Bull. nat. Hist. Miis. Land. (Geol.) 57(2): 83-93 



//^2^ 



00 



( ^-l 



y 



Issued 29 November 2001 



The Cenozoic Brachiopod Terebratula: its type 

 species, neotype, and other included species 



THE NATURAL 

 tflSTORY MUSEUM 



DAPHNE E. LEE 



Department of Geology. University of Otago. P.O. Bo.x 56. Diinedin. New Zealand 



C.H.C. BRUNTON 



Department of Palaeontology. The Natural History Museum. Cromwell Road, London SW7 5BD 



EMMA TADDEI RUGGIERO 



Dipartimento di Scienze della Terra della Universita' di Napoli 'Federico IT, Largo San Marcellino 10. 80138 

 Napoli. Italy 



MASSIMO CALDARA & ORONZO SIMONE 



Dipartimento di Geologia e Geofi'iica. Campus Universitario, Via Orahona 4, 70125 Bari, Italy 



S OCT 2001 



PB£ScWTHD 

 QEN£RAL UBRARY 



Synopsis. Terebratula terebratula (Linnaeus. 1758) has a long and complex history. The specimen now recognised as the type 

 was first illustrated by Colonna in 1616, and the first use of 'Terebratula' is attributed to Lhwyd, 1699. Colonna's specimen was 

 refigured by Klein, 1753, and the species /Inom/a terebratula was described by Linnaeus. 1 758. with reference to the Colonna and 

 Klein illustrations. The genus Terebratula was proposed by Mijller in 1776. and Anomia terebratula Linnaeus designated as the 

 type species by Lamarck in 1799. although it was not an originally included species. In spite of this history, the type of the genus 

 was never formally ratified, the whereabouts of the type specimen was unknown, and the age and exact position of the type locality 

 was uncertain. This paper summarises the history of Terebratula terebratula (Linnaeus) from \616. Anomia terebratula Linnaeus 

 is now accepted as the type species of Terebratula (ICZN ruling, 2000). We have collected new material from a locality near 

 Andria, Italy, from which Colonna collected specimens of Terebratula, and selected a neotype from the Calcarenite di Gravina 

 Formation which is Pliocene in age. Two existing species, Terebratula sinuosa (Brocchi) and T calabra Seguenza, are placed in 

 synonymy with T. terebratula. Three species are currently recognised in Terebratula. ranging in age from Miocene to Early 

 Pleistocene when the genus became extinct, probably because of ocean cooling in the Mediterranean region. 



INTRODUCTION 



The brachiopod genus Terebratula has a long and complex geologi- 

 cal and nomenclatural history. The nominal genus Terebratula was 

 proposed by Miiller in 1776, and as pointed out by Muir-Wood 

 (1955), it 'is the first valid post-Linnean brachiopod genus". 

 Terebratula terebratula (Linnaeus) is the name-bearer for the Order 

 Terebratulida which encompasses loop-bearing brachiopods of 

 Devonian - Recent age and includes most brachiopods living today. 



The name Terebratula has been widely used for over 200 years: 

 more than 850 specific names were applied to the genus between 

 1800-1850 alone (Sherborn, 1932). In spite of the ubiquity of the 

 name, the genus and species on which it was originally based have, 

 until recently, been little studied and many basic questions about 

 Terebratula terebratula remained unanswered. As pointed out nearly 

 a century ago (Buckman, 1907), not only was the type species of the 

 genus unconfirmed, but its type locality and age were uncertain. 



The present study had several aims. The first objective was to 

 ratify the type of the genus Terebratula for inclusion in the revised 

 Brachiopod Treatise, following the recommendation made by Muir- 

 Wood in the 1965 Treatise volume on the Order Terebratulida. 

 Secondly, we wished to summarise the complex nomenclatural 

 history of the genus and species, Terebratula terebratula. since it was 

 first illustrated and described in 1 6 1 6. The third aim was to locate the 

 type specimen and/or type locality of T. terebratula, or, if this proved 

 impossible, select a neotype to act as namebearer for the order. 

 Finally, we wished to describe the age and relationships of species 

 currently included in Terebratula. 



The first objective was achieved with an application to the Inter- 

 national Commission on Zoological Nomenclature to validate the 

 selection of Anomia terebratula Linnaeus as type-species of the 



genus Terebratula as designated by Lamarck in 1 799 (Lee & Brunton, 

 1998; Ruling of the Commission, September 2000). The remaining 

 objecfives are achieved in this paper. 



DERIVATION OF THE NAME 



The name Terebratula was first used in print by Lhwyd ( 1 699) (Little 

 etal.. 1 973 ), and is the oldest generic name in the Phylum Brachiopoda 

 in current use. Terebratula is "so-called from the perforated beak of 

 the ventral valve' (Little etal., 1973: 2265), and is a quasi-diminutive 

 of Latin terebratus, the past participle of terebrare 'to bore'. The 

 brachiopods listed as Terebratula in Lhwyd's catalogue of shells in 

 the collections of the Ashmolean Museum, Oxford, were 'poorly 

 figured specimens from Witney' (Muir-Wood, 1955: 2). The name 

 came into fairly common use in the 1 8th century, and some of the 

 numerous brachiopods referred to as Terebratula by other pre- 

 Linnean authors were mentioned by Muir-Wood (1955). 



The species name terebratula was first used in a valid binomial by 

 Linnaeus in his description of Anomia terebratula in Systema Natu- 

 rae (1758: 703). Linnaeus gave no illustration, but referred to figures 

 in Colonna (1616c), Lister (1678) and Klein (1753). The Colonna, 

 and Colonna/Klein, illustrations are reproduced in Figs 1, 2. 



THE COLONNA ILLUSTRATION OF 

 TEREBRATULA (FIG. 1) 



Fabio Colonna ( 1567-1650) (Fabius Columna) was bom in Naples 

 and was one of the first natural historians to use copper plates for 

 engraving botanical and zoological figures. He wrote extensively on 



©The Natural History Museum, 2001 



