86 



D.E. LEE, C.H.C. BRUNTON, E. TADDEI RUGGIERO, M. CALDERA AND O. SIMONE 



descript. 23. icon. 22. Altera imbricata. 24. altera margine imdosa. 

 ibid. & 25' . The two figures on the top of page 22 are joined under 

 the same caption 'Concha anomia vertice rostrato' . 



The figure in the centre of page 22, described on pages 24-25 as 

 'Concha altera Anomia striata [trilobos] rarior. I. Cap. XV.', is 

 considered as another shell and its numeration starts again from T". 



Much of the confusion over the identity of the two figures on the 

 top of page 22 stems from the fact that Colonna described the shell 

 placed on the upper right first, and on a following page discussed the 

 shell portrayed on the upper left, using a different practice from that 

 which became well established in the following centuries. The 

 confusion increased further when later authors assigned to these 

 figures two numbers that Colonna had never employed. Thus 

 Linnaeus (1758), following modern convention, designated the 

 strongly folded specimen on the upper left as f. 1 (ie., figure 1 ) and 

 that on the upper right as f 2 (ie., figure 2). Brocchi (1814), and the 

 present authors, unfil lately, have done the same. 



The problem of deducing which shell description accompanied 

 which illustration was compounded in a second edition of Colonna's 

 work, with the associated text from the earlier edition subdivided 

 into numbered paragraphs, which was reprinted posthumously in 

 1675 by J.D. Major. In this edition, the redrawn figure of the strongly 

 folded specimen from the upper left of page 22 in the 1616 edition 

 (i.e. Anomia terebratula of Linnaeus) appeared on page 32 within the 

 text referring to the Andria locality. Similarly, in the Dictionarium 

 this figure was noted by Major as that described in Cap. 12, of 

 Colonna (1616c: 23). 



A comparison of the three figures (Fig. 2 herein) shows that the 

 Klein figure listed by Linnaeus (Klein, 1753; Tab. Nostra XI. n. 74), 

 was redrawn from this later edition (Major, 1675 ) (Fab. Columnae de 

 Purp. Cap. XII $.3. pag. 32 [Klein 1753, p. 171-2]). 



Dollfus & Dautzenberg (1932) correctly interpreted the refer- 

 ences to Major's figures, but added to the confusion, by giving the 

 same citation for both the upper right and upper left figures: ie. 'p. 22, 

 fig.l (2e)'. These authors also assumed that the caption 'Altera' 

 pertained to the upper left figure, whereas it undoubtedly refers to the 

 figure in the center ('Altera [trilobos]'). because the captions in all 

 seven figures in Colonna's Purpura (pp. 13,16, 20, 22, 27, 30, 33) are 

 placed over, and never under, the relevant figures. 



DISCUSSION AND COMMENTS 



This discovery, as our manuscript was almost ready for submission 

 and after the ICZN had approved the selection of a neotype for 

 Terebratula terebratula (Linnaeus), raises some issues which need 

 further discussion. If indeed the provenance of the specimen selected 

 by Linnaeus as the type oi Anomia terebratula is unknown and the 

 specimen is lost, then the basis for the species, and consequently the 

 genus, family and superfamily would remain uncertain. 



However, since Lee & Brunton (1998) have already nominated a 

 neotype from Colonna's locality near Andria, the neotype locality 

 now becomes the type locality for the species, regardless of the 

 locality of the original Colonna brachiopod (International Code of 

 Zoological Nomenclature Article 75f). 



Is it possible then to determine the provenance of the original 

 specimen of Colonna selected by Linnaeus as the type of Anomia 

 terebratula! 



Firstly, it is obvious that Colonna's caption: 'Concha anomia 

 vertice rostrato' applies to both of the brachiopods figured on the 

 upper right and upper left of page 22, the first of which (that on the 



upper right) certainly came from Pliocene strata near Andria, and 

 that Colonna himself regarded these specimens as similar to one 

 another. 



Secondly, six original copies of Colonna (1616c) that we have 

 consulted have the numbers:'r, "4' and T' written beside the three 

 brachiopods figured on page 22. These numbers are not included in 

 Major's edition. Linnaeus ( 1758, 1767) and Brocchi (1814), if aware 

 of these numbers, used different, modern numbering (i.e. f. 1 for 

 Colonna's specimen 4 on the upper left), and Major (1675) and Lee 

 & Brunton (1998) considered that Colonna's description on page 23 

 referred to the figure on the upper left. 



Thirdly, even if the specimen of Colonna's (1616c; 22) upper left 

 figure is from an unknown locality, it was filled with white, loose 

 sediment (Colonna, 1 6 1 6c: 24), and it is possible that it was collected 

 from the same Pliocene calcarenites at Andria. 



From a close examination of the Colonna woodcut, it seems likely 

 that his specimen was somewhat deformed. It has a large, open fora- 

 men, and two strongly developed plicae/folds on the dorsal valve that 

 begin at an early stage of growth, and would have resulted in a strong 

 sulciplication(;;;a/-g;HCM«(yo5a) of the anteriorcommissure. The artist 

 (?Colonna himself) may have exaggerated the depth of the folds, 

 although the depiction of the other brachiopods on the plate seem to be 

 faithful to reality. No undeformed specimens collected by the authors 

 have folds as strongly developed as those depicted in the woodcut. 



From the many specimens of Terebratula collected by the authors 

 from Colonna's Andria locality and from elsewhere in Italy, it is 

 apparent that the brachiopods in any fossil assemblage/population 

 vary considerably in the degree of folding and may be rectimarginate 

 to biplicate or sulciplicate. Thus, both specimens labelled by Colonna 

 as Concha anomia vertice rostrato are species of Terebratula (sensu 

 lato). and given the wide variation in populations of Neogene 

 Terebratula, might be conspecific. Certainly, the specimens of 

 Terebratula terebratula figured by us in this paper (Figs 6-9) fall 

 somewhere in the middle of the two short-looped brachiopods 

 illustrated in Colonna's woodcut. The specimen of Pliocene 

 Terebratula terebratula from Monte Mario selected and figured by 

 Buckman (1907) and illustrated in the 1965 Treatise, is crushed and 

 deformed in a similar manner. In the Natural History Museum, 

 London, there are several collections of Terebratula from this same 

 horizon at Monte Mario, near Rome. These specimens, which are 

 almost certainly conspecific, vary from small rectimarginate (juve- 

 nile) specimens (labelled T. depressa) to large rectimarginate 

 individuals (named T. grandis), to examples with moderate 

 sulciplication (labelled T. ampulla or T terebratula). 



Thus, our selection of a neotype from a locality described and 

 collected by Colonna closely follows the recommendations of the 

 Code. 



It should be noted that Muir- Wood's ( 1955: fig. 2) caption for her 

 reproduction of the original Colonna figure is misleading. The caption 

 reads 'Reproduction of early drawings of Brachiopods figured as 

 'Concha anomia', and taken from Fabio Colonna's first edition ofde 

 Purpura, 1 6 1 6, p. 22. The first figure is of a specimen from Mte. Mario, 

 near Rome, and is probably of Tertiary (?Pliocene) age; the other two 

 figures may represent Jurassic forms'. However, as shown here, the 

 first figure is from an unknown locality, while the second specimen is 

 of Pliocene age and came from near Andria, not Rome. 



TYPE LOCALITY OF TEREBRATULA 

 TEREBRATULA 



Colonna ( 1616c: 23) described the locality from which he collected 

 his specimen of Terebratula shown on the top right of his figure 



