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A revision of the Lari (Aves, Charadriiformes) from the early Miocene of Saint-Grand-le-Puy (Allier, France)
Vanesa L. De Pietri , Loc Costeur , Marcel Gntert & Gerald Mayr
a a b a c

Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, CH-3005, Bern, Switzerland
b c

Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001, Basel, Switzerland

Forschungsinstitut Senckenberg, Division of Ornithology, Senckenberganlage 25, D-60325, Frankfurt am Main, Germany Available online: 11 Jul 2011

To cite this article: Vanesa L. De Pietri, Loc Costeur, Marcel Gntert & Gerald Mayr (2011): A revision of the Lari (Aves, Charadriiformes) from the early Miocene of Saint-Grand-le-Puy (Allier, France), Journal of Vertebrate Paleontology, 31:4, 812-828 To link to this article: http://dx.doi.org/10.1080/02724634.2011.586663

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Journal of Vertebrate Paleontology 31(4):812828, July 2011 2011 by the Society of Vertebrate Paleontology

ARTICLE

A REVISION OF THE LARI (AVES, CHARADRIIFORMES) FROM THE EARLY MIOCENE OF SAINT-GERAND-LE-PUY (ALLIER, FRANCE)
VANESA L. DE PIETRI,*,1 LOIC COSTEUR,2 MARCEL GUNTERT,1 and GERALD MAYR3 Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, CH-3005 Bern, Switzerland, vanesa.depietri@nmbe.ch; marcel.guentert@nmbe.ch; 2 Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001 Basel, Switzerland, loic.costeur@bs.ch; 3 Forschungsinstitut Senckenberg, Division of Ornithology, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany, gerald.mayr@senckenberg.de

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ABSTRACTThree species of gull-like birds were described by Milne-Edwards in the 19th century from the early Miocene of Saint-Gerand-le-Puy, France, two of which have been since then moved to the genus Laricola. These fossils are redescribed and revised in the present study. We further describe two new species of the taxon Laricola, L. intermedia, sp. nov., and L. robusta, sp. nov., as well as two species of a new taxon Sternalara, S. minuta and S. milneedwardsi, thus substantially increasing the number of charadriiform taxa known from this locality. To nd out how Laricola relates to modern taxa, we performed a phylogenetic analysis of 41 osteological characters, which resulted in Laricola being placed outside of Lariidae, as early representatives of the newly designated group Laromorphae, which encompasses terns, gulls, and skimmers. The results of our analysis have additionally enabled us to evaluate relationships between extant taxa of the Laromorphae.

INTRODUCTION Fossil gull-like birds (Charadriiformes) from the early Miocene (MN2) locality of Saint-Gerand-le-Puy in France were initially described by Milne-Edwards (1863, 18671868), who considered them to be members of the modern genus of gulls Larus. Currently, little is known about the phylogenetic afnities of these very abundant Miocene birds, though they can clearly be placed within the Lari, which is one of the three clades that, according to recent analyses of nuclear and mitochondrial sequence data (e.g., Ericson et al., 2003; Paton et al., 2003; Baker et al., 2007; Fain and Houde, 2007), make up the order Charadriiformes, the other two being Charadrii (plovers and allies) and Scolopaci (sandpipers and allies). The Lari comprise the Dromadidae (crab plover), Stercorariidae (skuas), Alcidae (auks), Laridae (gulls), Sternidae (terns), Rynchopidae (skimmers), and Glareolidae (pratincoles and coursers), with the Turnicidae (buttonquails) being the sister group to the whole clade (Ericson et al., 2003; Paton et al., 2003; Baker et al., 2007; Fain and Houde, 2007; Mayr, 2011). Although phylogenetic afnities within this group are largely consistent throughout the above-cited molecular studies, there is some disparity concerning some families. For instance, the position of Rynchopidae is variable in both molecular and morphological studies (Schnell, 1970; Ericson et al., 2003; Baker et al., 2007; Livezey and Zusi, 2007; Fain and Houde, 2007; Mayr, 2011) and it is thus uncertain whether skimmers are most closely related to terns, gulls, or are sister group of both. Also, recent research (e.g., Baker et al., 2007; see also Odeen et al., 2010) suggests that the Sternidae are a paraphyletic grouping, with Anous (noddies) and Gygis (white terns) being sister taxa to a clade including all other Sternidae, Laridae, and Rynchopidae. So far, no morphological assessment of the relationships within Lari has included any fossil representatives of this group.

*Corresponding

author.

The Paleogene fossil record of members of the Lari is poor (Mayr, 2009). Noteworthy among these fossil specimens are stem-group representatives of the Turnicidae from the Oligocene of France and Germany (Mayr, 2000; Mayr and Knopf, 2007). Fossils assigned to the Glareolidae were recovered from the early Oligocene of Belgium (Mayr and Smith, 2001). The purported occurrence of Laridae in the Eo-Oligocene deposits of Query, France (Mourer-Chauvire, 1982) has been withdrawn by Mourer Chauvire (2006). The Miocene record of the Lari is mostly European. Glareolidae are represented by Mioglareola gregaria and Glareola neogena from the early and middle Miocene of Germany, respectively (Ballmann, 1979), whereas Paractiornis is known from the early Miocene (MN3) of Nebraska (Wetmore, 1930; Olson and Steadman, 1978). Denite evidence of true Laridae having existed during the Miocene has not been convincingly presented yet. Larus teruelensis from the late Miocene of Mansuetos, Spain, is known from the distal end of a humerus and was originally identied as a scolopacid (Villalta, 1963). It was reassigned to the Laridae by Mlkovsky (2002). Larus dolnicensis from the early Miocene of Dolnice (MN4b), Czech Republic, is represented by a fragmentary distal end of a humerus (Svec, 1980), and was later allocated to the genus Mioglareola by Mlkovsky (2000). The existence of bones assignable to either Laridae or Sternidae, also from the early Miocene of Dolnice, was mentioned by Mlkovsky (1996), but no further information was provided on these specimens. In North America, Gaviota niobrara, from the late Miocene of Nebraska, is also known from the distal end of a humerus (Miller and Sibley, 1941). An alleged species of Larus from the middle Miocene of Lee Creek Mine, North Carolina, was described by Olson and Rasmussen (2001), but is only represented by the scapular half of a coracoid. The presence of some gull-like premaxillae from the earlymiddle Miocene (1619 Ma) of St. Bathans, New Zealand, was mentioned by Worthy et al. (2007). All of these Miocene fossils are too incomplete to be condently identied at genusor even familylevel, and may indeed be more basal than assumed.

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DE PIETRI ET AL.EARLY MIOCENE LARI FROM CENTRAL FRANCE The late Oligocene/early Miocene localities of Saint-Gerandle-Puy in France have yielded many thousands of avian specimens belonging to over 40 genera (see Cheneval, 1996), and certainly rank high in importance concerning early Miocene faunas worldwide. Due to its lacustrine deposits, the Saint-Gerandle-Puy area has produced an avifauna consisting primarily of taxa with a preference for aquatic habitats (e.g., Cheneval, 1984, 1996). Regarding charadriiforms, only members of the Scolopacidae and the Lari have been recorded from this location. The Lari of Saint-Gerand-le-Puy were originally described by Milne-Edwards (1863, 18671868) as Larus elegans, L. totanoides, and the larger L. desnoyersii. Because L. elegans and L. totanoides areaccording to Milne-Edwardss (18671868:360361) measurementssignicantly different in hind limb proportions from extant Laridae and Sternidae, Mlkovsky (2002) proposed to unite them under the genus Laricola, whereas he considered Larus desnoyersii as genus incertae sedis. Although it had already been suggested that these birds may have been too primitive for inclusion in the Laridae (Ballmann, 1976), until now it has not been determined whether this distinction is supported by derived osteological features. Laricola elegans has been allegedly also recovered from the early Miocene (MN4b) of Dolnice, Czech Republic (Mlkovsky, 2002), and members of Laricola have been reported from late Oligocene (MP 25 to MP 30) deposits in France (Mourer-Chauvire, 1995, 1996; Mourer-Chauvire et al., 2004). The last descriptive assessment of the Saint-Gerand-le-Puy Lari dates back to Alphonse Milne-Edwards in the late 1860s and a revision of these birds is long overdue. The ample avian collec tion from Saint-Gerand-le-Puy in the Natural History Museum in Basel, Switzerland, has presented us with the possibility of reevaluating these alleged larid-like specimens. Despite the vast amount of fossil material, the task of assigning the Saint-Gerand-le-Puy Lari a phylogenetic placement based on derived osteological traits has remained elusive, and only recently has progress been made towards reconciling the great number of phylogenetic hypotheses, based both on molecular and morphological data, proposed during the last 30 years for representatives of the Lari, among other Charadriiformes (Mayr, 2011). Here we present the rst attempt to include fossil Laricola from Saint Gerand-le-Puy in a phylogenetic context within the Lari, as well as to provide a descriptive overview and present state of knowledge for these fossils. We will further address the issue of the paraphyly of the Sternidae and the conicting position of Rynchopidae. Institutional AbbreviationsMNHN, Museum National dHistoire naturelle, Paris, France; NMB, Natural History Museum Basel, Switzerland; SMF, Forschungsinstitut Senckenberg, Frankfurt am Main, Germany. GEOLOGY The fossil site of Saint-Gerand-le-Puy (Fig. 1) is actually composed of several quarries in the vicinity of the villages of Saint-Gerand-le-Puy and Montaigu-le-Blin in central France (e.g., Cheneval, 1996). The old collections of the Natural History Museum Basel (together with others in Paris and Lyon) were gathered in the 19th and early 20th centuries and no direct relationship between the nds and their sedimentological context was ever recorded so that the exact provenance of the fossils is difcult to establish. This is not much of a problem for large taxonomic assessments, because the different quarries in the vicinity of both villages yielded Aquitanian faunas that were often ascribed to the MN2 (22.520.5 Ma; Steininger, 1999) biozone of the European Land Mammal Ages (Hugueney, 1974). Some localities of the Saint-Gerand-le-Puy area were neverthe less determined to be of late Oligocene age, such as Billy-Crechy,

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FIGURE 1. Geographic position of Saint-Gerand-le-Puy in the French Allier Region. The light grey zone shows the Aquitanian deposits (after Wattine et al., 2003). The region is mapped onto the palaeogeography of the Early Miocene (after Meulenkamp and Sissingh, 2003).

in which the lower layers date from the late Oligocene MP2930 (Hugueney et al., 2003). The exact stratigraphic provenance of the fossils in the old collections is not known in all cases. We cannot condently refer the studied material to any late Oligocene locality, and a late Oligocene age for some of the material is a possibility that cannot be excluded. Comprehensive sedimentological studies described the geology of the largest quarry at Montaigu-le-Blin (Bucher et al., 1985) or the formation of the lacustrine calcareous deposits of the area (Donsimoni, 1975; Wattine et al., 2003). The general geological context of the area (the Limagne sedimentary basin) is that of a subsident basin related to alpine orogeny such as many other Eocene and Oligocene rift-basins throughout Europe (the European-Rift System; Meulenkamp and Sissingh, 2003). The peculiarity of the early Miocene of the Limagne basin is the widespread calcareous sedimentation in a uvio-lacustrine context, associated with stromatolitic bioconstructions. For Montaigu-le-Blin, Bucher et al. (1985) suggested that most of the fossil accumulations were found within marls deposited as mud ows between the bioconstructions. These marls also show high concentrations of freshwater gastropods, attesting their lacustrine origin (Helicidae: Bucher et al., 1985; Wattine et al., 2003). The sedimentological succession in Montaigu-le-Blin shows regressive sequences of the lake level inducing facies changesfrom marls to sands and conglomeratesshowing an increasing inuence of the river system. Widely developed stromatolites often cap these sequences, testifying to the return of true lacustrine conditions (Donsimoni, 1975; Donsimoni and Giot, 1977; Wattine et al., 2003). Other fossil accumulations

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 4, 2011 SYSTEMATIC PALEONTOLOGY AVES Linnaeus, 1758 CHARADRIIFORMES Huxley, 1867 LAROMORPHAE, new taxon DiagnosisLeast inclusive clade including Laridae, Sternidae (including Anous and Gygis), Rynchopidae, and extinct Laricolidae, n. fam., but excluding Stercorariidae. Characterized by: marked impressio coracobrachialis (humerus, also in Stercorariidae); pronounced fossa musculi brachialis (humerus, except Anous). Differing from Stercorariidae in: presence of fossa caudalis (mandible); subovate facies articularis acrocoracoidea (furcula) extending over nearly entire width of scapus claviculae. LARICOLIDAE, n. fam. Type GenusLaricola Mlkovsky, 2002. DiagnosisSmall- to medium-sized members of Laromorphae, with proportionally longer and more slender legs than all extant representatives (Fig. 2). Further differing from other Laromorphae in: shallow facies articularis acrocoracoidea (furcula). Differing from Sternidae s.s., Laridae, and Rynchopidae in: wide and deeply concave trough-like processus maxillopalatinus (skull); absence of well-developed fenestra caudalis (mandible); absence of second fossa pneumotricipitalis (humerus). LARICOLA Mlkovsky, 2002 (Figs. 35) Type SpeciesLaricola elegans (Milne-Edwards, 1868). Included SpeciesLaricola elegans (Milne-Edwards, 1868); L. totanoides (Milne-Edwards, 1868); L. desnoyersii (MilneEdwards, 1863); L. intermedia, sp. nov.; L. robusta, sp. nov. RemarksMilne-Edwards (18671868) described Laricola based on comparisons with few extant Charadriiformes, most from the family Laridae. We present a more complete description of the fossil genus comparing it to other Laromorphae, and document previously overlooked anatomical details. The genus Laricola presently encompasses two species, Laricola elegans and L. totanoides (Mlkovsky, 2002). In this study we have identied two additional species, and further transferred Larus desnoyersii into the taxon Laricola. Anatomical differences within Laricola are, for the most part, of allometric nature. Many postcranial bones are only known for the very abundant Laricola elegans and, unless mentioned otherwise, we base the genus description on this species. L. elegans is among the most abundant fossil birds of Saint Gerand-le-Puy, so that most skeletal elements can be assigned to this species by their sheer number and gull-like morphology. For the same reasons and unless there existed a conict in size, we considered it most likely that other gull-like bones that were represented by smaller numbers (e.g., pelvis) belong to this abundant species rather than one of the rarer species present in the material. Description and Comparisons SkullThe skull of Laricola has so far only been incompletely known (Milne-Edwards, 18671868:pl. 57, g. 1), with several features in need of a thorough description. In our material there are two skulls, an exceptionally well-preserved and almost complete one (NMB s.g.18810; Fig. 3A, EF, H) and a cranium (NMB s.g.18812; Fig. 3P, RT), which differ marginally in size and minor morphological features. Overall, the complete skull resembles Larus minutus in proportions and shape. The beak is straight and intermediate between L. minutus and Sterna paradisaea. The tip, contrary to the condition found in Laridae, is barely decurved, though not as straight as in Sternidae s.s., reecting best the condition found in Anous stolidus. The distal portion of the

within uvio-deltaic calcareous sediments (also rich in freshwater gastropods: Valvatidae and Hydrobiidae to a lesser extent; see Wattine et al., 2003) seem to have preserved only aquatic birds, as far as vertebrates are concerned (Bucher et al., 1985) and again indicate freshwater lacustrine conditions. Overall, both the sedimentology and palaeontology at Saint Gerand-le-Puy testify to shallow freshwater conditions under varying climatic conditions with more or less humid periods changing the chemical conditions in the lake (with periods of low or high oxygenation). These changes are in turn responsible for the paleontological content (gastropods, ostracods, etc.; Wattine et al., 2003) of the different geologic formations. Bird fossils occur in different formations, either trapped in the marls between the bioconstructions (and in this case, aquatic, shore, or terrestrial species are known) or in the uvio-deltaic calcareous sediments (and here, only aquatic species are known). MATERIALS AND METHODS Materials

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The fossil specimens are deposited in NMB and, unless otherwise stated, the description of the material is based on this collection. Additional fossil material was consulted at MNHN. The following skeletons of recent Charadriiformes were available for comparisons. Sternidae: Thalasseus bergii (skull; NMB), Chlidonias niger (skull; NMB), Sterna paradisaea (NMB and SMF), S. saundersi (SMF), Anous stolidus (NMB and SMF), and Larosterna inca (SMF). Stercorariidae: Stercorarius parasiticus (NMB) and Catharacta skua (NMB). Laridae: Xema sabini (SMF), Rhodostetia rosea (SMF), Larus canus (NMB), L. argentatus (NMB), L. fuscus (NMB), L. hyperboreus (NMB), L. marinus (NMB), L. ridibundus (NMB), and Rissa tridactyla (NMB). Alcidae: Fratercula arctica (NMB), Alca torda (NMB), Uria lomvia (NMB), Alle alle (NMB), and Synthliboramphus antiquus (SMF). Glareolidae: Glareola pratincola (SMF). Rynchopidae: Rynchops niger (SMF). Charadriidae: Pluvialis apricaria (NMB) and Vanellus vanellus (NMB). Pluvianidae: Pluvianus aegyptius (NMB). Scolopacidae: Arenaria interpres (NMB), Philomachus pugnax (NMB), Scolopax rusticola (NMB), Numenius arquata (NMB), Limosa limosa (NMB), and Calidris canutus (NMB). Anatomical terminology follows Baumel and Witmer (1993). To comply with recent ndings from sequence data (see Introduction), Sternidae sensu stricto refers to a taxon excluding Gygis and Anous (see Baker et al., 2007), whereas we have retained the use of Sternidae for the traditional taxon including Anous and Gygis. Phylogenetic Analysis Forty-one characters (Supplementary Data, Appendix 1) were scored for the following genera: Stercorarius, Laricola, Anous, Sterna, Larosterna, Rynchops, Larus, Rissa, Xema, and Rhodostetia, with Limosa and Glareola chosen as outgroups. The matrix (Supplementary Data, Appendix 2) is largely based on Mayr (2011) from which 32 characters were retained and 9 were newly added. All characters were treated as unordered and assigned equal weight. Phylogenetic analyses were performed with the heuristic search modus of NONA 2.0 (Goloboff, 1993) through the Winclada 1.00.08 interface (Nixon, 2002), using the following commands: hold 10000, hold/10, mult 1000, and max . Unsupported nodes were collapsed. Bootstrap values were calculated under the following settings: 1000 replicates, with three searches holding one tree per replicate, and TBR branch swapping without max . Unless otherwise stated (see Results), characters were optimized using the unambiguous option in Winclada. Bremer supports were calculated in TNT 1.1 (Goloboff et al., 2003, 2008) using 10,000 replicates and retaining suboptimal trees by 20 steps.

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FIGURE 2. Simpson diagram showing the log differences of the greatest lengths (in mm) of some postcranial bones of the extinct Laricola elegans and the extant taxa Stercorarius parasiticus, Anous stolidus, Sterna paradisaea, Larosterna inca, Larus ridibundus, Rissa tridactyla, and Larus minutus. Anous stolidus was chosen as the standard taxon. The log difference for each bone refers to: log (measurement of compared taxon) log (measurement of standard taxon). Measurements for Laricola elegans are averaged. For a more detailed explanation of such diagrams, see Gohlich and Mourer-Chauvire (2005). Note the proportionally longer legs of Laricola elegans. Abbreviations: CMC, carpometacarpus; TBT, tibiotarsus, TMT, tarsometatarsus.

beak is not as narrow as in Sternidae. The narial openings are only slightly longer than those of S. paradisaea. These tend to be shorter in Sternidae, with a proportionally longer symphysis of the rostrum maxillare, whereas they are very long, extending throughout most of the length of the beak in Laridae and Stercorariidae (Fig. 3FG). Unlike in Laridae, the fossae glandularum nasales are shallow. The os lacrimale (Fig. 3A) is not as wide as in Sternidae s.s. and Rynchopidae and is thus in agreement with the condition in Laridae and Anous. The small anterior process (see Cracraft, 1968) that is characteristic of the Sternidae is not present in this specimen, although due to its fragility it is unlikely that it would have been preserved. The os ectethmoidale of NMB s.g.18810 (Fig. 3M) has a distinctive shape with a pointed tip and concave ventral border, matching the condition in A. stolidus and other terns (e.g., S. paradisaea). In NMB s.g.18812 (Fig. 3P), the ectethmoid is broader and lacks the concave ventral margin, as in Stercorariidae. The processus maxillopalatinus forms a wide and deep, laterally concave trough, as in A. stolidus and some species of the Stercorariidae (Fig. 3HJ). In Laricola, the maxillopalatine process is divided by a relatively wide bar of bone connecting its dorsal and ventral margins, and a similar condition is found in Stercorarius parasiticus (Fig. 3J). In Anous (Fig. 3I) the maxillopalatine process is laterally closed except for a round caudal foramen. In Sternidae S.S., Laridae, and Rynchopidae, the overall shape of the maxillopalatine process is less like that of a deep trough, being much atter. In Laridae (Fig. 3L), the maxillopalatine process is widely open with one or more thin bars of bone extending

along the bottom of the trough. In Sternidae and Rynchopidae, it is fenestrated by thin bars of bone (Fig. 3K). In NMB s.g.18810, the processus postorbitales are worn, but they show no indication of having been well developed. In NMB s.g.18812, they project laterally and caudally and are about the size of those of Larus minutus, albeit broader (Fig. 3RT). A maxillopalatine strut A in the sense of Zusi and Jehl (1970) is absent in Laricola. The processus laterales parasphenoidales (Fig. 3T) are well developed and ventrally protruding, and the processus suprameaticus is not longer than the processus zygomaticus (Mayr, 2011), both features in accordance with the condition in Laridae, Rynchopidae, and Sternidae. As in other Lari except the Turnicidae, processus basipterygoidei are absent. The fossae temporales are well marked, although not as enhanced as in some terns (e.g., Thalasseus bergii and S. paradisaea) and Laridae (e.g., Larus ridibundus). In these taxa, they almost reach the midline, but in the fossil they only reach over halfway, most closely resembling the morphology in Larus minutus. A fragment of a mandible (Fig. 3VW) assignable to Laricola elegans (MNHN Av.4847) is present in the MNHN collection. It lacks the well-developed fenestra caudalis, a condition shared with stercorariids and Anous, which indicates that its presence in gulls, skimmers, and some terns is plesiomorphic for Laromorphae. Also, a fossa caudalis, which is absent in skuas, is present in Laromorphae. VertebraeVertebrae of Laricola have not been previously described. In our material, only one cervical vertebra is likely to belong in Laricola, and does not differ from those within

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DE PIETRI ET AL.EARLY MIOCENE LARI FROM CENTRAL FRANCE Laromorphae. A couple of well-preserved thoracic vertebrae are also present within the material, which are indeed opisthocoelous and pleurocoelous. Opisthocoelous thoracic vertebrae are also present in Alcidae, Stercorariidae, and Laromorphae, whereas pleurocoelous vertebrae are absent in Stercorariidae and Alcidae (e.g., Mayr, 2011). CoracoidOur material includes numerous complete coracoids, two of which possess an intact processus lateralis (Fig. 5AB). In Laricola elegans and L. totanoides, this bone bears a close resemblance to the coracoid of the small gull Larus minutus. The pneumatic recess under the facies articularis clavicularis, characteristic of most gulls (Fig. 5E), is neither present in Laricola elegans nor L. totanoides. A coracoid of an alleged Miocene gull lacking this feature was described by Olson and Rasmussen (2001), who hypothesized that this might be a plesiomorphic trait of gulls. Indeed, this pneumaticity is absent from most, but not all, specimens of Laricola, and is thus a variable trait (it is present in Laricola desnoyersii, for example). FurculaIn Alcidae, Sternidae, Rynchopidae, and Laridae, the extremitas omalis of the furcula exhibits a distinct subovate facies articularis acrocoracoidea, almost extending over the entire width of the scapus claviculae (Mayr, 2011). This feature is also present in Laricola (Fig. 5GH). Nonetheless, the articulation facet is much shallower in the fossil than in the examined recent species, which is likely to be the plesiomorphic condition for Laromorphae. The scapula of Laricola matches thoroughly that of recent Laromorphae. SternumThere are several fragments of the sternum of Laricola present in the collection. Among these is a corpus sterni with a preserved rostral portion, whose preservation is so poor that barely any details are recognizable. One rostral portion of the sternum clearly shows the shape of the apex carinae (Fig. 5L), which is pointed and cranially projecting, as in Anous and some Laridae. HumerusA large number of complete and partial humeri in the collection can be referred to Laricola. A second fossa pneumotricipitalis is absent (Fig. 4A, C, H, K, N), a condition that within Laromorphae is only also present in Anous (Fig. 4X). Strauch (1978) mentioned that the absence of a second fossa pneumotricipitalis is a variable trait both in Anous and Rynchops, but we are unable to support this statement. The sulcus nervi coracobrachialis is very well marked in Laricola, a trait that is variable within Laridae and Sternidae. In agreement with recent Laridae, the fossa musculi brachialis is deep (Fig. 4D, F, I, J, O). Within Laromorphae, this fossa is shallow only in Anous. In Stercorariidae, Laricola, and Anous, there is a well-marked tubercle on the cranial surface of the distal end, between the base of the tuberculum supracondylare dorsale and the fossa m. brachialis (Fig. 4V), for the attachment of m. supinator, which is less marked in the rest of the Laromorphae. The humerus of Laricola otherwise matches that of other Laromorphae (for more details see Mayr [2011], Chu [1998], and Strauch [1978]). UlnaUlnae are fairly uniform throughout Laromorphae. The tuberculum ligamenti collateralis ventralis (Fig. 5M) is well developed throughout this group, but in most Laridae (except Larus hyperboreus) and Laricola it is more pronounced ventrally

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than in the examined Anous and Stercorariidae; in both these taxa the tuberculum is also slightly more elongated. In Rynchops, the tuberculum ligamenti collateralis ventralis is not developed at all. CarpometacarpusUnlike in recent Laridae, the carpometacarpus of Laricola bears a small processus intermetacarpalis (Fig. 5U). Otherwise this bone does not differ from that of Laromorphae. Milne-Edwards (18671868:354) mentioned that the spatium intermetacarpale is narrower in Laricola and Laridae than in terns. However, this is true only for very small terns, such as Sterna saundersi, and is probably a size related effect. Other Wing BonesThe radius of Laricola shows no peculiarities. In agreement with the condition found in Laromorphae and Stercorariidae, the phalanx proximalis digiti majoris is bifenestrated (Fig. 5VX). PelvisAlthough no complete pelvis is known from the NMB collection (Fig. 5ZAA), a specimen illustrated by MilneEdwards (18671868:pl. 56, g. 22) shows that the pelvis of Laricola possesses two rows of foramina transversaria, and thus matches that of smaller Laridae, such as Larus minutus, Xema sabini, and some Sternidae s.s. The one well-preserved partial pelvis present in the NMB collection shows that the recessus caudalis fossae (Fig. 5Y) is present in Laricola, and this recess is also present in Laridae and Anous stolidus (contra Strauch, 1978). It is absent in Sternidae (except Sterna caspia) and Rynchopidae. FemurThe femur of Laricola (Fig. 5BBCC) agrees with that of extant Laridae and Sternidae. There is a prominence just proximal of the condylus medialis (Fig. 5CC), which is also found in Sternidae and Rynchopidae, and is variable within Laridae, being present in some small species such as Larus minutus, but absent in Stercorarius and Anous. However, this prominence is developed as a ridge in Laricola and Laridae, whereas it is indeed a prominent tubercle in Sternidae and Rynchopidae. The tuberculum musculi gastrocnemialis lateralis is well developed, as in Laromorphae. TibiotarsusThe legs of Laricola are long and slender, longer than in all examined recent Laromorphae (Fig. 2). The tibiotarsi of Laromorphae and Stercorariidae are very uniform in appearance, and that of Laricola matches this morphology (Fig. 5DDFF). TarsometatarsusThe hypotarsus of Laricola exhibits a very prominent crista medialis hypotarsi and two distinct cristae intermediae hypotarsi (Fig. 5KKNN). This conguration agrees with that of Laridae, although the shape of the hypotarsus is very variable within Laromorphae, and some variation is also observed in Laricola. In Sternidae this sulcus is actually a canal, and some terns such as Thalasseus bergii and Anous stolidus have two closed canals, although given the current phylogeny of the group (Baker et al., 2007), these are unlikely to be homologous. Furthermore, the hypotarsus of the tern Larosterna resembles that of gulls and Laricola. One tarsometatarsus (Fig. 5NN) assigned to Laricola elegans bears a closed canal, but unlike in the Sternidae, in which the canal is centrally positioned, the orientation matches that of the sulcus in other Laricola specimens. The closed canal could thus be the result of ossication of cartilage that closes the

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FIGURE 3. Skulls and mandible of Laricola elegans in comparison to extant Lari. AD, dorsal view of Laricola elegans (A, NMB s.g.18810), Anous stolidus (B), Sterna paradisaea (C), and Larus canus (D). E, ventral view of the skull of Laricola elegans (NMB s.g.18810). FG, right lateral view of the skulls of Laricola elegans (F, NMB s.g.18810) and Larus canus (G). HQ, not to scale, details of the skull of Laricola elegans (H, M, NMB s.g.18810; P, NMB s.g.18812) in comparison to Anous stolidus (IN), Stercorarius parasiticus (JO), Sterna paradisaea (K), and Larus canus (LQ). RS, skull of Laricola elegans (NMB s.g.18812) in dorsal (R) and ventral (S) views. TU, occipital view of the skulls of Laricola elegans (T, NMB s.g.18812) and Larus canus (U). VW, fragment of right ramus mandibulae of Laricola elegans (MNHN Av.4847) in medial (V) and lateral (W) views. X, right ramus mandibulae of Larus canus in lateral view. Abbreviations: fgn, fossa glandulae nasalis; fm, fenestra mandibulae; ft, fossa temporalis; oe, os ectethmoidale; ol, os lacrimale; plp, processus lateralis parasphenoidales; pmp, processus maxillopalatinus; ppo, processus postorbitales; srm, symphysis rostrum maxillare.

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FIGURE 4. Humeri of the ve species of Laricola (AP) and the two species of Sternalara (QU) in comparison to extant Lari (VAA). AD, left humerus of Laricola elegans (NMB s.g.18878) in caudal view (A), cranial view (B), proximal end in caudal view (C), and distal end in cranial view (D). EG, left humerus of Laricola totanoides (NMB s.g.2180) in caudal view (E), cranial view (F), and distal end in cranial view (G). HI, right humerus of Laricola robusta, sp. nov. (NMB s.g.18465), in caudal (H) and cranial (I) views. JK, right humerus of Laricola intermedia, sp. nov. (holotype, MNHN SG 13744), in cranial (J) and caudal (K) views. LP, humeri of Laricola desnoyersii, left humerus (MNHN Av.4059) in cranial (L) and caudal (MN) views, and distal left humerus (NMB MA 1106) in cranial (O) and caudal (P) views. QR, right humerus of Sternalara milneedwardsi, sp. nov. (holotype, MNHN SG13742), in cranial (Q) and caudal (R) views. SU, humeri of Sternalara minuta, sp. nov., right humerus (holotype, NMB s.g.12727) in cranial (S) and caudal (T) views, and proximal end of left humerus (NMB M.A.1582) in caudal view (U). VX, right humerus of Anous stolidus in cranial (V) and caudal (WX) views. YZ, right humerus of Stercorarius parasiticus in cranial (Y) and caudal (Z) views. AA, right proximal end of humerus of Sterna paradisaea in caudal view. Abbreviations: ams, attachment for musculus supinator; crbi, crista bicipitalis; crdp, crista deltopectoralis; dfpn, dorsal fossa pneumotricipitalis; fmb, fossa musculi brachialis; fpn, foramen pneumaticum; td, tuberculum dorsale; tsd, tuberculum supracondylaris dorsalis; tsv, tuberculum supracondylare ventrale. Same scale bar for each type of skeletal element.

hypotarsal sulcus. One specimen assignable to Laricola desnoyersii (see below) also possesses this feature (Fig. 5LL). The foramen vasculare distale (Fig. 5II) is situated in a wider and better-marked fossa than in extant Laridae. This fossa is also well marked in the examined Stercorariidae, Sternidae, and Rynchopidae. In Laricola the trochlea metatarsi III projects farther distally than in all other examined Laromorphae, although some gulls (e.g., Larus marinus) do resemble Laricola in this

aspect. Rynchopidae and Sternidae, on the other hand, have a trochlea metatarsi III that does not project much farther distally. LARICOLA ELEGANS (Milne-Edwards, 1868) (Figs. 35, Table 1) Larus elegans Milne-Edwards, 1868:350357, pl. 56, gs. 1129; pl. 57, gs. 111; pl. 58, g. 1, reconstructed skeleton (original description).

DE PIETRI ET AL.EARLY MIOCENE LARI FROM CENTRAL FRANCE Laricola elegans (Milne-Edwards, 1868): Mlkovsky, 2002:136 (new combination). LectotypeMNHN Av.4134 (left humerus). Referred SpecimensAll specimens from NMB: NMB s.g.18810 (skull, with beak), NMB s.g.18812 (cranium), NMB s.g.21782, NMB s.g.1881, and NMB MA.1379 (partial skulls), MNHN Av.4847 (mandible, fragment of right ramus), NMB s.g.16334-49 (vertebrae), NMB s.g.16360-63, NMB s.g.16375, NMB s.g.16379, and NMB s.g.16378 (furculae), NMB s.g.15975, NMB s.g.18868, and NMB s.g.16385-16412 (scapulae), NMB s.g.18816 (left coracoid), NMB s.g.18827 (right coracoid), NMB s.g.13027 (left coracoid), NMB s.g.13030 (right coracoid), NMB s.g.13028 (left coracoid), NMB s.g.10608 (right coracoid), NMB s.g.18824 (left coracoid), NMB s.g.18815 (right coracoid), NMB s.g.18825 (left coracoid), NMB s.g.17023 (left humerus), NMB s.g.18885 (right humerus), NMB s.g.21935 (left humerus), NMB M.A.1091 (right humerus), NMB s.g.21936 (left humerus), NMB s.g.18878 (left humerus), NMB s.g.21933 (left humerus), NMB s.g.18987 (right ulna), NMB s.g.18983 (left ulna), NMB s.g.18995 (right ulna), NMB s.g.12951 (right ulna), NMB s.g.16139 (left ulna), NMB s.g.16579 (left ulna), NMB s.g.15913 (right ulna), NMB s.g.15665 (left carpometacarpus), NMB MA 3178 (left carpometacarpus), NMB s.g.16183 (left carpometacarpus), NMB M.A.2589 (left carpometacarpus), NMB M.A.2599 (left carpometacarpus), phalanx proximalis digiti majoris (NMB s.g.16768, NMB M.A.888), pelvis (NMB s.g.18814, NMB s.g.16328, NMB s.g.16329, NMB s.g.16330, NMB s.g.7516), NMB Ph.3537(left femur), NMB s.g.19038 (right femur), NMB s.g.16806 (left femur), NMB s.g.19127 (left tibiotarsus), NMB s.g.19164 (left tibiotarsus), NMB M.A.3144 (right tibiotarsus), NMB s.g.19127 (left tibiotarsus), NMB s.g.7331 (left tibiotarsus), NMB s.g.19138 (right tibiotarsus), NMB Ph.2073 (right tibiotarsus), NMB s.g.13223 (left tarsometatarsus), NMB s.g.19184 (left tarsometatarsus), NMB s.g.19186 (left tarsometatarsus), NMB s.g.13220 (right tarsometatarsus), NMB s.g.16904 (left tarsometatarsus). Locality and AgeAntoingt (late Oligocene, MP 25), France (Mlkovsky, 2002); Gannat (late Oligocene, MP 30), France (Mlkovsky, 2002); Saint-Gerand-le-Puy (late Oligocene?early Miocene, MN2), Allier, France. MeasurementsSkull, NMB s.g.18810: 61 mm long; maximum width excluding processus postorbitales: 17.24 mm; height including processus laterales parasphenoidales: 17.0 mm. Cranium, NMB s.g.18812: 30.36 mm long (as is; Fig. 3); maximum width excluding processus postorbitales: 17.90 mm; height including processus laterales parasphenoidales: 16.1 mm. For postcranial elements see Table 1. RemarksHundreds of postcranial bones are assignable to Laricola elegans, in the collections of both MNHN and NMB. These constitute a continuum varying greatly in size (Table 1), but show no obvious anatomical differences. Based on the postcranial skeleton alone it is impossible to discern between potential species within these ranges. Because of the large amount of fossil material (hundreds of bones), we have referred only to the best-preserved examples. From the syntype series in MNHN, we chose MNHN Av.4134, a left humerus, as lectotype. This bone is the one likely to have been gured by Milne-Edwards (1868:pl. 57, gs. 23). LARICOLA TOTANOIDES (Milne-Edwards, 1868) (Figs. 45, Table 1) Larus totanoides Milne-Edwards, 1868:358361, pl. 57, gs. 1217 (original description). Laricola totanoides (Milne-Edwards, 1868): Mlkovsky, 2002:137 (new combination).

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LectotypeMNHN Av.4004 (right humerus). Illustrated by Milne-Edwards, 18671868:pl. 57, gs. 1617. Referred SpecimensMNHN Av.9455 (right coracoid), NMB s.g18809 (right coracoid), NMB s.g.2180 (left humerus), NMB s.g.17047 (distal right humerus), NMB s.g.12946 (left ulna), NMB s.g.18994 (left ulna), NMB s.g.12947 (right ulna), NMB M.A.1602 (right ulna), NMB M.A.5181 (right carpometacarpus), NMB s.g.19012 (left carpometacarpus), NMB s.g.12959 (right carpometacarpus), NMB M.A.1098 (left femur), NMB s.g.21796 (right tibiotarsus). Locality and AgeSaint-Gerand-le-Puy, Allier, France; late Oligocene?early Miocene, MN2. Differential DiagnosisAll skeletal elements between 10% and 15% larger than the average for Laricola elegans. RemarksThe criteria chosen by Milne-Edwards (18671868) to distinguish between Laricola elegans and Laricola totanoidesnamely size and thickness of the bonesare largely unwarranted since these fall within a continuum (Table 1), and are therefore not discriminatory. From the original material examined by Milne-Edwards (18671868), only the ulnae and the illustrated humerus are large enough to be considered a different species from Laricola elegans. These clearly fall outside this range (Table 1), and we thus consider these specimens to be the true Laricola totanoides, even though we were unable to nd other distinguishing morphological features. This reasoning is nevertheless supported by the small number of bones of this size that are present in the collection. Given that there are no perceivable anatomical differences for all other bones falling within this continual size range, we have decided to unite all of these under Laricola elegans. We further designate as lectotype of L. totanoides the humerus (MNHN Av.4004), that was gured by Milne-Edwards (1868:pl. 57, gs. 1617), and labeled as type in the MNHN collection. One of the coracoids that we refer to this species was originally assigned to Totanus lartetianus, and it was the specimen illustrated by Milne-Edwards (18671868:pl. 63, gs. 1216) for this taxon. However, the presence of a foramen nervi supracoracoidei, as well as a dorsoventrally widened medial portion of the facies articularis sternalisboth features characteristic of Laromorphae and Stercorariidaeindicate that this bone is evidently not that of a scolopacid. Indeed, it matches exactly the coracoid of Laricola. Because it is marginally above the size range known for Laricola elegans (Table 1), we have referred it to L. totanoides. The carpometacarpus, tibiotarsus, and femur are only tentatively assigned to this species based on size. LARICOLA DESNOYERSII (Milne-Edwards, 1863) (Figs. 45, Table 1) Larus desnoyersii Milne-Edwards, 1863:161 (original description). Larus desnoyersii Milne-Edwards, 1868:344349; pl. 54, gs. 1526; pl. 56, gs. 110. Original MaterialMNHN Av.4044 (left coracoid), MNHN Av.4045 (right coracoid), MNHN Av.4046 (right coracoid; gured by Milne-Edwards, 1967: pl. 54, gs. 2223), MNHN Av.4048 (coracoid), MNHN Av.4050 (humerus, juvenile), MNHN Av.4061-67 (humeri), MNHN Av.4074 (ulna with distal end missing), MNHN Av.409598 (phalanx proximalis digiti majoris), MNHN Av.4106 (left tibiotarsus; juvenile), MNHN Av.4107 (left tibiotarsus; juvenile), MNHN Av.4108 (right tibiotarsus, juvenile), MNHN Av.4099 (left tarsometatarsus), MNHN Av.4100 (right tarsometatarsus), MNHN Av.4101 ( right tarsometatarsus). Newly Referred SpecimensNMB s.g.19325 (left coracoid, juvenile), NMB s.g.13024 (left coracoid, juvenile), NMB M.A.1106

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DE PIETRI ET AL.EARLY MIOCENE LARI FROM CENTRAL FRANCE (left humerus, proximal end missing), NMB s.g.7276 (left carpometacarpus, juvenile), NMB Gn.433 (left tarsometatarsus). Locality and AgeSaint-Gerand-le-Puy, Allier, France; late Oligocene?early Miocene, MN2. Differential DiagnosisApproximately 30% larger than Laricola totanoides. Differs from Laricola elegans and L. totanoides in having greater pneumaticity (i.e., a pneumatic recess) under the facies articularis clavicularis of the coracoid (Fig. 5D). RemarksBased on comparisons with the published illustrations, Olson (1985:180) hypothesized that the humerus of L. desnoyersii has a pneumatic foramen in the fossa pneumotricipitalis, and regarded this feature as potential evidence for afnities between L. desnoyersii and Stercorariidae. The bone is broken in this region, which, without proper examination of the specimen itself, gives the appearance of a pneumatic foramen, especially in Milne-Edwardss illustration. Our inspection of the original material in MNHN showed this hypothesis to be erroneous because there is no pneumatic foramen present in the fossa pneumotricipitalis, which is nevertheless very pronounced (Fig. 4N). We have referred Larus desnoyersii to the taxon Laricola because we were unable to nd any morphological features that are absent in Laricola. Although considerably larger than the other two species of Laricola previously described, such size differences within a genus are not unusual within Laromorphae, for instance in the extant genus Larus, the smallest species (L. minutus) and one of the largest (L. marinus) differ in size by over 60%. The rst description of Laricola desnoyersii dates back to 1863, in which Milne-Edwards referred to a distal end of a humerus. Unfortunately, he did not provide a catalog number nor did he gure the specimen, and an illustration of a distal end of a humerus is also absent in Milne-Edwards (18671868). There are several distal ends of humeri attributable to L. desnoyersii in MNHN (Av.406264), but because of the aforementioned lack of information, no holotype can be determined among these specimens. Mlkovsky (2002) considered the specimen mentioned in Milne-Edwards (1863) to be the lectotype for the species. However, we note that with no catalog number, a designation of this bone as a lectotype does not meet the requirements of the International Code of Zoological Nomenclature, which demand a type to be specied. If a lectotype were to be chosen, then we propose specimen MNHN Av.4059, a right humerus, which was fully illustrated and described by Milne-Edwards 18671868 (pl. 554, gs. 2425).

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The scapula and the pelvis referred to L. desnoyersii by MilneEdwards (18671868:pl. 56, gs. 12 and 57) do not belong to Charadriiformes. The single carpometacarpus in MNHN in good condition assigned to Laricola desnoyersii is likely to belong to the Scolopacidae. Description and Comparisons CoracoidWe have assigned two juvenile coracoids from NMB to Laricola desnoyersii based on size. Unfortunately many features are worn and few details can be recognized. The specimens in MNHN are much better preserved and, apart from a better-developed recess distal to the extremitas omalis (pneumaticity under the facies articularis clavicularis, see Fig. 5D)which could be size related, are not different from other Laricola. HumerusAlthough considerably larger, the humerus of Laricola desnoyersii matches that of Laricola elegans in proportions. The fossa m. brachialis of L. desnoyersii is not as deep as in most extant species of Larus, thus resembling Laricola elegans. The well-marked tubercle on the cranial surface of the distal end, between the base of the tuberculum supracondylare dorsale and the fossa m. brachialis, is present. UlnaOnly one ulna present in the Saint-Gerand-le-Puy collection in MNHN can be assigned to Laricola desnoyersii. All other ulnae in this collection either belong possibly to Scolopacidae, or to Anseriformes. CarpometacarpusCarpometacarpus NMB s.g.7276 (Fig. 5Q) is only tentatively assigned to this taxon based on size. Its edges are worn and most of the distal end is missing. Although the one complete specimen from MNHN is indeed larger (43.4 mm), MNHN Av.4092 bears a dorsally rounder condylus medialis and may belong to the Scolopacidae. Also, the synostosis metacarpi distalis is proportionally longer than in stercorariids and Laromorphae, thus resembling some Scolopacidae. The two other specimens from MNHN (Av.4091 and Av.4093) are too fragmentary for a reasonable identication. Unfortunately, the distinctive features of the distal end of NMB s.g.7276 are missing. The processus extensorius, however, is intact and resembles that of Laricola. Other Wing BonesOnly fragmentary specimens of the phalanx proximalis digiti majoris are present in the MNHN collection. However, it is clear that this bone was, as in all Laromorphae and Stercorariidae, bifenestrated. As in Laricola and other

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FIGURE 5. Postcranial skeletal elements of Laricola and Sternalara gen. nov. AB, right coracoid of Laricola elegans (NMB s.g.18825) in ventral (A) and dorsal (B) views; C, right coracoid of Laricola totanoides (NMB s.g.18809) in dorsal view; D, right coracoid of Laricola desnoyersii (MNHN 4045) in dorsal view; E, left coracoid of Larus ridibundus in dorsal view; F, left coracoid of Anous stolidus in dorsal view; GH, caudal aspect of right scapus claviculae (furcula) of Laricola elegans (NMB s.g.16378); I, caudal aspect of left scapus claviculae of Larus hyperboreus; J, right scapula of Laricola elegans (NMB s.g.19575) in lateral view; KL, sternum of Laricola elegans (NMB s.g.15879); M, ventral aspect of left ulna of Laricola totanoides (NMB s.g.12946); N, ventral view of right ulna of Laricola elegans (NMB s.g.18983); O, ventral view of right ulna of Laricola robusta (NMB M.A.2539); P, ventral view of left ulna of Sternalara minuta (NMB M.A.5161); Q, ventral view of left carpometacarpus of Laricola desnoyersii (NMB s.g.7275); RS, right carpometacarpus of Laricola totanoides (NMB M.A.5181) in dorsal (R) and ventral (S) views; TU, left carpometacarpus of Laricola elegans (NMB M.A.3178) in ventral (T) and dorsal (U) views; VW, left phalanx proximalis digiti majoris of Laricola elegans (NMB s.g.16768) in ventral (V) and dorsal (W) views; X, left phalanx proximalis digiti majoris of Laricola desnoyersii (MNHN Av.4098) in ventral view; Y, partial left portion of pelvis of Laricola elegans (NMB s.g.7516) in medial view; ZAA, synsacrum of Laricola elegans (NMB s.g.16329) in dorsal (Z) and ventral (AA) views; BB, left femur of Laricola intermedia (NMB M.A.5198) in caudal view; CC, right femur of Laricola elegans (NMB s.g.19038) in caudal view; DDEE, left tibiotarsus of Laricola elegans (NMB s.g.19127) in cranial (DD), proximal (EE, not to scale), and medial (FF, not to scale) views; GGHH, left tarsometatarsus of Laricola desnoyersii (Gn.433) in plantar (GG) and dorsal (HH) views; IIJJ, right tarsometatarsus of Laricola elegans (NMB s.g.13720) in plantar (II) and dorsal (JJ) views; KK, not to scale, right hypotarsus of Laricola desnoyersii (MNHN Av4101); LL, not to scale, left hypotarsus of Laricola desnoyersii (MNHN Av.4099); MM, not to scale, right hypotarsus of Laricola elegans (NMB s.g.13720); NN, not to scale, left hypotarsus of Laricola elegans (NMB s.g.16913); OOQQ, right carpometacarpus of Laromorphae indet. (NMB s.g.19007) in dorsal (OO, QQ, not to scale) and ventral (PP) views; RRSS, right humerus (MNHN Av.7318) of Laromorphae indet. in cranial (RR) and dorsal (SS) views. Abbreviations: ac, apex carinae; bb, bony bridge at sulcus interosseus; ch, hypotarsal canal; cih, cristae intermediae hypotarsi; cmh, crista medialis hypotarsi; faa, facies articularis acrocoracoidea; fm, femoral prominence; fvd, foramen vasculare distale; pi, processus intermetacarpalis; pii, processus internus indicis; pl, processus lateralis; pn, pneumatic recess; rcf, recessus caudalis fossae; sh, sulcus hypotarsi; tlcv, tuberculum ligamenti collateralis ventralis.

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TABLE 1.

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Skeletal measurements (in mm) of the long bones of three Laricola species: L. elegans, L. totanoides, and L. desnoyersi. Laricola elegans n Range 19.722.4 (21.3) 17.319.0 6.47.9 (6.9) 3.13.8 47.053.9 (48.4) 8.910.7 (9.9) 6.47.6 (7.0) 2.73.3 (3.0) 51.462.9 (56.9) 5.16.2 (5.9) 43.95.5 (4.3) 2.43.0 (2.3) 27.932.0 (30.0) 6.47.9 (3.2) 3.64.5 (3.2) 22.527.1 (22.2) 4.25.2 (4.0) 4.25.2 (4.4) 1.82.3 (1.8) 45.151.9 (52.0, maximum length) 3.55.1 (4.9) 3.94.4 (3.9) 1.62.0 (1.9) 31.837.7 (34.1) 4.15.0 (4.7) 4.04.6 (4.4) 1.41.9 (1.7) Mean 21.0 18.4 7.1 3.5 50.0 9.5 6.9 3.0 56.4 5.6 4.7 2.7 30.2 6.9 4.1 24.4 4.6 4.7 2.0 48.8 3.8 4.1 1.8 34.3 4.6 4.3 1.6 SD 0.8 0.6 0.4 0.2 2.0 0.4 0.3 0.6 3.4 0.3 0.4 0.2 1.4 0.35 0.23 1.3 0.3 0.3 1.1 2.8 0.4 0.1 0.1 2.0 0.2 0.2 0.1 n 1 1 1 1 1 1 1 1 1 2 1 4 4 4 4 3 3 2 1 1 1 1 1 1 1 1 Laricola totanoides Range 22.8 23.3 (22.0) 20.9 19.8 8.0 7.7 (7.5) 3.4 4.1 57(est) (51.0) (10.0) 7.68.0 (7.1) 3.6 (3.0) 65.166.3(est) (65.0) 5.66.5 (6.7) 4.85.5 (4.4) 2.83.0 (2.9) 33.134.6 (30.0) 7.47.5 (3.4) 4.64.9 (3.4) 29.1 (25) 5.1 (4.7) 5.4 (4.9) 2.3 (1.8) 54.0 () 5.4 () 4.4 (4.5) 2.2 (1.9) (37.2) (5.0) (4.5) (2.0) Mean 7.8 65.5 6.1 5.2 2.9 34.0 7.4 n 1 1 1 1 2 2 1 2 1 2 1 1 1 1 1 1 2 2 1 2 1 2 1 2 1 2 Laricola desnoyersi Range 32.0 (30.0) 29.1 10.0 (11.0) 5.5 84.385.3 (85.3) 14.515.6 11.2 10.411.3 5.3 4.44.9 94.5(est) (96.3) 9.6 4.6 39.4 (44.0) 8.9 (9.6) (6.0) 72.3 5.86.3 (6.1) 2.93.6 49.1 44.145.7 (45.7) 7.1 6.47.9 6.9 6.16.4 2.6 2.6 Mean 84.8 15.1 10.9 4.7 6.1 3.3 44.9 7.2 6.3 2.6

Coracoid Maximum length Medial length Distal width Width of processus acrocoracoideus Humerus Maximum length Proximal width Distal width

14 14 14 14 43 37 43 43 51 51 51 48 40 40 38 34 34 34 34 17 16 15 18 24 23 23 24

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Minimum width of shaft Ulna Maximum length Proximal width Distal width Minimum width of shaft Carpometacarpus Maximum length Proximal depth Distal depth Femur Maximum length Proximal width Distal width Minimum width of shaft Tibiotarsus Length from facies articularis Proximal width Distal width Minimum width of shaft Tarsometatarsus Maximum length Proximal width Distal width Minimum width of shaft

In parentheses are the measurements provided by Milne-Edwards (18671868). Note that for all bones except the ulna, the measurements listed by Milne-Edwards for Laricola totanoides overlap with those of Laricola elegans. All bones are from NMB, except those marked by an asterisk, which are from MNHN.

DE PIETRI ET AL.EARLY MIOCENE LARI FROM CENTRAL FRANCE


TABLE 2.

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Measurements (in mm) of the skeletal elements known for Laricola intermedia, L. robusta, Sternalara minuta, and S. milneedwardsi. Sternalara milneedwardsi SD 1.0 0.1 0.2 0.2 0.4 0.3 0.4 0.2 n 1 1 1 1 Range 133.9 25.6 18.1 7.7

Laricola intermedia n Humerus Maximum length Proximal width Distal width Minimum width of shaft Ulna Maximum length Proximal width Distal width Minimum width of shaft Femur Maximum length Proximal width Distal width Minimum width of shaft 1 1 1 1 2 2 2 2 Range 75.3 12.9 9.7 4.4 31.531.6 7.57.8 5.96.0 2.82.9 Mean 31.6 7.7 6.0 2.9 n 9 9 9 9 3 3 3 3

Laricola robusta Range 47.550.9 6.77.4 9.811.0 3.03.4 53.056.0 5.46.2 4.75.0 2.93.1 Mean 48.8 7.1 10.3 3.2 54.6 5.2 4.9 3.0 SD 1.1 0.3 0.4 0.2 1.5 0.3 0.2 0.1 n 6 6 6 6 4 4 4 4

Sternalara minuta Range 44.647.5 8.58.8 6.26.7 2.63.0 49.750.7 5.05.6 4.25.0 2.42.9 Mean 46.2 8.7 6.5 2.8 50.3 5.2 4.7 2.7

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All bones are from NMB, except those marked by an asterisk, which are from MNHN.

Laromorphae, the processus internus indicis is strongly elongated caudally (Fig. 5X). TibiotarsusUnfortunately, the tibiotarsi in MNHN belonging to L. desnoyersii are juveniles and/or badly preserved specimens. The distal end closely resembles that of other Laromorphae, exhibiting a wide sulcus extensorius. TarsometatarsusCorresponding with the condition in Laricola, the tarsometatarsus of L. desnoyersii is proportionally very long. MNHN Av.4101 was gured by Milne-Edwards (18671868:pl. 54, gs. 1618) and has a hypotarsus similar to that of Laricola elegans (Fig. 5KK), exhibiting an elongated crista medialis hypotarsi, two crista intermedia hypotarsi, and no closed canal for the tendon of the musculus digitorum longus. The hypotarsus of MNHN Av.4099 (Fig. 5LL) differs from this specimen in that it bears a closed canal for the tendon of the musculus digitorum longus, as is the case for one specimen we have assigned to Laricola elegans (Fig. 5NN). It should be noted that slight variations in length and orientation of the cristae of the hypotarsus are seen throughout the genus Larus, and indeed within Laricola (which partly may be due to the preservation of these delicate structures). NMB G.n.433 (Fig. 5GGHH) is only slightly larger than the ones from MNHN, and unfortunately the hypotarsus is worn, but matches Laricola in all other features, as do the specimens from MNHN. LARICOLA INTERMEDIA, sp. nov. (Figs. 45, Table 2) HolotypeMNHN SG.13744 (right humerus). Type Locality and AgeSaint-Gerand-le-Puy, Allier, France; late Oligocene?early Miocene, MN2. Differential DiagnosisSpecies of Laricola, which is distinguished from L. elegans and L. totanoides by its much greater size, being, however, smaller than L. desnoyersii (see Table 2). MeasurementsSee Table 2. Referred SpecimensNMB M.A.5197 (left femur), NMB M.A.5198 (left femur). EtymologyThe species name Laricola intermedia alludes to the intermediate size between Laricola elegans/L. totanoides and L. desnoyersii. RemarksThe femora are only tentatively assigned to this taxon based on size; they are smaller than what would be predicted for Laricola desnoyersii. Description and ComparisonsThe humerus MNHN SG.13744 (Fig. 4JK) is approximately 15% smaller than that of Laricola desnoyersii. A second fossa pneumotricipitalis is

absent and, as in Laricola desnoyersii, the fossa pneumotricipitalis that is present is very pronounced, but bears no pneumatic foramen. The sulcus nervi coracobrachialis is well marked; other features of the proximal end do not differ from those of other Laromorphae. The fossa musculi brachialis is deep, and the well-marked tubercle on the cranial surface of the distal end, between the base of the tuberculum supracondylare dorsale and the fossa m. brachialis, is present. The femora bear a well-marked tuberculum musculi gastrocnemialis lateralis, but the prominence just proximal of the condylus medialis that is present in some Laromorphae is not evident because the area is worn. LARICOLA ROBUSTA, sp. nov. (Figs. 45, Table 2) Larus totanoides Milne-Edwards, 1868:358 (original description). HolotypeMNHN Av.3777 (right humerus, labeled as Laricola totanoides). Type Locality and AgeSaint-Gerand-le-Puy, Allier, France; late Oligocene?early Miocene, MN2. Differential DiagnosisMember of Laricola with stout bones; differing from other Laricola in: enlarged crista bicipitalis and crista deltopectoralis, broader and better-developed processus supracondylaris dorsalis, proportionally shorter shaft of humerus (see Fig. 4, compare AB with HI). MeasurementsSee Table 2. Referred SpecimensNMB s.g.18887 (left humerus), NMB s.g.16467 (right humerus), NMB s.g.16514 (left humerus), NMB Ph.2013 (left humerus), NMB s.g.16554 (left humerus), NMB s.g.7209 (right humerus), NMB s.g.7206 (right humerus), NMB s.g.21 (right humerus), NMB s.g.18465 (right humerus). The following ulnae are only tentatively assigned to this taxon: NMB M.A.2539 (left ulna), NMB s.g.16581 (right ulna), NMB s.g.20 (right ulna). EtymologyLaricola robusta is named after its stout and robust skeletal elements. RemarksThe humeri that we have assigned to this taxon lie approximately in the mid-size range of Laricola elegans (see Table 1 and 2). The proportions of the shaft are unlike those for other Laricola, in which the shaft is proportionally longer than in this species (Fig. 4HI). Because of the wide size ranges (Table 1) for all long bones of Laricola elegans, it is difcult to separate material based on stoutness, and for that reason the referred ulnae can only be tentatively allocated to this taxon. In the

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 4, 2011 Description and ComparisonsThe humeri (Fig. 4SU) are smaller than those assigned to Laricola elegans. They differ from Laricola in that a second fossa pneumotricipitalis is present. This fossa is, nevertheless, not as deep as that of Laridae and Sternidae s.s. Also, it does not reach into the caput humeri, and is thus more caudally situated. The sulcus nervi coracobrachialis is distinct. The fossa musculi brachialis is deep, and the wellmarked tubercle on the cranial surface of the distal end, between the base of the tuberculum supracondylare dorsale and the fossa m. brachialis, is evident. Again, we have tentatively assigned the ulnae to this species based on size. Because Sternalara minuta is close in size to the smallest specimens of Laricola elegans, the smallest ulnae within the range assignable to the latter taxon may indeed belong to S. minuta. As already mentioned, ulnae within Laromorphae are fairly uniform, and it is thus not possibly to condently refer the smallest ulnae (e.g., NMB M.A.2550; 50.4 mm) to S. minuta, but we do acknowledge this possibility. STERNALARA MILNEEDWARDSI, sp. nov. (Fig. 4, Table 2) HolotypeMNHN SG.13742 (right humerus). Type Locality and AgeSaint-Gerand-le-Puy, Allier, France; late Oligocene?early Miocene, MN2. Differential DiagnosisAs for genus; differs from Sternalara minuta in: second fossa pneumotricipitalis not as wide (Fig. 4R). MeasurementsSee Table 2. EtymologyIn honor of palaeontologist Alphonse MilneEdwards (18351900). RemarksThe humerus belonging to this taxon is markedly larger than that of Sternalara minuta (Table 2). Still, because of the presence of a second fossa pneumotricipitalis, albeit not a deep one, as in extant Laromorphae, we tentatively assign this specimen to the same genus. Description and ComparisonsThe second fossa pneumotricipitalis (Fig. 4R) is not as deep as in extant Laridae or Sternidae, also not reaching into the caput humeri. Furthermore, it is not as wide as that of Sternalara minuta. The sulcus nervi coracobrachialis is well dened. As in Stercorariidae, Laricola and some other Laromorphae, the humerus bears a well-marked tubercle on the cranial surface of the distal end, between the base of the tuberculum supracondylare dorsale and the fossa m. brachialis (Fig. 4Q). This taxon differs from Sternalara minuta in that the processus supracondylaris dorsalis does not project as cranially as in the latter species (Fig. 4Q, S), and in that the processus supracondylare ventrale is distinctly more pronounced in Sternalara milneedwardsi. LAROMORPHAE INDET. Referred SpecimensNMB s.g.19007 (right carpometacarpus), NMB s.g.2044 (right carpometacarpus, distal end). Figure 4OOQQ. MeasurementsNMB s.g.19007: maximum length, 32.4 mm; proximal depth, 7.0 mm; distal depth, 4.4 mm; length of spatium, 20.5 mm. NMB s.g.2044: distal depth, 5.5 mm. RemarksIn size these carpometacarpi match the largest specimens of Laricola elegans. They differ from Laricola, nevertheless, in having a slightly less developed processus intermetacarpalis and in the presence of a bony bridge at the sulcus interosseus. Within the Lari, we have found the latter feature to be present only in Catharacta skua. There are no other features differentiating it from the carpometacarpus of the Laridae, Sternidae, and Stercorariidae. Referred SpecimenMNHN Av.7318 (right humerus), Figure 5RRSS. MeasurementsIn mm; greatest length: 72.2; distal width: 10.3; minimum width of shaft: 4.3.

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MNHN collection these elements are labeled Larus totanoides, to which they were assigned by Milne-Edwards (18671868), who described Larus totanoides as being much stouter than Laricola elegans in all bones. As we have shown, much of this difference is attributed to a wide size range of L. elegans, but in the case of the humerus, the different proportions allow for the construction of a new species. It is, nevertheless, not understandable why Milne-Edwards would have referred these specimens to Larus totanoides when the illustration of this taxon clearly depicts the much larger incomplete humerus already mentioned in the species section for Laricola totanoides. Description and ComparisonsThe humeri (Fig. 4HI) are stouter than those of Laricola elegans, with a broader and betterdeveloped processus supracondylaris dorsalis. The fossa musculi brachialis is deep, and the bone bears the well-marked tubercle on the cranial surface of the distal end, between the base of the tuberculum supracondylare dorsale and the fossa m. brachialis. They do not exhibit a second incipient fossa pneumotricipitalis (see below) and both the deltoid and bicipital cristae are better developed than in the other specimens of Laricola. The shaft is proportionally shorter than in other Laricola (compare with Fig. 4AB). The ulnae (Fig. 5O) do not differ from other ulnae that we have assigned to this genus, and it is because of their thickness in relation to their length (Table 1 and 2) that we have referred these to Laricola robusta. FAMILY incertae sedis STERNALARA, gen. nov. Type GenusSternalara. Type SpeciesSternalara minuta. Included SpeciesSternalara minuta, Sternalara milneedwardsi. Differential DiagnosisDiffers from Laricola in: presence of a second fossa pneumotricipitalis (humerus); differs from Laridae and Sternidae in: second fossa pneumotricipitalis not as marked, not reaching into the caput humeri. EtymologyThe genus name refers to a closer morphology of the humerus to more derived Laromorphae, such as Sterna and Larus. STERNALARA MINUTA, sp. nov. (Figs. 45, Table 2) HolotypeNMB s.g.12727 (right humerus). Type Locality and AgeSaint-Gerand-le-Puy, Allier, France; late Oligocene?early Miocene, MN2. Differential DiagnosisAs for genus; differs from Sternalara milneedwardsi in: second fossa pneumotricipitalis proportionally wider. Overall much smaller than S. milneedwardsi (around 65% smaller). MeasurementsSee Table 2. Referred SpecimensNMB M.A.1582 (left humerus), NMB s.g.2189 (left humerus), NMB s.g.17520 (right humerus), NMB s.g.5703 (right humerus), NMB M.A.2513 (right humerus). The following ulnae are only tentatively assigned to this taxon: NMB M.A.1611 (right ulna), NMB M.A.2550 (right ulna), NMB M.A.5161 (right ulna), NMB s.g.16570 (right ulna). EtymologyThe species name minuta refers to the small size of the new species. RemarksThis species is smaller than most individuals of Laricola elegans, with some bones overlapping with the smallest specimens. Although the humerus is very close to that of Sterna paradisaea, we refrain at this point from including it in the family Sternidae, particularly because of the difference in shape of the fossa pneumotricipitalis. In any case, a new genus is justied by the amount of time between this record and that of members of recent Sternidae (putative record dating back to the early Pliocene, see Olson and Rasmussen, 2001).

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FIGURE 6. AB, two most parsimonious trees (tree length = 61 steps, retention index = 0.67, consistency index = 0.68), showing character optimization. Black circles represent non-homoplastic apomorphies, white circles homoplastic ones. C, strict consensus cladogram of two most parsimonious trees. Bootstrap values are shown to the right of the branches, Bremer supports are shown to the left.

RemarksAlthough close in size to Laricola intermedia, it differs from Laricola in several aspects: the tuberculum supracondylare ventrale is situated more laterally in this specimen, making the ventral side appear much wider than in Laricola. Laricola also has a proportionally longer shaft. The tuberculum ventrale is much enlarged in this specimen, and the caput humeri is clearly more prominent. A second fossa pneumotricipitalis is absent. PHYLOGENETIC ANALYSIS Analysis of the character matrix resulted in two most parsimonious trees (tree length = 61 steps, retention index = 0.67, consistency index = 0.68), which differed only in the position of Laricola and Anous relative to the clade including Sternidae, Rynchopidae, and Laridae (Fig. 6AB). The strict consensus tree is shown in Figure 6C. Three characters were optimized as synapomorphies of the clade (Laricolidae + Anous + Sternidae + Rynchopidae + Laridae) in both most parsimonious trees: (1) beak, rostrum maxillare very long, occupying at least one third of the length of the beak (reversed in Laridae); (15) mandible, presence of fossa caudalis; and (20) furcula, extremitas omalis with subovate facies articularis acrocoracoidea that extends over nearly entire width of scapus claviculae. Laricola and Anous fall outside a clade including Sternidae, Rynchopidae, and Laridae. The position of Anous outside this clade agrees with results obtained in recent molecular phylogenies (Baker et al., 2007). The position of Laricola as sister taxon to all other Laromorphae is supported by the presence of a shallow facies articularis acrocoracoidea in the furcula (21), although it would be equally parsimonious to presume Anous in this position, which is endorsed by the presence of a shallow fossa musculi brachialis in the humerus (29). In the consensus tree (Fig. 6C) both Anous and Laricola result in a polytomy with the clade

including Sternidae, Rynchopidae, and Laridae. No autapomorphies were recovered for Laricola. The following characters were optimized as unambiguous synapomorphies for the clade Sternidae + Rynchopidae + Laridae, but are also present in Limosa: (5) skull, processus maxillopalatinus not forming a wide and deeply concave trough; (14) mandible, presence of well-developed fenestra caudalis; (26) humerus, presence of a well-developed second (dorsal) fossa pneumotricipitalis that undercuts caput humeri; and (30) humerus, absence of marked tubercle on cranial surface of distal end, between base of tuberculum supracondylare dorsale and fossa m. brachialis. A sister-group relationship between Sternidae s.s. and Rynchopidae is supported by one ambiguous synapomorphy (36; femur, distal end, crista supracondylaris medialis with distinct caudally projecting tubercle just proximal of condylus medialis) recovered only with ACCTRAN optimization, and by the following unambiguous synapomorphies: (3) skull, os lacrimale with maximum craniorostral width at least twice as much as maximum mediolateral width); (7) skull, processus maxillopalatinus, concave surface bridged by a pierced sheet of bone; and (34) pelvis, absence of recessus caudalis fossae. Character 3 is a unique synapomorphy, whereas characters 7 and 34 show homoplasy, with the same condition also present in Glareola, and Limosa and Glareola, respectively. The monophyly of Sternidae is supported by two unambiguously optimized characters: (15) mandible, fossa caudalis present and well-marked, chevron-shaped in ventral view; and (37) femur considerably longer than tarsometatarsus. The following homoplastic characters support Laridae: (1) beak, rostrum maxillare short; occupying less than one third of the length of the beak; (2) skull, os frontale, paired fossae glandularum nasales meeting at midline of skull; and (6) skull, maxillopalatine strut A present.

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JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 31, NO. 4, 2011 DISCUSSION ber of species and the combination of features present in all these species warrant the construction of a new family, Laricolidae. Sternalara has a more derived morphology of the humerus, with a second fossa pneumotricipitalis still differing from that of extant Laromorphae in being shallower and not reaching into the caput humeri, implying that it is unlikely for this taxon to have belonged in any recent families of Laromorphae. Whether Sternalara was more closely related to Laricola or to more-derived Laromorphae is difcult to say based on the material available, namely humeri, which is also why we considered this taxon unsuitable for inclusion in the phylogenetic analysis. Not much material from the original description of Laricola totanoides by Milne-Edwards can still be retained in this species, despite him mentioning several skeletal elements, many of which are labeled L. totanoides in the MNHN collection. We consider the depicted humerus and the ulnae with the dimensions specied by him as true L. totanoides because they are explicitly larger than the range we determined for Laricola elegans. All other bones fall within the large size spectrum of L. elegans, whereas the humeri and some ulnae clearly belong to a different species altogether, Laricola robusta. Concerning Laricola elegans, we acknowledge that the wide size range indicated for this bird may be too broad for a single species, and indeed this range of variation may encompass several species, which is primarily evinced in the variations found within the skull and the conguration of the hypotarsus. In living Laromorphae, size differences of this magnitudearound 20%are undocumented (see del Hoyo, 1996), but direct comparisons with the fossil record have not yet been performed. As explained in Geology (see above), the fact that the fossil collec tion from Saint-Gerand-le-Puy potentially spans a period ranging from the late Oligocene to the early Miocene may also explain the large size variability found within the material attributed to Laricola elegans. Incorporating Laricola into a phylogenetic analysis of extant taxa has demonstrated that this extinct taxon is an early offshoot of Laromorphae whose divergence predates the radiation of modern gulls, terns, and skimmers. The fossil record of crown group Laromorphae is fairly recent. Putative terns and gulls from Miocene deposits (e.g., Miller and Sibley, 1941; Grigorescu and Kessler, 1977; Mlkovsky, 1996) are based on frag mentary records, and cannot be assigned to any modern genus. Pliocene fossil evidence for Sternidae is minor, including carpometacarpi of two taxa from, most likely, the early Pliocene North America (Olson and Rasmussen, 2001). Pliocene fossils of Laridae appear to be more recurrent, particularly in North America: several specimens were described from the early Pliocene of North Carolina (Olson and Rasmussen, 2001); Emslie (1995) described two species (Larus perpetuus and L. lacus) from the Late Pliocene of Florida; and Larus elmorei is known from the earlymiddle Pliocene of Florida (Brodkorb, 1967). None of these fossils seems to signicantly depart from the morphology of extant gulls. The Quaternary fossil record of Laromorphae is far more abundant and diverse (e.g., Brodkorb, 1967, Mlkovsky, 2002). Fossils from Saint-Gerand-le-Puy are difcult to date with precision. The quarry in Montaigu-le-Blin and its fauna yielded fossils from biozone MN2 (from 22.5 to 20.5 Ma), but the uncertainty as to whether the old collections all come from this very place casts doubt on the precise dating of the fossil remains. Indeed, fossils were collected in the 19th and early 20th centuries from the whole area in various localities surrounding Saint-Gerand-le-Puy and no exact provenience was recorded. It is therefore possible that the different species of laromorphs present in the lake system during this time did not co-occur. This lake system was most certainly used as breeding grounds by several taxa of birds, including Laricola, as reected by the vast amount of juvenile individuals recovered. Whether these charadriiform birds were seasonal

Phylogenetic Afnities between Laricola and Other Laromorphae Laricola trenchantly differs from gulls in several skull features (e.g., length of rostrum maxillare, the shape of the fossae glandularum nasales, and processus maxillopalatini), mandible (absence of fenestra caudalis), and in few postcranial features (furcula with shallow facies articularis clavicularis, humerus with no second fossa pneumotricipitalis, and better marked attachment site for m. pronator brevis). Although Laricola is generally referred to gulls (Laridae) (e.g., Mlkovsky, 2002), this genus has been considered more primitive than modern Laridae by some authors (e.g., Ballmann, 1976; Olson, 1985), who based this statement on the plesiomorphic features of the humerus. As detailed above, our analysis supports a placement of Laricola outside a clade including Sternidae, Rynchopidae, and Laridae. Although the aim of our analysis was to place the fossil genus Laricola within a phylogenetic context of recent forms and not to provide an exhaustive assessment of the interrelationships of recent Laromorphae based on osteological characters, we note that our results also constitute the rst anatomical evidence for a paraphyletic Sternidae, which has not yet been recovered with anatomical data (e.g., Chu, 1998; Strauch, 1978). The following, presumably plesiomorphic, features set Anous apart from extant gulls, terns, and skimmers: the processus maxillopalatinus (skull) forming a wide and deeply concave trough (the maxillopalatine process is less concave in other extant Laromorphae, displaying an entirely different shape, see Fig. 3HL for comparisons), the absence of well-developed fenestra caudalis (mandible), the absence of a second fossa pneumotricipitalis (humerus), and the presence of a very distinct attachment site for m. pronator brevis (character 30; humerus). Although Anous clearly falls outside the clade comprising gulls, terns, and skimmers, our analysis did not resolve whether Anous or Laricola are more closely related to this group. Anous shares one plesiomorphic feature with Stercorariidae, namely the shallow fossa m. brachialis, whereas Laricola and Stercorariidae both bear a shallow facies articularis clavicularis on the furcula. Both morphological and molecular analyses have yielded conicting results concerning the position of Rynchops. In his phenetic study of the Lari, Schnell (1970) concluded that skimmers are overall more similar to terns, something that had already been speculated upon by Zusi (1962) based on the specialized feeding habits and corresponding morphological adaptations found in both these groups. Livezey and Zusis (2007) analysis of modern birds based on anatomical characters recovered Rynchopidae as sister group to the Laridae and Sternidae, as did the analysis based on nuclear DNA sequences by Ericson et al. (2003). Baker et al. (2007) recovered a sistertaxon relation between Laridae and Rynchopidae, albeit with low branch support. Our results are consistent with those of Fain and Houde (2007) who obtained substantial support for a sister-taxon relationship between Rynchopidae and Sternidae. We have found this relationship to be overall supported by four anatomical features (see Results), whereas this grouping was endorsed by only one character in Mayr (2011). Diversity of Early Laromorphae The fossil record indicates that the early Miocene diversity of Laromorphae is far higher than assumed previously. At least ve species of Laricola and two species of Sternalara, gen. nov., existed by the early Miocene. Among Laricola, L. elegans, L. totanoides, L. desnoyersii, and L. intermedia differ mainly in size, whereas L. robusta can be distinguished by its relatively shorter and stouter shaft of the humerus, as well as by betterdeveloped cristae bicipitalis and deltopectoralis. The sheer num-

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DE PIETRI ET AL.EARLY MIOCENE LARI FROM CENTRAL FRANCE visitors or permanent residents of the area can, however, only be speculated upon. ACKNOWLEDGMENTS We thank C. Meyer (NMB) for facilitating the study of the fossil specimens. We are grateful to R. Allain (MNHN) for his assistance during a visit to the avian fossil collection in MNHN. S. Hertwig and M. Schweizer (Naturhistorisches Museum Bern) provided useful comments on the phylogenetic analysis. M. Weick (NMB) assisted with the preparation of fossil specimens. We thank the Willi Hennig Society for subsidizing the free edition of TNT. A travel grant from Stiftung zur Forderung des Naturhistorischen Museums (Basel, Switzerland) to V.L.D. supported this research. Helpful comments by D. Ksepka (University of North Carolina, Chapel Hill, U.S.A.), C. Mourer-Chauvire (Universite Claude Bernard, Lyon, France), and T. Worthy (University of New South Wales, Sydney, Australia) improved the manuscript. LITERATURE CITED
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