A new actinopterygian fauna from the latest Cretaceous of Quintanilla la Ojada (Burgos, Spain)

Ana BERRETEAGA Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, SP-48080 Bilbao (Spain) and Universidad de Alcalá de Henares, Facultad de Ciencias, Departamento de Geología, Plaza San Diego s/n, SP-28801 Alcalá de Henares (Spain) [email protected]

Francisco José POYATO-ARIZA Universidad Autónoma de Madrid, Departamento de Biología, Unidad de Paleontología, Cantoblanco, SP-28049 Madrid (Spain ) [email protected]

Xabier PEREDA-SUBERBIOLA Universidad del País Vasco/EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, SP-48080 Bilbao (Spain) [email protected]

Berreteaga A., Poyato-Ariza F. J. & Pereda-Suberbiola X. 2011. — A new actinopterygian fauna from the latest Cretaceous of Quintanilla la Ojada (Burgos, Spain). Geodiversitas 33 (2): 285-301. DOI: 10.5252/g2011n2a6.

ABSTRACT We describe a new actinopterygian fauna from the uppermost Cretaceous of Quintanilla la Ojada (Burgos, Spain), in the Villarcayo Sinclynorium of the Basque-Cantabrian Region. It consists mostly of isolated teeth of pycnodon- tiforms (cf. Anomoeodus sp., Pycnodontoidea indet.), amiiforms (cf. Amiidae indet.) and teleosteans (elopiforms: Phyllodontinae indet., Paralbulinae indet.; KEY WORDS aulopiforms: Enchodontidae indet., plus fragmentary fi n spines of Acantho- Osteichthyes, Pycnodontiformes, morpha indet.). Paralbulinae teeth are the most abundant elements in the fossil Amiiformes, assemblage. All the remains are disarticulated and show intense post-mortem Elopiformes, Aulopiformes, abrasion. Th e fossil association has been found in dolomite sandstones that are Acanthomorpha, laterally correlated with the Valdenoceda Formation (Lower to basal Upper Maastrichtian, Maastrichtian) of the Castilian Ramp. Th e observed taphonomic features are Iberian Peninsula, Basque-Cantabrian coherent with the sedimentological interpretation of the fossiliferous beds as Region. coastal deltaic deposits.

GEODIVERSITAS • 2011 • 33 (2) © Publications Scientifi ques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 285 Berreteaga et al.

RÉSUMÉ Une nouvelle faune d’actinoptérygiens du Crétacé supérieur de Quintanilla la Ojada (Burgos, Espagne). Une nouvelle faune d’actinoptérygiens du Crétacé supérieur de Quintanilla la Ojada (Burgos, Espagne), localité située dans le Synclinorium de Villarcayo dans la région basco-cantabrique, est décrite ici. La plupart du matériel correspond à des dents isolées de pycnodontiformes (cf. Anomoeodus sp., Pycnodontoidea indet.), d’amiiformes (cf. Amiidae indet.) et de téléostéens (Elopiformes : Phyllo- dontinae indet., Paralbulinae indet. ; Aulopiformes : Enchodontidae indet., ainsi que des fragments d’épines d’Acanthomorpha indet.). Les dents de Paralbulinae MOTS CLÉS Osteichthyes, sont les éléments les plus abondants de l’association fossile. Tous les restes sont Pycnodontiformes, désarticulés et ont subi une abrasion post-mortem intense. Les fossiles ont été Amiiformes, Elopiformes, trouvés dans des dolomies sableuses qui sont corrélées latéralement avec la For- Aulopiformes, mation Valdenoceda (Maastrichtien inférieur à base du Maastrichtien supérieur) Acanthomorpha, de la Rampe Norcastillane. Les caractéristiques taphonomiques observées sont Maastrichtian, péninsule Ibérique, cohérentes avec l’interprétation sédimentologique des couches fossilifères comme région basco-cantabrique. ayant été déposées dans un milieu deltaïque côtier.

INTRODUCTION work in progress). Th e present paper aims to describe the actinopterygian fauna that occurs in this new ver- Upper Cretaceous marine localities that have yielded tebrate site, which consists mostly of teeth of pycno- actinopterygian remains are presently rather scarce dontiforms, amiiforms, elopiforms, and aulopiforms, in the sedimentary basins of Spain (see Poyato-Ariza plus fragmentary acanthomorph spines. et al. 1999a). One of the most important known sites is Albaina, located in the Laño quarry (Condado de Treviño; area situated within the Basque Country but MATERIAL AND METHODS administered by the province of Burgos), where pyc- nodontiform and teleostean fossils have been found Th e vertebrate fossil remains of Quintanilla la Ojada in Upper Maastrichtian marine levels (Poyato-Ariza (Burgos) have been collected since 2002. Some teeth et al. 1999b). In the same quarry, the Upper Campa- were picked up in the fi eld (or extracted from the nian to lowermost Maastrichtian fl uviatile deposits of matrix after bulk rock processing), but most of the the Laño site have yielded scarce lepisosteiform and remains were obtained by screen-washing of the teleostean remains (Cavin 1999). Th us, the osteich- sediment. About 300 kg of sediment were washed thyan fossils of the Laño quarry occur at two distinct through 2 mm, 1 mm and 0.5 mm stainless steel stratigraphic horizons and were accumulated in quite meshes with running water in order to search for diff erent circumstances; both Albaina and Laño are microremains. Th e fi ne fraction was washed again part of the Miranda-Treviño Synclinorium (Baceta using a hand sieve of 0.25 mm. Th e sorting of the et al. 1999). Recently, additional marine fi sh-bearing teeth and other microremains was made by means localities of Late Cretaceous age have been discovered of stereomicroscopes. Th e material is housed in the in the Basque-Cantabrian Region (López-Horgue & laboratory of the Departamento de Estratigrafía y Poyato-Ariza 2005, work in progress). Th e fossil as- Paleontología of the Universidad del País Vasco/ sociation of the Maastrichtian locality of Quintanilla Euskal Herriko Unibertsitatea (Bilbao) (EHUEP). la Ojada (Burgos) is composed of actinopterygian, About 200 actinopterygian isolated teeth and a few selachian, and reptile remains (Berreteaga et al. 2010, fi n spine fragments are known.

286 GEODIVERSITAS • 2011 • 33 (2) New actinopterygian fauna from Spain

N Paleozoic Bay of Biscay Permian and Triassic Santander Bayonne Keuper (diapir) Hettangian- San Sebastián Kimmeridgian (Marine Jurassic megasequence) C Bilbao CM Upper Tithonian to Barremian (Purbeck & AM Weald megasequences)

Ebro Aptian-Albian to Quintanilla la Ojada Reservoir Vitoria Lower Cenomanian * Pamplona Late Albian-Santonian Albaina P volcanics V * Tertiary Ebro U Middle Cenomanian to Tertiary Duero Basin Maastrichtian Basin Logroño Tertiary Iberian Burgos Peninsula Principal fault 50 km * Vertebrate sites C Cabuerniga fault P Pamplona fault Basque Massifs (emerged areas U Ubierna fault CM affected by erosion)

V Villela fault AM Asturian Massif

FIG. 1. — Geological map of the Basque-Cantabrian Region showing the location of the actinopterygian sites mentioned in the text.

LOCATION AND GEOLOGICAL SETTING drichthyans and osteichthyans (López-Horgue & Quintanilla la Ojada is located in the northern part of Poyato-Ariza 2005, work in progress). Burgos province (Losa Valley), approximately 22 km Th e Cretaceous-Tertiary deposits in the northwest east and 14 km northeast of Villarcayo and Trespaderne limb of the Villarcayo Synclinorium show several localities, respectively. Geologically, it is situated in hundred meters of siliciclastic and carbonate materi- the northern limb of the Villarcayo Synclinorium, in als arranged in transgressive-regressive cycles (Floquet the northern part of the Basque-Cantabrian Region. 1991, 1998; Berreteaga 2008). Time-equivalent Actinopterygian remains have been found in a quarry materials in the Miranda-Treviño Synclinorium are near the village of Quintanilla la Ojada. also arranged in comparable transgressive-regressive Th e Villarcayo and Miranda-Treviño synclinoria cycles (Baceta 1996; Baceta et al. 1999). Th e Maas- belong to the southern part of the Basque-Cantabrian trichtian deposits of both synclinoria have yielded Basin (Fig. 1). During the Late Cretaceous and abundant actinopterygian remains. Early Paleogene, the sedimentation throughout the study area comprised varied depositional environ- FACIES DESCRIPTION ments ranging from inner self to coastal and even Th e lower part of the Quintanilla la Ojada section continental ones (Floquet 1991; Pluchery 1995). consists of bioclastic calcarenites that include: bivalves, In Campanian-Maastrichtian time interval the gastropods, brachiopods, green and red algae, sponges, regression trend in the Basque-Cantabrian Basin oysters, echinoderms, bryozoans, serpulids, foramini- resulted in the progradation of deltaic siliciclastic fers, ostracods, and crustaceans (Floquet 1991). Th ese systems and the development of shallow marine calcarenites correspond to the top of the Tubilla del environments, inhabited by a fauna rich in chon- Agua Formation, as defi ned by Floquet et al. (1982).

GEODIVERSITAS • 2011 • 33 (2) 287 Berreteaga et al.

Th is formation is dated from the fi nal Santonian to (Sobrepeña Formation, Floquet 1991; Berreteaga probably early Campanian by foraminifera and am- et al. 2008a) (Fig. 2). monites (Floquet 1991, 1998; Gräfe 2005). Th e Tubilla del Agua Formation shows a gradual FACIES INTERPRETATION facies-transition to the overlaying siliciclastic unit. Th e Th e Rioseco Member is interpreted as an easterly latter is nearly 60 m in thickness in the Quintanilla progradational deltaic system (Floquet 1991; Ber- la Ojada site; it consists of well-rounded conglomer- reteaga 2008). Th e boundary between the Rioseco ates and sandstones of mainly quartz composition Member and the Valdenoceda Formation is an erosive arranged in meter-scale tabular sets with troughs surface defi ned by conglomerates, and is interpreted cross-stratifi cation, alternating with thin (cm thick) as a ravinement surface with conglomeratic transgres- levels of laminated greenish siltstones. Th is siliciclastic sive lag deposits. Th is surface is correlated with the unit shows important thickness variations across the basal discontinuity of the DC12 depositional cycle Villarcayo Synclinorium, decreasing their thickness (Fig. 2). Th is is one of the depositional transgressive- towards the North-West and the South. It reaches its regressive cycles defi ned in the North-Castilian Ramp maximum thickness (70-80 m) in the Quintanilla la by Floquet (1998). Th e base of the Valdenoceda Ojada and the Rioseco sections. Th e unit, named Rio- Formation represents the increasing infl uence of seco Member, is laterally equivalent to the Moradillo marine conditions in the system. Th e stratigraphical de Sedano, Quintanaloma, and Sedano formations, position of the Valdenoceda Formation suggests an which have been ascribed to the Campanian-Lower Early to basal Late Maastrichtian age (Floquet 1991; Maastrichtian and even basal Upper Maastrichtian Berreteaga 2008; Berreteaga et al. 2010). (Floquet et al. 1982; Floquet 1991, 1998; Berreteaga Time-equivalent transgressive-regressive cycles 2008). In the study area, these three formations show formed in similar conditions outcrop in the Miranda- minor lithological changes that are hardly noticeable Treviño Synclinorium (Pluchery 1995; Baceta 1996). because the zone corresponds to the area with the In the Albaina site (Condado de Treviño), vertebrate highest sedimentation rates of the Rioseco delta (Flo- fossils have not been discovered related to the DC12 quet 1991; Pluchery 1995; Berreteaga 2008). depositional cycle, but near the base of the DC13 Th e Rioseco delta sediments are overlain by lu- depositional cycle (Fig. 2). Th e shallow marine cal- tites or coarse gravel sized conglomerates containing carenites of Albaina are laterally equivalent to the abundant vertebrate fossils. Both lithologies are only Torme Formation (Pluchery 1995; Cappetta & Corral 50 cm thick and are overlain by sandy dolomites and 1999; Berreteaga 2008). In Albaina, vertebrate fos- organic-matter rich clays. Th ese materials (dolomites sils also occur associated with a transgressive surface and clays) are organised in metre-scale upward thin- (Intra-Maastrichtian Unconformity of Baceta et al. ning sequences. Th ese sequences are overlaid by a 1999) (Fig. 2). Th is discontinuity is correlated with 30-40 m thick massive dolomite body. Th e sequences the cycle boundary of CB13 (sensu Floquet 1998). and overlaid dolomite body are regarded as laterally Th e Albaina site is regarded as Upper Maastrich- equivalent to the Valdenoceda Formation (Floquet tian (Baceta et al. 1999); the selachian association et al. 1982; Floquet 1991; Berreteaga 2008). Moreo- indicates a Late but not latest Maastrichtian age ver, the type of the sequences is characteristic of the (Cappetta & Corral 1999). Valdenoceda Formation. Th e vertebrate fossils occur in the lower part of the Valdenoceda Formation, in TAPHONOMIC OBSERVATIONS the two fi rst meters, immediately above and closely Th e fact that only isolated teeth and fi n spines have related to the boundary between the Rioseco Mem- been unearthed from Quintanilla la Ojada obviously ber and the Valdenoceda Formation (Berreteaga indicates the occurrence of total disarticulation of et al. 2008b). the individuals, and, very probably, intense selective Finally, after a gradual increase in the clay content transport as well. Th is is supported by recurrent of the dolomites, there are nearly 30 m of variegated observation of noticeable, substantial abrasion clays corresponding to a coastal plain environment facets (e.g., Figs 3C1, D2; 4A1, D2). On the teeth,

288 GEODIVERSITAS • 2011 • 33 (2) New actinopterygian fauna from Spain

Quintanilla la Ojada Albaina section section Lower Sequences Thanetian Marine Floquet (1998) Landraves Fm. (base) CB-15 Distal Litoral Escaño Fm.

Danian s.l. Proximal DC-14 70 m Continental

Fresnedo Fm. Albaina CB-14 upper DC-13 Torme Fm. CB-15

upper? to Sobrepeña Fm. middle

10

DC-12 IMU

middle? Valdenoceda Fm. Dolomitic sandstones

Maastrichtian to lower Dolomicrites

100 Variegated clays Quintanilla la Ojada Sandstones CB-12 lower Conglomerates Sedano Fm. DC-11 Quintanaloma Limestones Fm. Marls CB-12 Campanian Clayed limestones Moradillo de 50

Rioseco Member Sedano s.s. Fm. Oblique laminations Gryphaea DC-10 Oolites Upper Tubilla del Aqua Santonian Fm. (top) Rudists 0 m Stromatolites and thrombolites 75 km Vertebrate fossil remains

FIG. 2. — Correlation of transgressive-regressive depositional cycles from the Late Cretaceous sections in Villarcayo Synclinorium (based on the Quintanilla la Ojada section) and Miranda-Treviño Synclinorium (based on the Albaina section). The Upper Cretaceous interval shows stratigraphical units, actinopterygian fossil-bearing units and transgressive-regressive depositional cycles. The cycles described from the Upper Santonian until the end of the Cretaceous are DC10-13, all of them are included in a transgressive-regressive megacycle called LTC4 by Floquet (1998). Abbreviation: IMU, Intra-Maastrichtian Unconformity. post-mortem abrasion facets are distinguishable Following the abrasion categories described by from anatomic wearing facets during lifetime of the Vullo (2007), 95% of the studied actinoptery- individual by: a) the aspect of their surface, which gian teeth show moderate to advanced abraded is, respectively, rough rather than smooth; b) the stage, indicating heavy transport. Such heavy size of each facet in relation to the total surface of transport must have been biostratinomic, since the tooth, which is extensive rather than limited, the results of geochemical analysis (e.g., dis- respectively; c) their pattern of distribution, which tribution of rare earth elements) in vertebrate is regular rather than irregular, and d) their situa- fossils of Quintanilla la Ojada allow us to reject tion, which occurs on mechanical contact surfaces the hypothesis of reelaborated bones in this site rather than on anatomical contact surfaces. (Berreteaga 2008).

GEODIVERSITAS • 2011 • 33 (2) 289 Berreteaga et al.

SYSTEMATIC PALAEONTOLOGY in major axis length. Th ree other teeth have sub- circular contour in occlusal view, 2.0 to 3.1 in Th e osteichthyan fi sh remains from Quintanilla la major axis length; EHUEP Q8 is the biggest one, Ojada are represented by isolated actinopterygian 7.3 × 5.7 mm, with the occlusal side fl attened by teeth mostly, plus a few specimens of fi n spines. abrasion (Fig. 3B). In general, isolated actinopterygian teeth provide limited taxonomic information, and usually present DISCUSSION considerable parataxonomic problems. In this par- Molariform teeth sensu Poyato-Ariza (2005a) are ticular locality, the abrasion of the material is yet typical of pycnodontiforms, and are always present another limitation for this type of study, as it often on the vomer and the prearticular of this neoptery- rubs out signifi cant taxonomic characters such as gian order. Both oval and subcircular molariform crests, ridges, pits, and so on. For these reasons, teeth are broadly present among Pycnodontiformes the taxonomic assessments in the present paper are (Poyato-Ariza & Wenz 2002), so that they cannot be to be taken with precaution, and are made mostly assessed to any particular taxon within this group. at higher-rank level. Lower-rank assignments are As for diversity, both morphologies are often present proposed only in the cases were the assessment is on a single individual (e.g., Poyato-Ariza & Wenz reasonably reliable and parataxonomic problems 2002: fi gs 21A, 22C; Poyato-Ariza 2003: fi g. 1C, are limited. In addition, the teeth are also classifi ed D), so they do not necessary indicate the presence according to morphotypes, as this classifi cation can of two distinct taxa. Th e molariform teeth of the be very useful for palaeoecological comparisons Pycnodontidae are relatively higher in lateral view with other fossil assemblages (e.g., Buscalioni et al. than those of the Sparidae, which have been cited 2008). to occur in the Late Cretaceous (e.g., Buff etaut et al. 1996), and whose crowns are much more fl attened. Class OSTEICHTHYES Huxley, 1880 Subclass ACTINOPTERYGII Cope, 1887 Division HALECOSTOMI Suborder PYCNODONTOIDEI Nursall, 1996 Regan, 1923 sensu Patterson 1973 Superfamily PYCNODONTOIDEA Order PYCNODONTIFORMES Berg, 1933 Agassiz, 1833 sensu Poyato-Ariza & Wenz 2002 Pycnodontiformes indet. Pycnodontoidea indet.

Morphotype 1: molariform teeth – type A Morphotype 2: incisiform teeth (Fig. 3A, B) (Fig. 3C-E)

MATERIAL EXAMINED. — Eight molariform teeth (EHUEP MATERIAL EXAMINED. — Nine incisiform teeth (EHUEP Q1-Q8). Q9-Q17).

DESCRIPTION DESCRIPTION Molariform teeth presenting a low crown, very Incisiform teeth present a high crown, enlarged enlarged, with great development of enamel, and and fl attened, with labial side more or less con- especially of dentine; the basal part, which is devoid vex, lingual side markedly concave, and occlusal of dentine, is consistently lost, as it remains inside border quite narrower than the base (labio-lingual the bone when the tooth is disarticulated. EHUEP dimension). See also Nursall (1996a), Poyato- Q7 is a very broken and heavily transported tooth, Ariza & Wenz (2002) and Poyato-Ariza (2005a) probably originally oval in shape (Fig. 3A); there for further details. In contrast with molariform are also three teeth with oval contour in occlusal teeth, incisiform teeth usually exhibit at least view and heavy abrasion facets, 1.6 to 2.5 mm part of their base together with their crown. In

290 GEODIVERSITAS • 2011 • 33 (2) New actinopterygian fauna from Spain

A1 A2 A3 B1

C1 C2 C3

F1

D1 D2 D3

F2

E 1 E2 E3

H3

G1 G2 H1 H2

FIG. 3. — Pycnodontiforms and amiiforms from Quintanilla la Ojada (Burgos): A, B, Pycnodontiformes indet., molariform teeth EHUEP Q7 in basal (A1), occlusal (A2) and lateral (A3) views and EHUEP Q8 in occlusal (B1) view; C-E, Pycnodontoidea indet., incisiform teeth EHUEP Q14, Q16, Q15 in labial (1), lingual (2) and mesiodistal (3) views; F, Pycnodontoidea indet., branchial tooth EHUEP Q20 in lateral (F1) and apical (F2) views; G, cf. Anomoedus sp., molariform tooth EHUEP Q23 in occlusal (G1) and basal (G2) views; H, cf. Amiidae indet., conic-styliform tooth EHUEP Q26 in mesiodistal (H1), (H2) and occlusal (H3) views. Scale bars: A, H, 1 mm; B, G, 4 mm; C-F, 2 mm.

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Quintanilla la Ojada, incisiform teeth present high. In this locality, they are scanter than other intense abrasion, which has considerable reduced pycnodont teeth, and range in size from 0.6 to their original size, and even veiled their original 2.8 mm in maximum dimension. morphology (e.g., Fig. 3C). Th e parallel canals of the dentine layer are often clearly visible. Th e DISCUSSION bigger tooth (Fig. 3E) is 7.6 mm high (crown is Th is tooth morphotype has for a long time been 5.5 mm high), 4.2 mm in maximum width, and assessed to “Stephanodus” Zittel, 1883, but this 5.2 mm in labiolingual dimension; the smallest is clearly a parataxon, and must not be used in teeth are about 1.2 mm in height and maximum formal taxonomy until its in-depth revision is width. accomplished. Th is tooth morphotype is found in the branchial chamber of a number of Pycno- DISCUSSION dontoidea, and is not known outside the super- Th e inclination of the occlusal border from the family (Kriwet 1999; Poyato-Ariza & Wenz 2002, horizontal plane (Fig. 3D1, D2, E1, E2) was most 2005; FJPA, pers. obs.), so that their occurrence likely caused by wearing during life time of the in this locality does not necessarily indicate the individual (FJPA, pers. obs. on articulated pycno- presence of another distinct taxon of this super- dontoid specimens; for a list of taxa and material, family. Although elongated branchial teeth have see Poyato-Ariza & Wenz 2002, 2004), but the been reported outside the Pycnodontoidea, the uniform, intense abrasion observed is post-mortem type of broad pharyngeal tooth herein reported (see criteria above). Th e presence of fully incisiform has never been found in non-pycnodontoid pyc- premaxillary and dentary teeth is a typical diagnostic nodontiforms. Similar pharyngeal teeth have character of the Pycnodontoidea (Poyato-Ariza & been, nonetheless, found in other groups, such Wenz 2002). Although it is reverted in some mem- as Semionotiformes (e.g., Th ies 1989). Since they bers of the Pycnodontoidea, such as Nursallia Blot, are associated to pycnodont teeth, and since re- 1987 (Poyato-Ariza & Wenz 2002) and Akromystax mains of other candidate groups are absent, they Poyato-Ariza & Wenz, 2005, and is therefore not are more confi dently assessable to pycnodontoids an autapomorphy, it is not known in any taxon than to other groups. outside this superfamily; it is therefore a synapo- morphy of this group, and consequently a reliable taxonomic character. Family PYCNODONTIDAE Agassiz, 1833 sensu Nursall 1996b Genus Anomoeodus Forir, 1887 Pycnodontoidea indet. cf. Anomoeodus sp.

Morphotype 3: branchial teeth Morphotype 4: molariform teeth – type B (Fig. 3F) (Fig. 3G)

MATERIAL EXAMINED. — Th ree branchial teeth (EHUEP MATERIAL EXAMINED. — Th ree broken and incomplete Q18-Q20). teeth (EHUEP Q21-Q23).

DESCRIPTION DESCRIPTION Th ese branchial teeth are hook-shaped, pedicellate, Molariform teeth whose major axis, in occlusal and very fl attened, markedly laterally compressed; view, is about three times longer than the minor they present a wide base and a conspicuous, stout axis; their medial edge is tapering, and the gen- anterior process (Fig. 3F). Th ey are usually higher eral outline, in occlusal view, is curved and sort than long, but they can also be wider than high; one of drop-shaped. Th e occlusal surface is eroded, of these three teeth is 1.0 mm wide and 0.6 mm so that it is smooth, and the presence of crests,

292 GEODIVERSITAS • 2011 • 33 (2) New actinopterygian fauna from Spain

ridges, or any other kind of ornamentation can- curved in lateral view. Th e crown presents no striae, not be estimated. Th e biggest preserved fragment lateral crests, projections, or processes whatsoever. (Fig. 3G) is 5.1 mm wide and 9.8 mm long (pre- Th e enamel is thin, smooth, and translucid. Most served portion). specimens are broken, yet they do not show heavy abrasion facets. Th is type of teeth is relatively rare DISCUSSION in this locality. Th e only complete known tooth Th is crown morphology, only found in prearticular measures 1.2 mm of total height and 0.4 mm of teeth, is described in detail and compared with other maximum diameter; the glossy crown is 0.6 mm molariform shapes in Poyato-Ariza & Wenz (2002: high (Fig. 3H). Other teeth are more or less frag- 174, 175). Only two pycnodont genera, reliably mentary: the preserved part of the biggest one is identifi able from articulated remains, bear this type 1.9 mm high (glossy crown, maybe incomplete), of molariform, tapering-edged teeth (Poyato-Ariza & so that its total height would be at least 3.8 mm, as Wenz 2002). One of them is Ichthyoceros Gayet, estimated from comparison of crown height to total 1984, from the Cenomanian of the Lebanon, but height in the complete tooth just described. the size of these teeth in this genus is signifi cantly smaller than the teeth from Quintanilla la Ojada. DISCUSSION Th e other genus where this type of teeth is known, Small conic-styliform teeth of this kind are typical and for which are typical, is Anomoeodus, known of small to medium-sized fi shes with ichthyofagous from the Early Cretaceous of Uña (Kriwet 1999) diet and predatory habits. When compared with and from the Late Cretaceous of the Netherlands other geographically close ichthyofagous forms, and England (e.g., Poyato-Ariza & Wenz 2002: they appear to be typical of the family Amiidae 143). In both cases, the teeth are ornamented, with (e.g., Grande & Bemis 1998), which includes both a central ridge and/or light to strong crenulations marine and freshwater fossil forms plus the Recent (Kriwet 1999; Poyato-Ariza & Wenz 2002). Th e Amia calva Linnaeus, 1766, which is an abundant teeth from Quintanilla la Ojada are devoid of any freshwater North American species. Th e teeth of ornamentation, but this is probably due to abra- other typical small- to medium-sized ichthyo fagous sion during the fossilization process. In addition, fi shes from the Late Jurassic-Cretaceous, such as the specimens are broken and incomplete, so their the Aspidorhynchidae, Ionoscopidae or Allothris- assessment to cf. Anomoeodus, based on the general sopidae, are quite diff erent in curvature, height to morphology combined with their relatively big size, diameter ratio, morphology of the occlusal end of is to be taken with precaution. crown, etc. (e.g., Th ies & Mudroch 1996; Brito 1997). Other ichthyofagous Mesozoic fi shes, such as the Ichthyodectidae, bear much bigger teeth. Order AMIIFORMES Other similar teleostean teeth seem to present a Hay, 1929 sensu Grande & Bemis 1998 not so glossy and traslucid crown (Martín-Abad & Family AMIIDAE Bonaparte, 1838 Poyato-Ariza pers. obs.). Pending a thorough revi- cf. Amiidae indet. sion of this type of teeth from a reliable sample of articulated material, it is herein considered as cf. Morphotype 5: conic-styliform teeth Amiidae indet. (Fig. 3H)

MATERIAL EXAMINED. — 11 teeth (EHUEP Q24- Subdivision TELEOSTEI Müller, 1846 Q34). sensu Patterson & Rosen 1977 Order ELOPIFORMES DESCRIPTION Greenwood, Rosen, Weitzman & Myers, 1966 Small, conic-styliform teeth with pointed, well- Superfamily ALBULOIDEA defi ned enameloid, glossy crown, straight to slightly Greenwood, Rosen, Weitzman & Myers, 1966

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Family PHYLLODONTIDAE Sauvage, 1875 DISCUSSION sensu Estes 1969 Phyllodont teeth, unlike pycnodont teeth, grow Subfamily PHYLLODONTINAE Sauvage, 1875 as replacement sets (e.g., Estes 1969). In phyl- sensu Estes 1969 lodontines, the replacement sets grow very closely Phyllodontinae indet. arranged, hence the remarkable fl atness of each individual tooth. However, no complete phyllodont Morphotype 6: fl attened teeth dentitions are known from this locality (unlike (Fig. 4A) Albaina: Poyato-Ariza et al. 1999b). Although the individual tooth morphology of this type of tooth MATERIAL EXAMINED. — Th ree teeth (EHUEP Q35- is consistent with that of the genus Phyllodus Agas- Q37). siz, 1839, generic assessments within this group are mostly based on characters of complete tooth DESCRIPTION plates, and therefore, isolated teeth can only be Th ese teeth are somewhat similar to molariform assessed, with the necessary precautions, to the teeth because their single crown is robust, stout, Phyllodontinae. and subcircular in occlusal view (Fig. 4A1). But this morphotype is much more fl attened, so that their thickness, measured as their occluso- Subfamily PARALBULINAE Estes, 1969 basal dimension, is less than one third of their Paralbulinae indet. diameter in occlusal view (Fig. 4A3), quite lower than that of molariform teeth (Fig. 3A3). Th eir Morphotype 7: globular teeth dentine and enamel are less developed than those (Fig. 4B, C) of molariform teeth, and they lack the small central cavity that all molariform teeth exhibit MATERIAL EXAMINED. — 122 teeth (EHUEP Q38- Q160). when observed in basal view (Figure 4A2 ver- sus Figure 3A1, respectively). In Quintanilla la Ojada, fl attened teeth are very rare and only DESCRIPTION three specimens are known. Specimen EHUEP High, robust teeth of globular shape in overall Q35 measures 2.3 × 2.9 mm in occlusal view, morphology, circular in occlusal view and nearly and 0.9 mm thick in occluso-basal dimension. It circular in lateral view, except for their insertion shows life-wearing facets, indicating that it was base. Th ey present a well-developed basilar foramen probably a functional rather than a replacement with strong, striated borders (Fig. 4C2). Th e enamel tooth, and also strong transport facets, with loss is thick, heavy, opaque, and blackened. Th ese teeth of the whole central region. Specimen EHUEP are, by far, the most abundant actinopterygian re- Q36 (Fig. 4A) is the biggest one, as it measures mains in Quintanilla la Ojada. Th ey usually show 6.6 × 4.7 mm in occlusal view, and 1.9 mm in life-wearing facets, and especially transport facets; occluso-basal dimension. Th is specimen is more whenever accessible to observation (some teeth irregular in contour, with concave occlusal sur- with less intense wearing and abrasion), certain face and convex basal surface. It shows abrasion ornamentation can be observed in form of faint, facets, but apparently no life-wearing surfaces, diverging crests with tiny intercalated tubercles suggesting it was a replacement, non-functional (Fig. 4B1). Th ese teeth range from 0.6 to 2.3 mm tooth. Finally, specimen EHUEP Q37 measures in diameter, although the majority of them meas- 5.0 × 4.1 mm, and is heavily eroded by abra- ures around 1.5 mm. sion. In all three cases, abrasion prevents from observing any tooth ornamentation; the irregular DISCUSSION exposures (Fig. 4A1) correspond to the inner tis- Paralbulinae teeth, like Phyllodontinae teeth, sular structure. grow as replacement sets, although the sets, and

294 GEODIVERSITAS • 2011 • 33 (2) New actinopterygian fauna from Spain

A1 A2 B1

A3

B2

C1 C2

B3

D D 1 2 D3

E1 E2 E3

G1 F1 F2 F3

G2

FIG. 4. — Teleosteans from Quintanilla la Ojada (Burgos): A, Phyllodontinae indet., fl attened tooth EHUEP Q36 in occlusal (A1), basal (A2) and lateral (A3), views; B, C, Paralbulinae indet., globular teeth EHUEP Q61-62 in occlusal (1), basal (2) and inclined lateral (3) views; D-F, Enchodontidae indet.; D, caniniform teeth EHUEP Q175 (morphotype A); E, EHUEP Q186 (morphotype B); F, EHUEP Q193 (morphotype C) in labial (1), lingual (2) and mesiodistal (3) views; G, Acanthomorpha indet., fragmentary fi n spine EHUEP Q200 in posterior (G1) and anterior (G2) views. Scale bars: A, D-F, 4 mm; B, G, 1 mm; C, 2 mm.

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consequently the teeth, are quite diff erent in the specimen prevents from knowing their precise morphology and arrangement, much less compact shape (simply curved or sigmoid). Wearing would than phyllodontine sets (e.g., Estes 1969), and, prevent from observing striae, had these been consequently, individual teeth are quite higher, present. Th e material is too fragmentary to provide nearly spherical. As for phyllodontine teeth, no accurate size range estimations: the preserved part complete paralbuline dentitions are known from of the biggest specimen (Fig. 4D) is 18.9 mm; its this locality (unlike, again, Albaina). Although maximum width at about the mid-length of the the individual tooth morphology is consistent preserved part of the crown. with that of the genus Paralbula Blake, 1940, in- cluding the development of the basilar foramen, DISCUSSION generic assessments, within this group as well, As for molariform and incisiform teeth above, are greatly based on characters of complete tooth the term caniniform is used as a descriptive and plates. Consequently, these isolated teeth can only ecomorphological term only, and does not have be assessed, with the necessary precautions, to the any kind of taxonomic or phylogenetic implication Paralbulinae. on itself. Th is type of caniniform teeth, so strong and elongated, is typical of the Enchodontidae, who present surprisingly big teeth with this mor- Order AULOPIFORMES Rosen, 1873 phology on their dentary, dermopalatine, and Family ENCHODONTIDAE Lydekker, 1889 ectopterygoid bones (e.g., Goody 1969; Chalifa Enchodontidae indet. 1996; Poyato-Ariza et al. 1999b). Assessment of these teeth to a particular genus within the Encho- Morphotype 8: caniniform teeth – type A dontidae is not reliable, since key characters such (Fig. 4D) as size ranges, degree of elongation, and complete lateral profi le are not accurately observable in this MATERIAL EXAMINED. — 28 teeth (EHUEP Q161-Q183, material, and it is only the combination of these Q205-Q209). characters, together with the presence of crests on the cutting edges (Poyato-Ariza et al. 1999b) DESCRIPTION that allows consistent assessment to Enchodus Enchodontidae material from this locality is quite Agassiz, 1835 (or to any other particular genus). fragmentary and very worn out. Th e term canini- Th ese teeth are therefore considered as Encho- form is used for teeth that are stout, strong, very dontidae indet. elongated, and laterally compressed, presenting cutting edges and pointed to slightly blunt oc- clusal end. Th ose of this locality present a base Morphotype 9: caniniform teeth – type B that is not expanded, but is well marked, bearing (Fig. 4E) tiny pits. Th e contour of the crown seems more or less straight in labial and lingual views, and MATERIAL EXAMINED. — Five teeth (EHUEP Q184- slightly curved backwards in mesial and distal Q188). views. Th eir incompleteness and bad state of preservation, though, prevent from knowing their DESCRIPTION precise degree of curvature, or possible sigmoid Unlike caniniform teeth of type A, the contour shape when complete. Th e lingual side is always of the crown in caniniform teeth type B is clearly quite fl atter than the labial side, which is rather sigmoid in labial and lingual views (Fig. 4E), in- convex. Some better preserved remains (Fig. 4D) stead of straight, and they are straight in mesial exhibit sharp crests developed on their cutting and distal views instead of slightly curved back- edges (mesial and distal). Th ese crests are also wards (Fig. 4E3). In addition, both sides, lingual slightly curved, although the incompleteness of and labial, are relatively fl at, rather than convex.

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Th e biggest specimen, 15.1 mm preserved high Superorder ACANTHOPTERYGII Klein, 1885 and 5.4 mm of maximum width (Fig. 4E); all sensu Johnson & Patterson 1993 specimens are fragmentary, incomplete. Crests on Clade ACANTHOMORPHA Rosen, 1973 its cutting edges are not observable, but this may sensu Johnson & Patterson 1993 be due to abrasion. Acanthomorpha indet. DISCUSSION (Fig. 4G) See above for caniniform teeth – type A. MATERIAL EXAMINED. — Seven fragments of spines (EHUEP Q197-Q203). Morphotype 10: caniniform teeth – type C (Fig. 4F) DESCRIPTION Th e fragments of fi n spines are very partial: only MATERIAL EXAMINED. — Eight teeth (EHUEP Q189- the basal parts are preserved. Th ey are apparently Q197). fragile, and quite worn out. Th e widest fragment measures 2.4 mm in maximum base width. When DESCRIPTION better preserved, the base of these fi n spines shows: Th ese teeth are scarcer and quite similar in general one pair of well-developed basal processes; one pair shape, although relatively less elongated, that is, of large, comparatively bigger lateral processes; and comparatively larger, than those of type A and B; a small anterior median process (Fig. 4G). Th ey unlike type A and B, the maximum width occurs at seem to present one single median cavity on their the base of the tooth. Type B teeth are also appar- posterior side. Th e visible ornamentation consists ently smaller. Th e contour of type C teeth is clearly of one pair of big, deep basal grooves. sigmoid in labial and lingual views (Fig. 4F1, F2), and straight in mesial and distal views (Fig. 4F3). DISCUSSION Th e only complete tooth of this (and the other Th is is the only fi sh material other than teeth known enchodontid) type is 7.0 mm high and 2.9 mm in from Quintanilla la Ojada. Johnson & Patterson maximum width (Fig. 4F). (1993) affi rm that true fi n spines, that is, azygous, unsegmented, and laterally fused, are a diagnostic char- DISCUSSION acter of the Acanthomorpha (i.e. all Acanthopterygii Th e three diff erent types of caniniform teeth excluding the Atherinomorpha). Th eir ornamentation, probably represent at least two distinct taxa, as crests, processes, cavity, and general morphology are the diff erence in meso-distal contour (sigmoid quite diff erent from polypteriform or siluriform fi n in B and C here versus straight in type A) might spines. Th e precise morphology, variability, and taxo- be signifi cant at specifi c, and even generic level nomic distribution of the fi n spines among Cretaceous (in combination with other characters that are acanthomorphs are at present very poorly understood, not observable in this material). In contrast, type and consequently the scarce material from this local- C might correspond to younger individuals of ity is referred to Acanthomorpha indet. the same taxon than type B, or even to smaller teeth of the same individual: it is typical of en- DISCUSSION chodontids to present the fi rst (most anterior) tooth remarkably big and elongated on the den- Osteichthyan fi shes are, in general, potentially mis- tary, dermopalatine, and ectopterygoid bones. leading palaeoenvironmental indicators (Poyato- As a consequence, the occurrence of a third Ariza et al. 1998), so that the mere presence of any morphotype is not necessarily an indication of particular taxon is not a necessary indication of any the presence of a third distinct taxon within the particular environment. Not even the presence of Enchodontidae of this locality. pycnodontiform remains is a necessary indication of a

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marine palaeoenvironment, since these fi shes are also and even in relative abundance (Table 1). For instance, known from continental palaeoenvironments (Nursall phyllodontine teeth are rare, much scarcer than the 1996b; Poyato-Ariza et al. 1998; Poyato-Ariza 2005a- extremely abundant paralbuline teeth in both locali- c). Only the ensemble of the community, studied ties. However, whenever the taphonomic bias and the in combination with all other fauna and fl ora, and state of preservation allow observation of signifi cant especially in combination with sedimentological and characters, some diff erences in lower rank taxa can geochemical data, is palaeoecologically reliable. Th is be found. For instance, the generic assessments that especially applies to the fi sh material of this locality, can be made for pycnodonts are diff erent in both where the remains are so partial that most of them localities (cf. Anomoeodus sp. from Quintanilla la cannot be assessed to any low rank taxon. Ojada, also reported from Uña [Kriwet 1999]; cf. In overall, there are 10 diff erent tooth morphotypes Paramicrodon sp. from Albania). Interestingly, the known from Quintanilla la Ojada. Th is number of most derived forms present in both Quintanilla la diff erent morphotypes, nonetheless, does not neces- Ojada and Albaina are diff erent low rank taxa of the sarily correspond to the same number of distinct taxa. same high rank taxon (Acanthomorpha). Th e base Th e number of pycnodont taxa (4 morphotypes) can of the acanthomorph fi n spines from Quintanilla be estimated in a minimum of 2, but no certainty la Ojada, with their big pair of lateral processes and of additional diversity can be provided (see particu- smaller anterior median process, are quite diff erent lar discussions above). Th ere are probably no more from the Albaina acanthomorph fi n spines, which show than two distinct taxa of Enchodontidae indet. (see very faint lateral processes and comparatively bigger above). Consequently, this accounts for a reliable median process (Poyato-Ariza et al. 1999b: fi g. 5C). estimated diversity of 7 taxa, as inferred from these Th e visible ornamentation, consisting of one pair of 10 tooth morphotypes. With the addition of the big, deep basal grooves, is also quite diff erent from the fi n spines, which belong to a totally diff erent taxon small, shallow wavy grooves of the Albaina material. altogether, the whole actinopterygian diversity for Th is means that the corresponding acanthomorph Quintanilla la Ojada is estimated in a minimum forms of both localities, although not identifi able at of 8 diff erent taxa. Th ese correspond to 5 major present, are clearly distinct taxa. Th ere are, then, at taxa, order or higher. Two of these high rank taxa least a couple of signifi cant diff erences at low rank level are non-teleostean ones: corresponding to 3 lower within the general similarity at high rank level. rank taxa, and the other three are teleostean ones, Th e assemblage of actinopterygian remains from corresponding to 5 lower rank taxa. Quintanilla la Ojada, as that of Albaina and other Th e better assemblage comparisons (with the precau- localities of comparable age, shows a larger di- tions explained above) are made with the geographically versity and greater abundance of teleostean taxa, and temporally close locality of Albaina (Poyato-Ariza within the mixture of primitive, non-teleostean et al. 1999b). Th e higher rank taxa present in both and derived teleostean taxa that is characteristic to localities are the same (with the exception of the all of them. Th e most reliable ecomorphotype for Amiiformes, only known in Quintanilla la Ojada), ichthyofaunal comparisons is the durophagous one. although this is obviously a refl ection of their similar Th e teeth with this function are heavy, stout, dense, taphonomic bias, in addition to their obvious ichthyo- and extremely resistant, and are consequently the faunal similarities. Th at is, they have the same high best, more consistently, and more abundantly pre- rank taxa in common, but other actinopterygians that served in this kind of isolated-remains assemblages certainly must have formed part of the correspond- biased by strong, diff erential transport during the ing original communities (e.g., small teleosts with biostratinomic and/or fossil-diagenetic processes. In villiform or absent teeth), are not preserved due to Quintanilla la Ojada, this ecomorphotype includes destruction during taphonomic processes. But as far as both non-teleosts (pycnodonts: morphotypes 1 and they can be compared with the necessary precautions, 4) and teleosts (phyllodonts: morphotypes 6 and the actinopterygian assemblages from Quintanilla la 7), but teleosts (especially Paralbulinae) are largely Ojada and Albaina are very similar in composition, dominant in number, by far, over non-teleosts.

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Th is is interesting because Cretaceous ichthyofaunas TABLE 1. — Actinopterygian taxonomic distribution from the Maas- trichtian localities of Albaina (Miranda-Treviño Synclinorium) and from the Iberian area, in general, are characterized by Quintanilla la Ojada (Villarcayo Synclinorium). The table indicates this mixture of non-teleostean forms, mostly relicts the number of specimens (mainly isolated teeth) by taxon and/or from Jurassic times, and teleostean forms, which are morphotype. For more details about the actinopterygian fauna of Albaina see Poyato-Ariza et al. (1999b). in the process of replacing them. Th is replacement is especially remarkable and intense during the Late Cretaceous, by the end of which teleosts are experi- Quintanilla encing the huge evolutionary radiation that would la Ojada Albaina eventually make them the dominating fi sh forms, by Pycnodontiformes indet. 8 15 far, in any aquatic environment from this time on. Pycnodontoidei indet. A 1 Pycnodonts, which are the dominating durophagous Pycnodontoidei indet. B 14 Pycnodontoidea indet. 9 actinopterygians during the Jurassic and the Early (morphotype 1) Cretaceous, saw their geographic distribution pro- Pycnodontoidea indet. 3 gressively reduced during the Late Cretaceous, and (morphotype 2) became extinct after the Eocene/Oligocene, leaving no Cf. Anomoeodus sp. 31 known relatives (see Poyato-Ariza 2005a for further Cf. Paramicrodon sp. 29 details). Semionotiformes, another group that includes Cf. Amiidae indet. 11 typically durophagous forms, consistently present in Cf. Elopiformes indet. 2 Early Cretaceous fi sh communities, are totally absent Phyllodontinae indet. 3 in Quintanilla la Ojada (and Albaina, for that matter). Phyllodus sp. 6 Th e actinopterygian remains of Quintanilla la Ojada, Paralbulinae indet ~120 with their mixture of dominating teleostean forms Paralbula sp. 559 plus relict, non-teleostean forms, is a nice example of Enchodontidae indet. 28 this in-process ichthyofaunal replacement that would (morphotype A) Enchodontidae indet. 5 radically change the actinopterygian composition of (morphotype B) the aquatic ecosystems in the Iberian area during the Enchodontidae indet. 8 Late Cretaceous. (morphotype C) Enchodus sp. A 15 Enchodus sp. B 146 Acanthomorpha indet. 7 8 CONCLUSIONS Cf. “Stephanodus” sp. 2 Total specimens 205 798 Brackish vertebrate fossil remains, most of them iso- lated teeth, found in the latest Cretaceous of Quin- tanilla la Ojada (Burgos, northern Spain) correspond actinopterygian-bearing beds of Quintanilla la Ojada to actinopterygians, selachians, and marine reptiles. are closely related to a transgressive lag at the base of Actinopterygians are mainly represented in number of the Valdenoceda Formation, which is attributed to specimens by teleosts (paralbulines, enchodontids, and the Lower Maastrichtian to basal Upper Maastrichtian in lesser extent phyllodontines and acanthomorphs), on the basis of the regional stratigraphy. Th e Albaina but pycnodonts and amiids are also present. Diff er- unit is laterally equivalent to the Torme Formation, ent tooth morphotypes (i.e. molariform, incisiform, which is considered to be Upper but not uppermost branchial, conic-styliform, fl attened and globular Maastrichtian on the basis of the selachian association. teeth) are known. Th e actino pterygian diversity for Th ese Cretaceous Iberian ichthyofaunas are charac- this locality is estimated in a minimum of 8 diff erent terized by a mixture of dominating teleosteans and taxa. Th e fossil association of Quintanilla la Ojada relict, non-teleostean forms (such as pycnodonts), (Villarcayo Synclinorium) is similar to that discov- and constitute an example of replacement process ered in Albaina (Miranda-Treviño Synclinorium), of the actinopterygian composition in the aquatic both located in the Basque-Cantabrian Region. Th e ecosystems during the Latest Cretaceous.

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Acknowledgements BRITO P. 1997. — Révision des Aspidorhynchidae (Pisces, We are grateful to Marc Floquet for his help and sup- Actinopterygii) du Mésozoïque : ostéologie, relations port, Mikel López-Horgue for useful comments and phylogénétiques, données environnementales et bio- géographiques. Geodiversitas 19 (4): 681-772. to Eneko Iriarte for fi nding the Quintanilla la Ojada BUFFETAUT É., COSTA G., LE LOEUFF J., MARTIN M., site and fi eld assistance. We also thank Lionel Cavin RAGE J.-C., VALENTIN X. & TONG H. 1996. — An for his valuable contribution, which has improved Early Campanian vertebrate faune from the Villeveyrac the manuscript, and an anonymous referee. Th is Basin (Hérault, southern France). Neues Jahrbuch für paper is a contribution to the projects CGL2005- Geologie und Paläontologie, Monatshefte 1: 1-16. BUSCALIONI A., FREGENAL M. A., BRAVO A., POYATO- 01121 (FJPA), CGL2004-02338, CGL2007-64061/ ARIZA F. J., SANCHÍZ B., BÁEZ A. M., CAMBRA MOO O., BTE and CGL2010-18851/BTE (AB, XPS) of the MARTÍN CLOSAS C., EVANS S. E. & MARUGÁN LOBÓN Ministerio de Educación y Ciencia (currently Min- D. J. 2008. — Th e vertebrate assemblage of Buenache isterio de Ciencia e Innovación) of Spain, and to de la Sierra (Upper Barremian of Serrania de Cuenca, the research project GIC07/14-361 and IT-320-10 Spain) with insights into its taphonomy and palaeo- ecology. Cretaceous Research 29: 687-710. of the Gobierno Vasco/EJ (AB, XPS). CAPPETTA H. & CORRAL J. C. 1999. — Upper Maas- trichtian selachians from the Condado de Treviño (Basque-Cantabrian region, Iberian Peninsula). Estudios REFERENCES del Museo de Ciencias Naturales de Álava 14 (número especial 1): 339-372. BACETA J. I. 1996. — El Maastrichtiense superior, Paleo- CAVIN L. 1999. — Osteichthyes from the Upper Cre- ceno e Ilerdiense basal de la Región Vasco-Cantábrica: taceous of Laño (Iberian Peninsula). 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Submitted on 12 December 2008; accepted on 24 August 2010.

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