THE CENOZOIC BRACHIOPOD TEREBRATULA 



91 



Type specimens. The holotype, the specimen figured by Colonna 

 (1616c). is lost. We here nominate as neotype (Fig. 6a-c). an entire 

 specimen from the Calcarenite di Gravina Formation near a locality 

 collected by Colonna, in the collection of the Natural History Mu- 

 seum. London (BMNH BG152), collected 22 March 1998 by M. 

 Caldara & O. Simone. Dimensions of the neotype are: length 55.4 

 mm, width 43 mm, thickness 30mm. A well-preserved complete 

 topotype with the dimensions: length 50.8 mm, width 42 mm, 

 thickness 28 mm, is also figured (Figs 7a-c). Two further topotypes. 

 one with a complete loop (Figs 9a. b), and a dorsal valve with a 

 strongly-M-shaped anterior commissure (Figs 8a, b) are also 

 illustrated. 



Material. The brachiopods from the Calcarenite di Gravina For- 

 mation are frequently broken across mid-valve, or are separated 

 valves. Most specimens are infilled with coarse, white, hard, moder- 

 ately cemented calcarenite. and few have a complete anterior 

 commissure. A number of topotypes ( BG 1 53- 1 6 1 . BG 1 94- 1 96) are 

 held in The Natural History Museum. London. 



Type locality. The type locality is an outcrop of Calcarenite di 

 Gravina Formation, of Pliocene age, on the east side of a small dry 

 valley, about 200m north of the church of Madonna dei Miracoli 

 (41°13'52"N; 16°16'00"E), about 2km west of Andria, Puglia, Italy. 



Age. Late Miocene (Tortonian), Pliocene. 



Distribution. Puglia, Calabria. Tuscany. Emilia (Piacentino). 

 Abruzzi, Spain (Alicante). 



Description. Shell of medium to large size, biconvex, anterior 

 commissure rectimarginate, uniplicate to sulciplicate, two broad 

 plicae may be developed; beak short, massive, suberect; foramen 

 large, mesothyrid to permesothyrid; symphytium narrow, partly 

 concealed. Pedicle collar short; hinge teeth with moderately swollen 

 bases. Cardinal process semielliptical, moderately protuberant; outer 

 hinge plates narrow; no inner hinge plates; crural bases fused to 

 socket ridges to form a deep V-shaped trough, crural processes long; 

 loop broadly triangular, about 0.3 valve length, transverse band 

 narrow, forming medially flattened low arch. 



Terebratula ampulla (Brocchi, 1814) 



1814 Anomia ampulla Brocchi: 466 



Age. Pliocene. 



Distribution. Italy (Emilia (Piacentino), Tuscany, from Brocchi's 

 list), but not Calabria (see Seguenza, 1871). 



Comments. A medium-sized, strongly bisulcate species. 



Terebratula scillae Seguenza, 1871 



1 87 1 Terebratula scillae Seguenza: 39 



Age. Early Pleistocene. 



Distribution. Calabria, Puglia, Sicily. 



Comments. The largest species of Terebratula (up to 95mm in 

 length), which formed extensive shellbeds in the Early Pleistocene. 



ECOLOGY AND EXTINCTION OF 

 TEREBRATULA ^^ 



Terebratula grandis Blumenbach, 1803, now included in the genus 

 Pliothyrina (see Cooper, 1983), may be the ancestor of Terebratula 



sensu stricto. This large species from the Oligocene of Germany 

 needs further study. Pliothyrina appears to have been widespread in 

 northern Europe and England (see collections in the Natural History 

 Museum, London), whereas Terebratula was abundant in the Med- 

 iterranean region. Both groups became extinct in the Plio-Pleistocene. 



Terebratula was widely distributed in the Mediterranean region 

 from the Miocene until the early Pleistocene. It lived in circalittoral 

 environments on muddy, biodetrital seafloors. attached to substrates 

 which included bivalves and other brachiopods. During the 

 Messinian, when the Mediterranean basin became too saline to 

 support normal marine life, the brachiopod fauna disappeared from 

 the region. With flooding of the Atlantic sea into the Mediterranean 

 basin, the Mediterranean was recolonised by a brachiopod biota with 

 Atlantic affinities (Logan, 1979). This does not account for the 

 reappearance of Terebratula, which may have survived in refugia. 



In the Pliocene, Terebratula was an important component of a 

 widespread brachiopod fauna which mcXndtd Aphelesia and Megerlia 

 (Gaetani, 1986;TaddeiRuggiero, 1994, 1995). The youngest records 

 are of very large Terebratula scillae which formed vast shellbeds in 

 the Early Pleistocene (Taddei Ruggiero, 1986, 1994). The extinction 

 of Terebratula by the Middle Pleistocene appears to be related to the 

 reduction in sea temperatures as Pleistocene cooling proceeded. 



Acknowledgements. Professor Maurizio Gaetani gave initial introduc- 

 tions to geologists in Italy. We thank Eileen Brunton and Sarah Long at the 

 The Natural History Museum, London, for help with bibliography and for 

 photography of the brachiopods. We also thank Professor John Barsby, 

 University of Otago. for translations of Colonna's work, Bill Lee and Doug 

 Campbell for improvements to the manuscript. Andrew Grebneff for speci- 

 men preparation and Ewan Fordyce for photography. The Royal Society of 

 New Zealand contributed towards travel to the British Museum (Natural 

 History) in 1991, and funding from the University of Otago Division of 

 Sciences, and the Department of Scienze della Terra of the University of 

 Napoli. is gratefully acknowledged. Professor Giovanni Aliotta, II University 

 of Napoli, gave us new information on Colonna, Professor Roberto Taddei, 

 University of Napoli. translated some passages oi Purpura, and the Library of 

 the University and the Department of Biologia Vegetale of the University of 

 Napoli 'Federico 11' allowed photographic reproduction of Colonna's Purpura. 



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 Brunton, C.H.C. & Cocks, L.R.M. 1967. Tcrehratulina d'Orbigny, 1 847 (Brachiopoda): 



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