In: MARTINS, A.M.F. 



International Workshop of Malacology,


São Miguel, 1988). Açoreana, 


COSTELLO, M.J., C. EMBLOW & R. 

WHITE (eds.), 2001. European 

Register of marine Species. A checklist 

of the marine species in Europe 

and a bibliography of guides to their 

identification. Patrimoines Naturels, 

50: 463 pp. 









HAWKINS, S.J., L.P. BURNAY, A. NETO, 

R. TRISTÃO da CUNHA & A.M. de 

FRIAS MARTINS, 1990. A description 

of the zonation patterns of molluscs 

and other biota on the south coast of 

São Miguel, Azores. In: MARTINS, 

A.M.F. (ed.), The Marine Fauna and 

Flora of the Azores (Proceedings of the 


Malacology, São Miguel, 1988). 


MORTON, B., 1990. The intertidal ecology 

of Ilhéu de Vila Franca – a 

drowned volcanic crater in the 







MORTON, B. & J.C. BRITTON, 2000. The 

origins of the coastal and marine flora 

and fauna of the Azores. 





Açores, 249pp. Sociedade Afonso 


WELLS, F.E., 1995. An investigation of 

marine invertebrate communities in 

the sediments of Ilhéu de Vila Franca 

off the island of São Miguel, Azores. 

In: MARTINS, A.M.F. (ed.), The Marine 

Fauna and Flora of the Azores 



Biology, São Miguel, 1991). Açoreana, 

Supplement [4]: 57-65.

208 AÇOREANA 

2009, Sup. 6: 201-210 

APPENDIX 1. Complete benthic faunal data from quantitative sedimentary samples 

WVF WVF WVF Island Island Island Island Island WVF WVF WVF WVF 

Sample 019 020 021 24.1 24.3 24.6 24.7 24.9 034 035 036 039 TOTAL 

SIPUNCULA 

Golfingia minuta 2 2 

ANNELIDA 

Glycera capitata 9 1 6 6 22 

Glycera tesselata 3 3 

Glycinde nordmanni 1 1 

Eumida cf. bahusiensis 1 1 

Scoloplos armiger 1 1 

Spio armata 2 2 

Armandia polyophthalma 24 7 7 4 42 

Myriochele oculata 4 70 74 

Ditrupa arietina 1 3 4 

ARTHROPODA 

Crustacea 

Mysidacea 

Gastrosaccus normani 22 1 1 18 8 50 

Anchialina agilis 1 1 2 

Amphipoda 

Harpinia laevis 14 5 14 33 

Synchelidium haplocheles 

5 5 

Erichthonius punctatus 1 1 

Microdeutopus versiculatus 6 1 2 5 1 15 

Ampithoe rubricata 1 1 

Caprella penantis 1 1 

Isopoda 

Eurydice affinis 1 1 17 19 

Decapoda 

Atyaephyra desmaresti 1 1 

Processa edulis 1 1 

Crangon trispinosus 2 2 

Parthenope expansa 1 1 

MOLLUSCA 

Bivalvia 

Ervilia castanea 1 1 2 

Moerella donacina 1 1 

Solemya sp. 1 1 

ECHINODERMATA 

Echinocardium flavescens 

4 3 7 

Echinocyamus pusillus 1 3 1 5 

No. of Species 16 7 2 1 0 0 0 0 6 10 10 1 28 

No. of Individuals 91 14 2 17 0 0 0 0 13 53 109 1 300


APPENDIX 2. Faunal data from qualitative dredge samples (Mollusca not included) 

Sample WVF005 WVF006 WVF007 WVF008 WVF015 WVF016 WVF040 WVF041 TOTAL 

SIPUNCULA 

Golfingia margaritacea 1 1 

Golfingia minuta 1 1 

ANNELIDA 

Harmothoe spp. 4 3 7 

Glycera capitata 1 2 4 7 

Glycera tesselata 1 3 4 

Glycinde nordmanni 1 1 

Nereis pelagica 3 3 

Nereis diversicolor 2 2 

Eulalia cf. expusilla 1 1 

Spio armata 1 1 

Armandia polyophthalma 1 1 

Pisione remota 1 1 

Onuphis eremita 6 1 7 

Hyalinoecia tubicola 1 1 2 

ARTHROPODA 

Pycnogonida 

Achelia echinata 9 9 

Anoplodactylus virescens 1 1 

Anoplodactylus amora 3 3 

Crustacea 

Mysidacea 

Gastrosaccus normani 1 1 4 2 8 


Ampelisca spinipes 1 1 

Erichthonius punctatus 1 1 

Erichthonius difformis 7 1 4 12 

Microdeutopus versiculatus 

2 2 

Melita gladiosa 2 2 

Lembos websteri 1 1 


Eurydice affinis 1 1 3 5 

Tanaidacea 

Paratanais martinsi 6 6 


Processa edulis 1 2 3 

Crangon trispinosus 6 6 

Anapagurus laevis 1 1 

Paguridae indet. 1 2 3 

Galathea intermedia 1 3 4 

Scyllarus arctus 1 1 

Ebalia tuberosa 7 3 10 

Liocarcinus arcuatus 1 1 

Liocarcinus marmoreus 4 4 

Liocarcinus pusillus 1 1 

Pilumnoides inglei 1 1 

Parthenope expansa 1 1 2 

Macropodia rostrata 1 1 

PHORONIDA 

Phoronis muelleri 1 1 

ECHINODERMATA 

Echinocardium flavescens 2 2 

Echinocyamus pusillus 1 1 

No. of Species 7 10 10 4 10 8 5 6 42 

No. of Individuals 15 12 26 6 17 24 20 12 132

210 AÇOREANA 

2009, Sup. 6: 201-210 

APPENDIX 3. Faunal data from algal samples 

Sample WVF011 Island 24.2 Island 24.4 Island 24.5 Island 24.10 TOTAL 

SIPUNCULA 

Golfingia margaritacea 1 1 

ANNELIDA 

Harmothoe spp. 4 2 1 7 

Nereis pelagica 2 2 

Platynereis dumerilii 13 47 162 129 351 

Nainereis cf. laevigata 2 2 

Euphrosyne armadillo 1 1 

Polyophthalmus pictus 4 2 5 1 12 

Eupolymnia nebulosa 1 1 

Fabricia stellata 4 12 13 29 

ARTHROPODA 

Pycnogonida 

Achelia echinata 4 1 5 

Callipallene emaciata 6 6 12 

Anoplodactylus amora 0 

Anoplodactylus pygmaeus 

1 1 2 

Anoplodactylus angulatus 1 5 9 15 

Crustacea 


Ampelisca aequicornis A 3 3 

Ampelisca aequicornis B 3 3 

Erichthonius punctatus 12 1 8 3 24 

Erichthonius difformis 42 4 43 51 140 

Microdeutopus versiculatus 26 1 51 37 115 

Melita gladiosa 3 1 4 

Lembos websteri 13 13 

Ampithoe rubricata 3 1 13 7 1 25 

Dexamine cf. spinosa 12 27 31 25 95 

Hyale nilssoni 8 9 82 85 184 

Hyale perieri 4 4 

Corophium acutum 2 55 57 

Caprella penantis 2 4 2 8 

Caprella acanthifera 18 17 21 56 


Paranthura costana 1 1 11 9 22 

Eurydice affinis 18 18 

Dynamene bidentata 3 22 13 38 

Cymodoce truncata 3 7 5 15 

Janira maculosa 1 7 8 

Tanaidacea 

Tanais grimaldii 4 7 194 110 315 

Leptochelia caldera 15 2 17 

Cumacea 

Cumella limicola 6 11 11 28 


Thoralus cranchi 1 1 

Clibanarius erythropus 4 4 

Paguridae indet. 5 5 

Pilumnus hirtellus 2 2 

MOLLUSCA 

Gastropoda 

Setia subvaricosa 1 5 6 12 

Rissoa guernei 1 1 

No. of Species 26 16 25 27 3 41 

No. of Individuals 196 129 719 606 7 1657


SHELL OCCUPANCY BY THE HERMIT CRAB CLIBANARIUS ERYTHROPUS 

(CRUSTACEA) ON THE SOUTH COAST OF SÃO MIGUEL, AÇORES 

Pedro Rodrigues 1 & Roshan K. Rodrigo 2 

1 


Azores, Portugal. e-mail: pedrorodrigues@uac.pt 

2 

Faculty of Science, Department of Zoology, University of Colombo, Colombo 7, Sri Lanka 

ABSTRACT 

The importance of gastropod shells to hermit crabs is well known. The strong association 

between hermit crabs and their adopted shells influences almost all aspects of their 

biology and there is a strong correlation between the sizes of the shell and the crustacean. 

The hermit crab Clibanarius erythropus is abundant on the rocky shores of the Açores. 

However, there are few references to the ecology of this species. The aim of the present 

study was thus to evaluate the occupancy of mollusc shell species by C. erythropus on the 

south coast of the island of São Miguel. Shells of Columbella adansoni had the highest occupancy 

rate followed by Mitra cornea and Stramonita haemostoma. The significant differences 

in the estimated shell volumes available for C. erythropus suggests that juveniles choose 

Nassarius incrassatus, medium sized hermits prefer Pollia dorbignyi, C. adansoni and M. 

cornea, and the largest adults opt for S. haemastoma shells. 

RESUMO 

A importância das conchas de gastrópodes para os bernados-eremitas é bem 

conhecida. A forte associação entre os bernardos-eremita e as suas adoptadas conchas 

influencia grandemente quase todos os aspectos da sua biologia, existindo uma forte 

correlação entre o tamanho da concha e o tamanho do crustáceo. O bernado-eremita 

Clibanarius erythopus é uma espécie muito abundante nas costas rochosas. Todavia, são 

raras as referências à ecologia e à biologia desta espécie. O presente estudo teve como 

objectivo avaliar a ocupação de conchas de espécies de moluscos por C. erythropus na costa 

sul de São Miguel. Conchas de Columbella adansoni obtiveram a maior taxa de ocupação, 

seguidas de Mitra cornea e de Stramonita haemastoma. As diferenças significativas no 

volume de concha estimado acessível a C. erythropus sugerem que os juvenis escolhem 

conchas de Nassarius incrassatus, os de tamanho médio preferem conchas de Pollia 

dorbignyi, C. adansoni e M. cornea, e os adultos maiores optam por conchas de S. 

haemastoma. 

INTRODUCTION 

The importance of gastropod shells to 

hermit crabs is well known; such 

shells particularly supplying protection 

against predators (Vance, 1972) and physical 

stresses (Reese, 1969) due to the fact 

that the crabs have a soft, vulnerable 

abdomen. Shells can be found either 

empty, obtained by confrontations 

between individual crabs, or by removal 

of the gastropod (Elwood & Neil, 1992). 

The availability of shells is the main factor 

limiting populations of hermit crabs 

(Kellog, 1976) and they usually partition 

shell resources according to size 

(Bertness, 1981; Scully, 1983; Neil, 1985) 

and/or their own body size (Mitchell, 

1975), shell shape, weight and volume 

(Reese, 1963; Kuris & Brody, 1976; 

Conover, 1978). Interspecific competition 

among hermit crabs for shell resources 

may also be avoided by partitioning habitats. 

Partitioning also occurs between

212 AÇOREANA 

2009, Sup. 6: 211-216 

hermit crabs and other species such as 

sipunculans (Morton & Britton, 1995). 

Hermit crabs that occupy large shells 

can better resist to desiccation, thermal 

stress and predation (Rittschof et al., 

1995). However, heavy shells may limit 

reproduction and growth because of the 

high-energy cost of locomotion (Bertness, 

1981). For another hand, small shells render 

the crabs more vulnerable to predation 

and may reduce their growth rate 

(Hazlett, 1981; Angel, 2000). The strong 

association between hermit crabs and 

their adopted shells influences greatly 

almost all aspects of their biology 

(Hazlett, 1981) and there is a strong correlation 

between shell and crab sizes 

(Abrams et al., 1986; Botelho & Costa, 

2000; Sant’Anna et al., 2006). 

The hermit crab Clibanarius erythropus 

(Latreille, 1818) is a common species 

along the Mediterranean shores and 

Atlantic coasts from United Kingdom to 

the Açores (Ingle, 1993). In the Açorean 

archipelago, C. erythropus is abundant on 

rocky shores, including tide pools 

(Morton et al., 1998). However, references 

to the general ecology and biology of this 

species are few (Gherardi, 1991). 

The aim of the present study was to 

evaluate the occupancy of mollusc shell 

species by Clibanarius erythropus in the 

south coast of São Miguel island. 


FIGURE 1. Location of the sampling sites on 

São Miguel. A. Ilhéu de Vila Franca do Campo; 

B. Vila Franca do Campo Harbour; C. Roída da 

Praia; D. Ponta da Galera; E. Baía do Cruzeiro. 

The study was carried out on the 

south coast of São Miguel during July 

2006 as part of the 3 rd International 

Workshop on the Malacology and Marine 

Biology of the Açores covered in Vila 

Franca do Campo, São Miguel, Açores. 

Five populations of Clibanarius erythropus 

were studied for shell occupancy 

(Figure 1). 

The hermit crabs were hand-sampled 

by snorkelling for ten minutes to standardize 

capture effort and brought alive 

to the laboratory where individuals and 

shells were identified according to Wirtz 

(1995) and Morton et al. (1998). The occupied 

shells were measured (total shell 

length and shell width) using vernier callipers 

to the nearest 0.01 mm. The volume 

of each shell was estimated according to 

Morton & Britton (1995) by squaring the 

shortest linear measurement (width) and 

multiplying this value by the longest linear 

measurement (length). 

A Chi-square test was applied to the 

five most common occupied shell species. 

The relationship between those and the 

estimated volume was also subjected to 

analysis of variance (ANOVA) against the 

null hypothesis that different shell species 

have the same volume available for 

Clibanarius erythropus. 

RESULTS 

Eleven species of gastropod shells 

occupied by Clibanarius erythropus were 

collected and frequency of occupation at 

each site is identified in Table 1. 

Columbella adansoni Menke, 1853, Mitra 

cornea (Lamarck, 1811) and Stramonita 

haemastoma (Linnaeus, 1766) were the 

most commonly occupied shells, followed 

by Pollia dorbignyi (Payraudeau, 

1826) and Nassarius incrassatus (Ström, 

1768).

RODRIGUES & RODRIGO: SHELL OCCUPANCY BY CLIBANARIUS ERYTHROPUS 213 

TABLE 1. The frequency of mollusc shells occupied by Clibanarius erythropus from the different 

sampling sites on São Miguel. 

Mollusc shell Site A Site B Site C Site D Site E TOTAL 

Pollia dorbignyi (Payraudeau, 1826) 6 1 0 34 0 41 

Columbella adansoni Menke, 1853 54 41 51 67 31 244 

Stramonita haemastoma (Linnaeus, 1766) 6 15 81 25 15 142 

Mitra cornea (Lamarck, 1811) 1 46 0 27 84 158 

Nassarius incrassatus (Ström, 1768) 10 4 5 4 0 23 

Calliostoma lividum Dautzenberg, 1927 1 0 0 0 0 1 

Coralliophila meyendorffii (Calcara, 1845) 0 4 0 0 0 4 

Melarphe neritoides (Linnaeus, 1756) 0 0 0 1 0 1 

Littorina striata King & Broderip, 1832 0 0 0 0 5 5 

Jujubinus sp. 0 0 0 0 1 1 

Bittium cf. latreillii (Payraudeau, 1826) 0 0 0 0 2 2 

TOTAL: 78 111 137 158 138 622 

The Chi-square test, comparing the 

total number of the five commonest occupied 

shells from the different sampling 

sites, indicated a significantly different 

proportion of species occupied by 

Clibanarius erythropus (χ 2 = 386.1, p

214 AÇOREANA 

2009, Sup. 6: 211-216 

ability of certain gastropods influences 

their pattern of shell utilization in the natural 

habitat as the crabs tend to be opportunistic 

with regards to the shells they 

inhabit (Botelho & Costa, 2000). The significant 

differences in the estimated shell 

volume available for Clibanarius erythropus, 

the presence of empty shells (personal 

observations), and that the crab 

requires different shell sizes atdifferent 

stages of their growth (Morton & Britton, 

1995), suggests that juvenile C. erythropus 

individuals choose Nassarius incrassatus 

shells, middle size individuals prefer 

Pollia dorbignyi, Columbella adansoni and 

Mitra cornea shells, and larger adults opt 

for Stramonita haemastoma shells. This is 

in accordance with the study of Botelho & 

Costa (2000) where it was reported that 

hermit crabs of < 8.6 mm occupied all 

shell species, whereas those > 8.6 mm 

were found only in S. haemastoma shells. 

Smaller crabs occupied Littorina striata 

and N. incrassatus shells. 

ACKNOWLEDGMENTS 

We are grateful to Professor António 

M. de Frias Martins for the opportunity to 

undertake this research, and Professor 

Brian Morton for the review of this paper. 

This project was supported by the Third 

International Workshop on Malacology 

and Marine Biology, Vila Franca do 

Campo, São Miguel, Açores 

The experiments performed for the 

present study comply with the laws of the 

country in which they were performed. 


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Stephanie Naglschmid, Stuttgart.


A CONSERVATIONAL APPROACH ON THE SEABIRD POPULATIONS OF ILHÉU 

DE VILA FRANCA DO CAMPO, AZORES, PORTUGAL 

Pedro Rodrigues 1 , Joana Micael 1 , Roshan K. Rodrigo 2 & Regina T. Cunha 1 

1 


Azores, Portugal. e-mail: pedrorodrigues@uac.pt 

2 

Faculty of Science, Department of Zoology, University of Colombo, Colombo 7, Sri Lanka 

ABSTRACT 

This study was performed to identify the seabird species occurring on Ilhéu de Vila 

Franca do Campo (IVFC) off São Miguel island, Azores, giving a special emphasis on the 

description of their ecology and threats. Flush counts, ground searches and raft counts 

were conducted and two types of natural habitats were identified. The results confirmed 

the nesting of two endangered species and revealed three other possible breeders enhancing 

the importance of the islet for the protection and conservation of Azorean seabird populations. 

Although playing an important role on the conservation of Cory’s shearwater 

and Common tern populations, the islet can be threatened by continuous habitat degradation 

by human disturbance. The implementation of a habitat restoration program is 

suggested for the islet in a near future. 

RESUMO 

Foi feito um levantamento das espécies de aves marinhas que ocorrem no Ilhéu de Vila 

Franca do Campo, com especial ênfase para a sua ecologia e ameaças. Foram realizadas 

contagens visuais, procura de ninhos e contagem de jangadas ao longo de todo o ilhéu e 

zonas adjacentes, confirmando a nidificação de duas espécies ameaçadas e a possibilidade 

de nidificação de outras três. A identificação de dois tipos de habitats naturais evidencia 

a importância do ilhéu para a protecção e conservação das populações de aves marinhas. 

Apesar do importante papel na conservação das populações de cagarros e garajauscomuns, 

o ilhéu de Vila Franca do Campo continua a ser ameaçado por uma contínua 

degradação dos seus habitats devido à acção humana. Este trabalho sugere, num futuro 

próximo, a implementação de medidas proteccionistas e um programa de restauração dos 

habitats naturais deste ilhéu. 

INTRODUCTION 

The Azorean archipelago, located in the 

north Atlantic Ocean, has always been 

recognized as an interesting place for 

birds, mainly seabirds, not only due to the 

coast line with steep scarps but also to its 

geographical location (N36-39º, W25-31º) 

that represents an ornithological transition 

between temperate and tropical 

zones (Monteiro et al., 1996a, b). 

Thirteen seabird species are known to 

occur in the Azores. The regular breeders 

are Bulwer’s petrel (Bulweria bulwerii), 

Cory’s shearwater (Calonectris diomedia 

borealis), Manx shearwater (Puffinus puffinus), 

Little shearwater (Puffinus baroli), 

Band-rumped storm-petrel (Oceanodroma 

castro), Monteiro’s storm petrel 

(Oceanodroma monteiroi), Yellow-legged 

gull (Larus michaelis atlantis), Common 

tern (Sterna hirundo) and Roseate tern (S. 

dougallii). There are two occasional 

breeders, Red-billed tropicbird (Phaethon 

aethereus) and Sooty tern (Onychoprion fuscatus), 

a possible breeder, Cape Verde 

petrel (Pterodroma feae), and a possible former 

breeder, White-faced storm-petrel 

(Pelagodroma marina) (Le Grand et al., 

1984; Monteiro et al., 1996a).

218 AÇOREANA 

2009, Sup. 6: 217-225 

The archipelago accounts for the 

largest population of Cory’s shearwater of 

the world with more than 180.000 couples 

(79% of the European population) 

(Rodrigues & Nunes, 2002). Also representative 

are the populations of Bandrumped 

storm-petrel, 915 to 1240 couples 

(around 25% of the European population), 

Little shearwater, 800 to 1500 couples 

(around 20% of the European population) 

(Monteiro et al., 1999), Roseate tern 

with more than 1000 couples (60% of the 

European population) and Common tern, 

around 2000 couples (5% of the European 

population) (Rodrigues & Nunes, 2002). 

All these seabird species have a vulnerable 

status, except Roseate tern which 

is in Danger, and the Common tern with a 

Favourable Conservation status 

(Rodrigues & Nunes, 2002). 

Most seabird populations breeding in 

Azores have been suffering dramatic historical 

declines as a consequence of major 

habitat degradation, mainly by human 

activities from the late 15 th century on, 

and the introduction of mammalian 

FIGURE 1. Ilhéu de Vila Franca do Campo with identified habitats, burrows and nests cavities. 

C - Vegetated Sea Cliffs of the Macaronesian Coasts habitat; H - Endemic Macaronesian Heaths 

habitat; F - Forests of iron trees; V – vineyards; + Common tern (CT) nests; * Cory’s shearwater (CS) 

burrows/nests cavities; CS destroyed nest; CS abandoned nest; CS abandoned egg; CT 

destroyed nest; CT abandoned nest; CT abandoned egg; Area of public permitted access.

RODRIGUES ET AL: SEABIRD CONSERVATION AT ILHÉU DE VILA FRANCA 219 

predators (Monteiro et al., 1996b), so they 

tend to breed on isolated islets and sea 

cliffs, free of predators and human disturbance, 

with natural habitats without invasive 

alien species (Monteiro et al., 1996b; 

Ramos et al., 1997; Groz & Pereira, 2005). 

One of these islets is the Ilhéu de Vila 

Franca do Campo (IVFC) (Figure 1), 1.2 

km south of São Miguel island. 

Several studies were published on the 

islet’s biota, describing its general topography 

and biological characteristics, 

mainly about its marine life and coastal 

ecology (Martins, 1978, 1995, 2004; 

Morton, 1990; Britton, 1995; Backeljau et 

al., 1995; Morton et al., 1998) revealing the 

ecological and geological importance of 

the place. 

Some surveys targeting Cory’s shearwater 

and terns were made in the Azores 

(del Nevo et al., 1993; Bolton, 2001 and 

Monteiro et al., unpublished report) but 

none of them went to the islet. Monteiro 

et al. (1999) estimated 0 to 10 Bandrumped 

storm-petrel couples breeding on 

the islet. 

Due to the unique importance of the 

IVFC as an agglomeration of various 

micro-ecosystems, the Azorean 

Government established it as a Nature 

Reserve in 1983, as provided by the 

Regional Legislative Decree nº 3/83/A, of 

March 3 rd , and some rules were implemented 

with the intention to preserve 

and protect the islet; the public access was 

restricted to the lagoon, the marine area 

of the reserve was extended to 30 m deep, 

and fishing and any underwater activity 

were forbidden. In spite of these protection 

measures, the flora and fauna in the 

islet and its lagoon have suffered a significant 

negative impact caused by the flood 

of people mostly during the summer 

(Morton et al., 1998). The islet is popular 

for recreation and between May and 

September a boat brings in about 400 persons 

a day. 

This study was developed to identify 

the breeding species of seabirds on IVFC, 

giving special emphasis to habitat characterization 

and threats, thus contributing 

to the conservation and protection of 

the islet’s natural habitats and their 

seabird populations. 


The study was carried out on the 

IVFC (N37º42.30’, W25º26.52’), during 

July 2006, in the course of the 3 rd international 

Workshop on Malacology and 

Marine Biology in Vila Franca do 

Campo, São Miguel, Azores (Figure 1). 

The islet is a drowned volcanic crater 

with a surface area of 61.640 m 2 , accessed 

by a narrow channel, but with several 

fissures also connecting the lagoon with 

the sea outside. The main length of the 

islet is 420 m from east to west, reaches 

an altitude of 62 meters, and is divided 

into two portions: the Big islet, that constitutes 

almost all the islet’s area and the 

Small islet on the northeast side. There 

are also several rocks of different sizes, 

outstanding the Farilhão with 32.5 m, 

and Baixa da Cozinha with 19.4 m. On 

the islet there is an internal lagoon connected 

to the open sea trough chaps and 

underwater tunnels. Inside the lagoon 

there is a small pier where people enter 

the islet (Martins, 2004). 

The habitat characterisation of the 

islet was made in loco following the 

Interpretation Manual of European 

Union Habitats (European Habitats 

Committee, 1991). 

Three different methods were conducted 

to identify the seabird species of 

the islet, and to estimate species abundance 

and number of breeding pairs: 

1. Flush counts 

Six boat rides, of 15 minutes each, 

were undertaken for flush counts of 

seabirds and to observe the external

220 AÇOREANA 

2009, Sup. 6: 217-225 

coast zone of the islet. The method consisted 

on counting the number of individuals 

visible from the boat (eye view), 

at different hours of the day, three rides 

in the morning and three on the afternoon. 

Species were identified and all the 

individuals standing on the islet or flying 

over the sea were counted. 

2. Ground searches 

Two ground searches were made for 

occupied nests of terns and 

Porcellariiformes (Figure 1). Signs of 

occupation included the presence of an 

adult, eggs or chicks in a visible nest 

chamber. In sites where a nest chamber 

was not visible, a burrow was considered 

to be occupied if it was of sufficient size 

to accommodate a bird or if there was 

one or more evidences of occupation 

(e.g. faeces; shed breast feathers, excavated 

soil or absence of obstructing vegetation 

or spider webs in the burrow’s 

entrance). 

3. Raft counts 

During the breeding season, Cory’s 

shearwater characteristically form flocks 

(called “rafts”) on the sea, with the number 

of individuals increasing by the end 

of the day, waiting for nightfall to fly up 

to the breeding colony (Mallet & 

Coghlan, 1964). According to Richdale 

(1963) and Skira (1991) over half of the 

adult shearwaters at a colony are 

expected to be non-breeders. 

Raft counts were conducted for 

Cory’s shearwater from two places, one 

on the top of the south side of the islet, 

and another from Ponta de São Pedro 

(N37º42.39’, W25º26.41’) on the coast of 

São Miguel island and in front of the 

islet; raft counts were made by the end of 

the day, starting exactly at 08:00 pm, for 

45 minutes long, using binoculars (10 X 

50). 

All the methods applied in this study 

involved two replicates from two different 

observers. 

RESULTS 

Habitats 

Two types of natural habitats were 

identified on the IVFC (Figure 1): i) 

Vegetated Sea Cliffs of the Macaronesian 

Coasts habitat, dominated by the endemic 

fescue (Festuca petraea), with associated 

plants such as rush (Juncus acutus), 

wild carrot (Daucus carota), seaside goldenrod 

(Solidago sempervirens) and rock 

samphire (Crithmum maritimum); ii) 

Endemic Macaronesian Heaths habitat 

dominated by the endemic green heather 

(Erica azorica), with associated plants 

such as laurel (Laurus azorica), Myrica 

faya, Cyrtomium falcatum, Holcus rigidus 

and Euphorbia azorica. This last habitat 

presents plenty of exotic plants such as 

the giant reed (Arundo donax), used in the 

past for protective barriers of the vineyards, 

tamarisks (Tamarix gallica), brambles 

(Rubus ulmifolius) and australian pittosporum 

(Pittosporum undulatum). 

On the higher southern and western 

inner slopes of the Big islet there are two 

small “forests” of iron trees (Metrosideros 

tomentosa) and vineyards (Vitis labrusca), 

although they are no longer cultivated. 

Seabirds 

There was evidence of two species of 

seabirds breading on the islet (Figure 1), 

Cory’s shearwater and Common tern, 

but three more species were registered, 

Little shearwater, Band-rumped stormpetrel 

and Roseate tern, although at present 

without any breeding evidence. 

Table 1 shows avifauna abundance 

determined through the different methods 

used. 

Cory’s shearwater breeds on Big islet 

with Vegetated Sea Cliffs of the 

Macaronesian Coasts habitat where 34 

burrows and nests cavities were found, 

and on slopes of Endemic Macaronesian 

Heaths habitat that exhibited 10 burrows


TABLE 1. Seabird abundance from flush and raft counts and number of couples from occupied 

nests/burrows on the Ilhéu de Vila Franca do Campo. SD = Standard Deviation. 

Species 

Flush counts 

(individuals) 

Mean ± SD 

Ground searches 

(couples) 

Raft counts 

(individuals) 

Mean ± SD 

Cory’s shearwater 154 ± 22.6 44 395 ± 21.2 

Common tern 325 ± 35.4 80 - 

Roseate tern 2 - - 

Little shearwater 1 - - 

Band-rumped storm-petrel 1 - - 

and nests cavities. The three nests found 

on Small islet were destroyed or abandoned. 

Common terns breed on two external 

rocks of the islet, Baixa da Cozinha (10 

nests) and Farilhão (19 nests), and on the 

external coast of the Small islet with 


Coasts habitat (41 nests). 

Occasional observations on the populations 

of Cory’s shearwater and 

Common terns allowed identification of 

chicks being reared, incubating parents, 

destroyed and abandoned nests and 

abandoned eggs (Table 2). It was not possible 

to identify the exact number of eggs 

laid per couple of Common terns. 

Evidences of human disturbance were 

found near potential seabird nests, 

including recreation, wastes, and vandalism 

such as broken eggs and burrows 

destruction. There were an equivalent 

percentage of abandoned nests in both 

populations (11%) and about 18% of 

Cory’s shearwater nests and 5% of 

Common tern nests were destroyed. 

Nearly 7% of Cory’s shearwater eggs and 

1% of Common tern eggs were abandoned 

by the progenitors. 

DISCUSSION 

The Vegetated Sea Cliffs of the 

Macaronesian Coasts habitat of the IVFC 

are well preserved probably because they 

occur on the external rocks of the islet and 

on inaccessible cliffs, but also because 

they are highly influenced by salt-water 

spray where exotic plants cannot grow. 

According to Sjögren (1973) the association 

Festucetum petreae is characteristic of 

Azores coastal habitats, occurring mainly 

in the sea cliffs. The characteristic species 

of this association is the common endemic 

Festuca petraea, which usually develops 

coastal prairies. Other species are found 

in this association, such as the common 

Solidago sempervirens and Crithmum maritimum, 

the less common Tolpis succulenta 

and even rarest endemic plants like 

Azorina vidalii and Myosotis maritima. 

The Endemic Macaronesian Heaths 

habitat was much degraded with exotic 

plants such as Arundo donax, Lantana 

TABLE 2. Number of observations on Cory’s shearwater and Common tern nests on Ilhéu de Vila 


Species 

Used 

nests 

Destroyed 

nests 

Abandoned 

nests 

Abandoned 

eggs 

Incubating 

couples 

Rearing 

chicks 

Cory’s shearwater 44 8 5 3 9 6 

Common tern 80 4 9 1 37 11

222 AÇOREANA 

2009, Sup. 6: 217-225 

camara and Pittosporum undulatum. 

According to the Convention on 

Biological Diversity, invasive alien species 

which introduction and/or spread threatens 

biological diversity are now considered 

the second cause of biodiversity loss 

at a global level, after direct habitat 

destruction (Shine et al., 2000; Groz & 

Pereira, 2005). 

The methods used in this study to 

estimate species abundance and number 

of breeding pairs were effective because 

they complemented each other giving a 

broader view of the colonies. These 

results confirmed nesting of two endangered 

species and listed three other possible 

breeders revealing the importance of 

the IVFC for the protection and conservation 

of seabirds. 

Cory’s shearwater breeds generally in 

the inaccessible sea cliffs of the islet and 

less in slopes with Macaronesian vegetation. 

The differences between the numbers 

of burrows and nests found in the 


Coasts habitat and in the Endemic 

Macaronesian Heaths habitat are probably 

related to human disturbance and 

degradation of this last habitat with invasive 

alien species that are a threat to 

seabird populations (Groz & Pereira, 

2005). Adults arrive in colonies by late- 

February, lay eggs from late-May to early- 

June and hatching is relatively synchronous, 

most chicks hatching between 18 

and 31 July (Granadeiro, 1991). Young 

birds fledge from late-October to early- 

November (Monteiro et al., 1996b). They 

lay only one egg per year and do not 

replace it if it is damaged or lost soon 

after laying, so there is an ecological significance 

when a couple looses their egg, 

all the dispended energy is lost and it is 

one less breading couple. 

Common terns breed in inaccessible 

rocks with Vegetated Sea Cliffs of the 

Macaronesian Coasts habitat, where 

human disturbance is almost absent, and 

where they can avoid possible predators. 

Adults arrive in colonies by early-April, 

egg laying occurs from early-May to mid- 

June and they stay in colonies until late- 

September (Monteiro et al., 1996b). A 

clutch of 2-4 (usually 3) eggs is laid. One 

brood per season is typical but re-nesting 

is common when the first nest is 

destroyed (Peterson, 1988). This study 

indicates that the colony on IVFC represents 

at least 18% of the Azorean total population. 

Although being a protected species, 

populations of these seabirds are becoming 

smaller than in the past, they have been 

chased and hunted by humans, and suffered 

with predation from introduced 

mammals and from deforestation 

(Monteiro et al., 1996b). 

The suite of terns and shearwaters is of 

major international conservation 

importance (Rodrigues & Nunes, 2002). 

Their breeding distributions in the Atlantic 

Ocean are concentrated in Europe and 

most species have small world populations 

(Monteiro et al., 1999) being classified as 

globally threatened species (Collar et al., 

1994). The Azorean population of Cory’s 

shearwater decreased around 50% between 

1996 and 2001 (Rodrigues & Nunes, 2002), 

and a similar situation was observed for 

Common tern populations, estimated 

around 4000 breeding couples for 1992 (del 

Nevo et al. 1993), against 2000 breeding 

couples for year 2000 (Rodrigues & Nunes, 

2002), probably due to the impact of continuous 

lost of their habitats. 

People’s access to nests and burrows 

represents an important threat over 

seabirds in the islet, and several impacts 

from recreation and vandalism, are leading 

to their distraction from normal activities; 

parents spend less time tending young 

birds or eggs, flying away from nests, leaving 

eggs or chicks vulnerable to predators 

and amenity, nests are destroyed and


seabirds entirely abandon the colonies. 

These impacts can affect Azorean seabirds, 

particularly Common terns, since the 

colony of IVFC represents one of the 

largest in the Azores. 

Due to the importance of the Azores 

archipelago for seabirds in Europe, it is 

fundamental to protect every natural habitat 

and to conserve the few areas where 

they breed, usually steep scarps and islets 

(Monteiro et al, 1996b, 1999; Ramos et al., 

1997). 

In conclusion, results from the present 

work indicate that IVFC plays an important 

role on the conservation of Cory’s 

shearwater and Common tern populations, 

since hundreds of couples of these two 

species breed on the islet. But this importance 

can be threatened by the continuous 

degradation of their habitats. 

Seabirds become mature at a late age, 

experience low annual fecundity, often 

refrain from breeding, and enjoy annual 

adult survival rates as high as 98%. This 

suite of life history characteristics limits the 

capacity for seabird populations to recover 

quickly from major perturbations, and presents 

important conservation challenges 

(Russell, 1999). 

An urgent management plan for IVFC 

is necessary, in order to conserve the natural 

habitats of the islet and protect their 

seabird populations. 

According to Groz & Pereira (2005), 

islets habitats are expected to respond 

rapidly to habitat restoration, so it is urgent 

to implement an habitat restoration program 

and a major efficient islet control to 

people access, in order to improve seabirds 

breeding conditions on the islet. A welldesigned 

legal and institutional framework 

is essential to provide a basis for effective 

eradication and control measures of alien 

plant species (Shine et al., 2000), control of 

soil erosion and multiplication and reintroduction 

of native flora. A mark-recapture 

analysis is useful for tracking demographic 

changes in a population over time 

(i.e., assessing population size, adult survival, 

and juvenile recruitment) (Brichetti et 

al., 2000) and could be used to evaluate the 

effect of the habitat restoration programme. 

Moreover, seabirds are believed to constitute 

useful samplers of the marine environmental 

since they have been recognized 

as potentially useful and economical indicators 

of the status of marine environment 

and, in particular, the status of commercially 

important prey stocks (Furness & 

Greenwood, 1993). 


We are grateful to Professor António M. 

de Frias Martins for the opportunity to 

undertake this research. 

This project was supported by the 

Third International Workshop of 

Malacology and Marine Biology, Vila 

Franca do Campo, São Miguel, Azores, July 

2006. 

The experiments performed for the present 

study comply with the laws of the 

country in which they were performed. 


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Remembering 

Joseph C. Britton (1942-2006) 

with Brian Morton and Fred Wells at his side, at the 2nd International Workshop of Malacology 

and Marine Biology, Vila Franca do Campo (1991) 

Joseph Cecil Britton passed away on 29 th November 2006. His life intersected the 

Azores in 1991, when he attended the 2nd International Workshop of Malacology and 

Marine Biology, and stayed linked to these islands from then on. I first met Joe at one 

of the Hong Kong workshops in 1989. Always polite and respectful, a true companion 

in science, beer and joie de vivre, Joe’s good humour made him a valued member of the 

team of scientists he joined here in the mutual search for knowledge. He knew that he 

was path finding, but this only increased his determination to help Azorean scientists 

and students understand their rich marine cultural heritage. His accumulated wealth of 

information about marine ecology flowed easily through his elegant discourse (albeit 

with a true Texan twang) and his calm devotion to work fostered in everyone a sense of 

duty. One of Joe Britton’s greatest contributions to the Azores, and one in which I was 

privileged to be part of, was the book Coastal Ecology of the Açores, one of a series of 

monographs created by Brian Morton, about the shores of Hong Kong and of Texas. 

Other works about the ecology and conservation of the Azores followed and, at the time 

of his death, he was compiling a huge list of publications related to the marine biology 

of the Azores. The completion of this task would form a lasting memorial to him and 

his work. Joe Britton was a member of the Editorial Board of Açoreana. A true friend, 

Joe will be missed, but his memory will be with us forever.

AÇOREANA é a revista da Sociedade Afonso Chaves 

- Associação de Estudos Açoreanos e visa publicar 

trabalhos devotados principalmente às diversas áreas da 

história natural dos Açores. AÇOREANA está indexada em 

BYOSIS, é enviada para Zoological Record e distribuída em 

regime de troca por bibliotecas de vários países. 

Os manuscritos, em Português, Francês ou Inglês, 

incluirão um RESUMO e tradução deste numa daquelas 

línguas. O formato conformar-se-á com o de números 

posteriores a 2000. Todas as partes do manuscrito (texto, 

referências, tabelas, legendas) serão dactilografadas a dois 

espaços. Nomes de géneros e espécies serão sublinhados; 

todas as outras indicações serão deixadas ao critério do 

editor. As ilustrações deverão ser executadas de forma a 

permitir uma utilização eficiente do espaço útil (página, 

125x180 mm; coluna, 62x180 mm); letras e números 

deverão permanecer perfeitamente legíveis após a redução. 

As referências no texto seguirão uma das seguintes formas: 

‘Dance (1986) descreveu …‘ ou ‘ … (Morton, 1965) …‘ ou ‘… 

(Nobre, 1924, 1930; Martins, 1989a, b; Hawkins et al., 1990; 

Martins & Ripken, 1998;).’ A bibliografia é listada 

alfabeticamente e os nomes dos autores repetidos sempre 

que necessário; os nomes das revistas são apresentados por 

extenso. A listagem bibliográfica (BIBLIOGRAFIA 

CITADA) seguirá o formato dos números posteriores a 

2000, conforme se exemplifica: 

HOUBRICK, R.S., 1990. Anatomy, reproductive biology 

and systematic position of Fossarus ambiguus (Linné) 

(Fossarinae: Planaxidae; Prosobranchia). In: 

MARTINS, A.M.F. (ed.), The Marine Fauna and Flora of 

the Azores (Proceedings of the First International Workshop 

of Malacology, São Miguel, 1988). Açoreana, Supplement 

[2]: 59-73. 

LEAL, J.H., & P. BOUCHET, 1991. Distribution patterns and 

dispersal of prosobranch gastropods along a seamount 

chain in the Atlantic Ocean. Journal of the Marine 

Biological Association of the United Kingdom, 71 (1): 11- 

25. 

MORELET, A., 1860. Notice sur l’Histoire Naturelle des Açores 

suivie d’une description des Mollusques terrestres de cet 

Archipel, 216 pp. J.-B. Baillière et Fils, Paris. 

Tabelas e ilustrações virão após BIBLIOGRAFIA 

CITADA, constando no manuscrito o lugar apropriado para 

a sua integração; as ilustrações devem ser numeradas em 

série única; as legendas das figuras serão apresentadas 

separadamente após as ilustrações. 

Aceitam-se notas curtas que não deverão exceder três 

páginas dactilografadas a dois espaços. Normalmente não 

comportarão sumário ou subtítulos. 

Para salvaguardar a interpretação na 

eventualidade de desformatação do material digital, uma 

cópia impressa completa com tabelas, legendas e ilustrações 

será submetida ao editor, juntamente com uma cópia em 

CD ou via e.mail. 

SEPARATAS. O primeiro autor receberá 50 separatas 

grátis; cópias adicionais a preço de custo poderão ser 

requisitadas no acto da devolução das provas. 

CORRESPONDÊNCIA. Enviar para o editor, Prof. 

António M. de Frias Martins, Sociedade Afonso Chaves - 

Associação de Estudos Açoreanos, Apartado 258, 9501-903 

Ponta Delgada, São Miguel, Açores, Portugal. E.mail: 

afonsochaves.sac@gmail.com 

AÇOREANA is the journal of the Sociedade Afonso 

Chaves - Associação de Estudos Açoreanos and aims at the 

publication of works devoted mainly to the various areas 

of the natural history of the Azores. AÇOREANA is 

indexed by BIOSIS, sent to Zoological Record, and distributed 

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References are listed alphabetically, the authors’ names 

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listing should follow the format of the issues after 2000, 

according to the examples: 

HOUBRICK, R.S., 1990. Anatomy, reproductive biology 

and systematic position of Fossarus ambiguus (Linné) 

(Fossarinae: Planaxidae; Prosobranchia). In: 

MARTINS, A.M.F. (ed.), The Marine Fauna and Flora 

of the Azores (Proceedings of the First International 

Workshop of Malacology, São Miguel, 1988). Açoreana, 


LEAL, J.H., & P. BOUCHET, 1991. Distribution patterns 

and dispersal of prosobranch gastropods along a 

seamount chain in the Atlantic Ocean. Journal of the 

Marine Biological Association of the United Kingdom, 71 

(1): 11-25. 

MORELET, A., 1860. Notice sur l’Histoire Naturelle des 

Açores suivie d’une description des Mollusques terrestres 

de cet Archipel, 216 pp. J.-B. Baillière et Fils, Paris. 

Tables and illustrations should come after LITERA- 

TURE CITED, but there should be in the manuscript an 

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REPRINTS. The first author receives 50 reprints free 

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CORRESPONDENCE. Manuscripts and 

correspondence related to the journal should be 

addressed to the editor, Prof. António M. de Frias 

Martins, Sociedade Afonso Chaves — Associação de 

Estudos Açoreanos, Apartado 258, 9501-903 Ponta 

Delgada, São Miguel, Açores, Portugal. E.mail: 

afonsochaves.sac@gmail.com

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