Sobre la identidad de un canino superior masculino de hominoideo de la Cuenca del Vallès-Penedès figurado por Pickford (2012)-On the identity of a hominoid male upper canine from the Vallès-Penedès Basin figured by Pickford (2012)

Pickford (2012) inadvertently figured—with an incorrect catalogue number—an unpublished male upper canine of a hominoid primate from the Vallès-Penedès Basin (without specifying its locality of provenance), and attributed it to the Late Miocene fossil great ape Hispanopithecus laietanus. In fact, this canine comes from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM), and shows greater similarities with Middle Miocene hominoids than with Hispanopithecus. Here we report the correct catalogue number of this specimen (IPS41714) and provide details on its locality of provenance (ACM/C4-Ap) as well as estimated age (11.9 Ma). On morphologic grounds, we further justify the attribution of this specimen to Dryopithecus fontani, previously recorded by craniodental remains at another ACM locality with the same age, and also correct several other minor errors.


Introduction
In a recently-published paper on the taxonomic identity of the hominoid dental remains from Neuhausen and other Bohnerz localities (Swabian Alb, Germany), Pickford (2012) emphasized the role of Middle and Late Miocene hominoids from the Vallès-Penedès Basin for understanding the diversity of European fossil hominoids.In his Figure 12, Pickford (2012) compared the male upper canine SMNS 47445 from Germany with several of the available Miocene specimens from the Vallès-Penedès.Middle Miocene taxa from this basin include Pierolapithecus catalaunicus, Anoiapithe-cus brevirostris and Dryopithecus fontani from several localities of the Abocador de Can Mata (ACM) local stratigraphic series (Moyà-Solà et al., 2004, 2009a,b), whereas Late Miocene taxa include Hispanopithecus laietanus and Hispanopithecus crusafonti (Begun et al., 1990;Begun, 1992;Golpe Posse, 1993;Moyà-Solà & Köhler, 1995;Alba et al., submitted).Pickford (2012) provisionally maintained the latter species within Dryopithecus, concluding that it might be a junior synonym of Udabnopithecus garedziensis, and further noting similarities with Anoiapithecus brevirostris and Neopithecus brancoi (considered a nomen dubium by Casanovas-Vilar et al., 2011, but a taxonomically valid taxon by him).Leaving aside Pickford's (2012) particular taxonomic attributions of several specimens, we generally agree with his contention that the paleodiversity of European Miocene hominoids has been underestimated during the last decades (see also Casanovas-Vilar et al., 2011).The aim of this paper is not to discuss Pickford's (2012) taxonomic attributions, but rather to clarify several factual mistakes regarding the identity of several male upper canines figured by this author.

Catalogue numbers
In Pickford's (2012) Figure 12 legend, the letters denoting the H. laietanus IPS18000 canine from Can Llobateres 2 and that of D. fontani IPS35026 from ACM/C3-Ae (see also our Fig.1F-I) are transposed, the former corresponding in fact to his Fig. 12D, and the latter to his Fig. 12E.It is also noteworthy that the canine from Can Llobateres 1 figured by Pickford's (2012) Fig. 12G should be given a different catalogue number than IPS1801.Certainly, this specimen has borne a label with this number for at least 15 years, and has appeared as such in print: Ribot et al. (1996), in particular, referred to it as IPS1801, whereas Moyà-Solà & Köhler (1995) referred to it as IPMC1801 and Alba et al. (2001) as CLL1801.As noted by Alba et al. (submitted), the introduction of modern IPS catalogue numbers in the early 1990s generated some confusion in the literature.Modern catalogue numbers were first used in print by Harrison (1991), who did not provide a modern number for this canine (previously referred to as IPS41 following the older numbering system, e.g.Crusafont Pairó & Golpe Posse; Golpe Posse, 1993).Harrison (1991) explicitly asserted that IPS1801 was the number of a ruminant incisor, which is still present in the ICP collections (DMA, pers.obs.).Hence, as noted by Alba et al. (submitted: Table S1), the hominoid canine specimen should be in fact labeled IPS1765.The latter is the correct number according to the ICP records, even though Harrison (1991) missed this modern catalogue number for some unknown reason, subsequently being incorrectly labeled as IPS1801.
The most serious inadvertent mistake made by Pickford (2012), however, refers to the specimen depicted in his Figure 12H (see our Figure 1A-E).According to the legend, this specimen would be labeled as IPS14741, being attributed to Hispanopithecus laietanus.Throughout the text, Pickford (2012) did not refer specifically to this specimen, although he attributed the Can Llobateres canines to H. laietanus, thus implicitly assuming that this specimen came from Can Llobateres.Neither the catalogue number, nor the taxonomic attribution, nor the locality of provenance are correct.First, according to ICP records, IPS14741 corresponds to a coprolite from the early Pleistocene site of Cal Guardiola.The canine depicted in Pickford's (2012) Figure 12H corresponds in fact to IPS41714, which was originally labeled with the field number ACM14741.No doubt, the similarity between the two numbers is responsible for Pickford (2012) mistaking the ACM field number for the correct catalogue number (preceded by the acronym 'IPS', corresponding to the collections of the Institut Català de Paleontologia Miquel Crusafont).In any case, this mistake ultimately resulted in incorrect taxonomic and locality assignments for this specimen, which are corrected here.
This situation is particularly unfortunate, given that Pickford was allowed to study, but not to publish (either describe or figure) this particular specimen, because it was already under study by the two authors of the present paper.In the acknowledgments of his paper, Pickford (2012) thanked one of us (SMS) for granting him access to the Vallès-Penedès specimens, and he did not include in his paper any of the other unpublished specimens that he was allowed to study.Nevertheless, given the factual inaccuracies that follow from his inadvertent mistake, we feel compelled to clarify both its stratigraphic provenance and taxonomic attribution (see below).We refrain from providing a detailed description of this specimen, since a comparative description of all available hominoid male upper canines from the Vallès-Penedès Basin will be submitted elsewhere (Alba et al., in prep.).

Comparisons and taxonomic attribution
IPS41714 (Figure 1A-E; see also Pickford, 2012: Fig. 12H) resembles other Miocene hominoid homologous teeth from the Vallès-Penedès Basin, which correspond to several dryopithecine genera.It should be noted that here 'dryopithecine' is employed as the semi-formal designation of the subfamily Dryopithecinae, unlike Pickford's (2012) usage of the same term as an informal designation Estudios Geológicos, 68(1), 149-153, enero-junio 2012.ISSN: 0367-0449.doi:10.3989/egeol.40900.180 On the identity of a hominoid male upper canine from the Vallès-Penedès Basin 151 that does not imply any taxonomic rank, and which should be better substituted by 'dryopith'.Similarities between IPS41714 and the upper canines of Vallès-Penedès dryopithecines include the presence of secondary, vertical enamel crenulations on both the lingual and buccal faces and the presence of a well-developed mesiolingual sulcus originating close to the crown base (except in Hispanopithecus crusafonti).IPS41714, however, clearly differs from Late Miocene taxa (Hispanopithecus spp.; see Pickford, 2012: Figs. 12C ,D,G) by displaying a more buccolingually-compressed crown and root, with the former showing a clearly oval occlusal profile (broader towards its mesial portion), instead of the more elliptical and less buccolingually-compressed occlusal contour of Hispanopithecus.IPS41714 also differs from the latter by displaying a shorter and less vertical root, which is markedly tilted distalwards (thus forming a relatively abrupt angle relative to the main basal-apical crown axis), and by further displaying a lingually curved crown relative to the root (instead of being tilted buccalwards).
In the various features mentioned above, IPS41714 most closely resembles the Middle Miocene taxa Dryopithecus (Fig. 1F-I; see also Pickford, 2012: Fig. 12E), Pierolapithecus (Pickford, 2012: Fig. 12F) and Anoiapithecus (Pickford, 2012: Fig. 12A), although on the basis of size and shape, it is attributable to the first of these, given its similarities to the two available canines from the partial face and palate IPS35026 from ACM/C3-Ae (Fig. 1F-I), attributed to D. fontani by Moyà-Solà et al. (2009a;see also Pickford, 2012).Admittedly, IPS41714 is less completely preserved and shows a somewhat more advanced degree of wear, but their morphological features fit very well in spite of the slightly smaller overall dimensions of IPS41714.In contrast, compared to the holotype specimen of Anoiapithecus brevirostris, the two D. fontani individuals display larger dimensions, a less buccolingually-compressed crown, and a larger buccolingual diameter (clearly situated on the mesial portion of the crown and progressively tapering distally).In both size and morphology, the canines attributed to D. fontani resemble more closely those of the holotype of Pierolapithecus catalaunicus than that of Anoiapithecus, although Pierolapithecus differs from Dryopithecus by displaying a relatively more brachyodont crown with a broader (less tapering) distal portion.Unfortunately, small sample sizes preclude a more secure assessment of intraspecific variability.However, given the association of distinctive canine features with other diagnostic features distinguishing the three above-mentioned hominoid taxa (see Moyà-Solà et al., 2004, 2009a,b, and Alba et al., 2010, for further details), an attribution of IPS41714 to D. fontani is justified.