A revision of the Lari (Aves, Charadriiformes) from the early Miocene of Saint-Gérand-le-Puy...

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A revision of the Lari (Aves, Charadriiformes) from theearly Miocene of Saint-Gérand-le-Puy (Allier, France)Vanesa L. De Pietri a , Loïc Costeur b , Marcel Güntert a & Gerald Mayr ca Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, CH-3005, Bern,Switzerlandb Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001, Basel, Switzerlandc 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, Loïc Costeur, Marcel Güntert & Gerald Mayr (2011): A revision of the Lari (Aves,Charadriiformes) from the early Miocene of Saint-Gérand-le-Puy (Allier, France), Journal of Vertebrate Paleontology, 31:4,812-828

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Journal of Vertebrate Paleontology 31(4):812–828, July 2011© 2011 by the Society of Vertebrate Paleontology

ARTICLE

A REVISION OF THE LARI (AVES, CHARADRIIFORMES) FROM THE EARLY MIOCENEOF SAINT-GERAND-LE-PUY (ALLIER, FRANCE)

VANESA L. DE PIETRI,*,1 LOIC COSTEUR,2 MARCEL GUNTERT,1 and GERALD MAYR3

1Naturhistorisches Museum der Burgergemeinde Bern, Bernastrasse 15, CH-3005 Bern, Switzerland, [email protected];[email protected];

2Naturhistorisches Museum Basel, Augustinergasse 2, CH-4001 Basel, Switzerland, [email protected];3Forschungsinstitut Senckenberg, Division of Ornithology, Senckenberganlage 25, D-60325 Frankfurt am Main, Germany,

[email protected]

ABSTRACT—Three species of gull-like birds were described by Milne-Edwards in the 19th century from the early Mioceneof Saint-Gerand-le-Puy, France, two of which have been since then moved to the genus Laricola. These fossils are redescribedand 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 increasingthe number of charadriiform taxa known from this locality. To find out how Laricola relates to modern taxa, we performeda phylogenetic analysis of 41 osteological characters, which resulted in Laricola being placed outside of Lariidae, as earlyrepresentatives of the newly designated group Laromorphae, which encompasses terns, gulls, and skimmers. The results ofour 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 initiallydescribed by Milne-Edwards (1863, 1867–1868), who consideredthem to be members of the modern genus of gulls Larus. Cur-rently, little is known about the phylogenetic affinities of thesevery abundant Miocene birds, though they can clearly be placedwithin the Lari, which is one of the three clades that, according torecent analyses of nuclear and mitochondrial sequence data (e.g.,Ericson et al., 2003; Paton et al., 2003; Baker et al., 2007; Fain andHoude, 2007), make up the order Charadriiformes, the other twobeing Charadrii (plovers and allies) and Scolopaci (sandpipersand allies).

The Lari comprise the Dromadidae (crab plover), Ster-corariidae (skuas), Alcidae (auks), Laridae (gulls), Sternidae(terns), Rynchopidae (skimmers), and Glareolidae (pratincolesand coursers), with the Turnicidae (buttonquails) being the sis-ter group to the whole clade (Ericson et al., 2003; Paton et al.,2003; Baker et al., 2007; Fain and Houde, 2007; Mayr, 2011). Al-though phylogenetic affinities within this group are largely consis-tent throughout the above-cited molecular studies, there is somedisparity concerning some families. For instance, the position ofRynchopidae is variable in both molecular and morphologicalstudies (Schnell, 1970; Ericson et al., 2003; Baker et al., 2007;Livezey and Zusi, 2007; Fain and Houde, 2007; Mayr, 2011) andit is thus uncertain whether skimmers are most closely related toterns, gulls, or are sister group of both. Also, recent research (e.g.,Baker et al., 2007; see also Odeen et al., 2010) suggests that theSternidae are a paraphyletic grouping, with Anous (noddies) andGygis (white terns) being sister taxa to a clade including all otherSternidae, Laridae, and Rynchopidae. So far, no morphologicalassessment of the relationships within Lari has included any fos-sil representatives of this group.

*Corresponding author.

The Paleogene fossil record of members of the Lari is poor(Mayr, 2009). Noteworthy among these fossil specimens arestem-group representatives of the Turnicidae from the Oligoceneof France and Germany (Mayr, 2000; Mayr and Knopf, 2007).Fossils assigned to the Glareolidae were recovered from the earlyOligocene of Belgium (Mayr and Smith, 2001). The purportedoccurrence 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. Glare-olidae are represented by Mioglareola gregaria and Glareolaneogena from the early and middle Miocene of Germany, respec-tively (Ballmann, 1979), whereas Paractiornis is known from theearly Miocene (MN3) of Nebraska (Wetmore, 1930; Olson andSteadman, 1978). Definite evidence of true Laridae having ex-isted during the Miocene has not been convincingly presentedyet. Larus teruelensis from the late Miocene of Mansuetos, Spain,is known from the distal end of a humerus and was originallyidentified as a scolopacid (Villalta, 1963). It was reassigned to theLaridae by Mlıkovsky (2002). Larus dolnicensis from the earlyMiocene of Dolnice (MN4b), Czech Republic, is represented bya fragmentary distal end of a humerus (Svec, 1980), and waslater allocated to the genus Mioglareola by Mlıkovsky (2000).The existence of bones assignable to either Laridae or Stern-idae, also from the early Miocene of Dolnice, was mentioned byMlıkovsky (1996), but no further information was provided onthese specimens. In North America, Gaviota niobrara, from thelate Miocene of Nebraska, is also known from the distal end ofa humerus (Miller and Sibley, 1941). An alleged species of Larusfrom the middle Miocene of Lee Creek Mine, North Carolina,was described by Olson and Rasmussen (2001), but is only repre-sented by the scapular half of a coracoid. The presence of somegull-like premaxillae from the early–middle Miocene (16–19 Ma)of St. Bathans, New Zealand, was mentioned by Worthy et al.(2007). All of these Miocene fossils are too incomplete to be con-fidently identified at genus—or even family—level, and may in-deed be more basal than assumed.

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DE PIETRI ET AL.—EARLY MIOCENE LARI FROM CENTRAL FRANCE 813

The late Oligocene/early Miocene localities of Saint-Gerand-le-Puy in France have yielded many thousands of avian specimensbelonging to over 40 genera (see Cheneval, 1996), and certainlyrank high in importance concerning early Miocene faunasworldwide. Due to its lacustrine deposits, the Saint-Gerand-le-Puy area has produced an avifauna consisting primarily oftaxa with a preference for aquatic habitats (e.g., Cheneval,1984, 1996). Regarding charadriiforms, only members of theScolopacidae and the Lari have been recorded from this location.The Lari of Saint-Gerand-le-Puy were originally described byMilne-Edwards (1863, 1867–1868) as “Larus” elegans, “L.”totanoides, and the larger “L.” desnoyersii. Because “L.” ele-gans and “L.” totanoides are—according to Milne-Edwards’s(1867–1868:360–361) measurements—significantly differentin hind limb proportions from extant Laridae and Sternidae,Mlıkovsky (2002) proposed to unite them under the genusLaricola, whereas he considered “Larus” desnoyersii as genusincertae sedis. Although it had already been suggested thatthese birds may have been too primitive for inclusion in theLaridae (Ballmann, 1976), until now it has not been determinedwhether this distinction is supported by derived osteologicalfeatures. Laricola elegans has been allegedly also recoveredfrom the early Miocene (MN4b) of Dolnice, Czech Republic(Mlıkovsky, 2002), and members of Laricola have been reportedfrom 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-PuyLari dates back to Alphonse Milne-Edwards in the late 1860s anda revision of these birds is long overdue. The ample avian collec-tion from Saint-Gerand-le-Puy in the Natural History Museumin Basel, Switzerland, has presented us with the possibility of re-evaluating these alleged larid-like specimens.

Despite the vast amount of fossil material, the task of assigningthe Saint-Gerand-le-Puy Lari a phylogenetic placement based onderived osteological traits has remained elusive, and only recentlyhas progress been made towards reconciling the great number ofphylogenetic hypotheses, based both on molecular and morpho-logical data, proposed during the last 30 years for representativesof the Lari, among other Charadriiformes (Mayr, 2011). Here wepresent the first attempt to include fossil Laricola from Saint-Gerand-le-Puy in a phylogenetic context within the Lari, as wellas to provide a descriptive overview and present state of knowl-edge for these fossils. We will further address the issue of theparaphyly of the Sternidae and the conflicting position of Ryn-chopidae.

Institutional Abbreviations—MNHN, Museum Nationald’Histoire naturelle, Paris, France; NMB, Natural History Mu-seum Basel, Switzerland; SMF, Forschungsinstitut Senckenberg,Frankfurt am Main, Germany.

GEOLOGY

The fossil site of Saint-Gerand-le-Puy (Fig. 1) is actuallycomposed of several quarries in the vicinity of the villages ofSaint-Gerand-le-Puy and Montaigu-le-Blin in central France(e.g., Cheneval, 1996). The old collections of the Natural HistoryMuseum Basel (together with others in Paris and Lyon) weregathered in the 19th and early 20th centuries and no direct re-lationship between the finds and their sedimentological contextwas ever recorded so that the exact provenance of the fossilsis difficult to establish. This is not much of a problem for largetaxonomic assessments, because the different quarries in thevicinity of both villages yielded Aquitanian faunas that wereoften ascribed to the MN2 (22.5–20.5 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,

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

in which the lower layers date from the late Oligocene MP29–30(Hugueney et al., 2003). The exact stratigraphic provenanceof the fossils in the old collections is not known in all cases.We cannot confidently refer the studied material to any lateOligocene locality, and a late Oligocene age for some of thematerial is a possibility that cannot be excluded.

Comprehensive sedimentological studies described the geol-ogy of the largest quarry at Montaigu-le-Blin (Bucher et al.,1985) or the formation of the lacustrine calcareous deposits ofthe area (Donsimoni, 1975; Wattine et al., 2003). The generalgeological context of the area (the Limagne sedimentary basin)is that of a subsident basin related to alpine orogeny such asmany 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 thewidespread calcareous sedimentation in a fluvio-lacustrine con-text, associated with stromatolitic bioconstructions.

For Montaigu-le-Blin, Bucher et al. (1985) suggested that mostof the fossil accumulations were found within marls deposited asmud flows between the bioconstructions. These marls also showhigh concentrations of freshwater gastropods, attesting theirlacustrine origin (Helicidae: Bucher et al., 1985; Wattine et al.,2003). The sedimentological succession in Montaigu-le-Blinshows regressive sequences of the lake level inducing facieschanges—from marls to sands and conglomerates—showingan increasing influence of the river system. Widely developedstromatolites often cap these sequences, testifying to the returnof true lacustrine conditions (Donsimoni, 1975; Donsimoniand Giot, 1977; Wattine et al., 2003). Other fossil accumulations

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within fluvio-deltaic calcareous sediments (also rich in freshwatergastropods: Valvatidae and Hydrobiidae to a lesser extent; seeWattine et al., 2003) seem to have preserved only aquatic birds,as far as vertebrates are concerned (Bucher et al., 1985) andagain indicate freshwater lacustrine conditions.

Overall, both the sedimentology and palaeontology at Saint-Gerand-le-Puy testify to shallow freshwater conditions undervarying climatic conditions with more or less humid periodschanging the chemical conditions in the lake (with periods of lowor high oxygenation). These changes are in turn responsible forthe paleontological content (gastropods, ostracods, etc.; Wattineet al., 2003) of the different geologic formations. Bird fossils occurin different formations, either trapped in the marls between thebioconstructions (and in this case, aquatic, shore, or terrestrialspecies are known) or in the fluvio-deltaic calcareous sediments(and here, only aquatic species are known).

MATERIALS AND METHODS

Materials

The fossil specimens are deposited in NMB and, unless other-wise stated, the description of the material is based on this collec-tion. Additional fossil material was consulted at MNHN.

The following skeletons of recent Charadriiformes were avail-able for comparisons. “Sternidae”: Thalasseus bergii (skull;NMB), Chlidonias niger (skull; NMB), Sterna paradisaea (NMBand SMF), S. saundersi (SMF), Anous stolidus (NMB and SMF),and Larosterna inca (SMF). Stercorariidae: Stercorarius parasiti-cus (NMB) and Catharacta skua (NMB). Laridae: Xema sabini(SMF), Rhodostetia rosea (SMF), Larus canus (NMB), L. argen-tatus (NMB), L. fuscus (NMB), L. hyperboreus (NMB), L. mar-inus (NMB), L. ridibundus (NMB), and Rissa tridactyla (NMB).Alcidae: Fratercula arctica (NMB), Alca torda (NMB), Urialomvia (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), Philomachuspugnax (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 findings from sequence data (see Intro-duction), Sternidae sensu stricto refers to a taxon excluding Gy-gis and Anous (see Baker et al., 2007), whereas we have retainedthe use of “Sternidae” for the traditional taxon including Anousand Gygis.

Phylogenetic Analysis

Forty-one characters (Supplementary Data, Appendix 1) werescored for the following genera: Stercorarius, Laricola, Anous,Sterna, Larosterna, Rynchops, Larus, Rissa, Xema, and Rhodoste-tia, with Limosa and Glareola chosen as outgroups. The ma-trix (Supplementary Data, Appendix 2) is largely based on Mayr(2011) from which 32 characters were retained and 9 were newlyadded. All characters were treated as unordered and assignedequal weight. Phylogenetic analyses were performed with theheuristic search modus of NONA 2.0 (Goloboff, 1993) throughthe Winclada 1.00.08 interface (Nixon, 2002), using the followingcommands: hold 10000, hold/10, mult∗1000, and max∗. Unsup-ported nodes were collapsed. Bootstrap values were calculatedunder the following settings: 1000 replicates, with three searchesholding one tree per replicate, and TBR branch swapping with-out max∗. Unless otherwise stated (see Results), characters wereoptimized using the unambiguous option in Winclada. Bremersupports were calculated in TNT 1.1 (Goloboff et al., 2003, 2008)using 10,000 replicates and retaining suboptimal trees by 20 steps.

SYSTEMATIC PALEONTOLOGY

AVES Linnaeus, 1758CHARADRIIFORMES Huxley, 1867

LAROMORPHAE, new taxon

Diagnosis—Least inclusive clade including Laridae, “Stern-idae” (including Anous and Gygis), Rynchopidae, and extinctLaricolidae, n. fam., but excluding Stercorariidae. Characterizedby: marked impressio coracobrachialis (humerus, also in Sterco-rariidae); pronounced fossa musculi brachialis (humerus, exceptAnous). Differing from Stercorariidae in: presence of fossa cau-dalis (mandible); subovate facies articularis acrocoracoidea (fur-cula) extending over nearly entire width of scapus claviculae.

LARICOLIDAE, n. fam.

Type Genus—Laricola Mlıkovsky, 2002.Diagnosis—Small- to medium-sized members of Laromor-

phae, with proportionally longer and more slender legs thanall extant representatives (Fig. 2). Further differing from otherLaromorphae in: shallow facies articularis acrocoracoidea (fur-cula). 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 Mlıkovsky, 2002(Figs. 3–5)

Type Species—Laricola elegans (Milne-Edwards, 1868).Included Species—Laricola elegans (Milne-Edwards, 1868);

L. totanoides (Milne-Edwards, 1868); L. desnoyersii (Milne-Edwards, 1863); L. intermedia, sp. nov.; L. robusta, sp. nov.

Remarks—Milne-Edwards (1867–1868) described Laricolabased on comparisons with few extant Charadriiformes, mostfrom the family Laridae. We present a more complete descrip-tion of the fossil genus comparing it to other Laromorphae,and document previously overlooked anatomical details. Thegenus Laricola presently encompasses two species, Laricolaelegans and L. totanoides (Mlıkovsky, 2002). In this study wehave identified two additional species, and further transferred“Larus” desnoyersii into the taxon Laricola. Anatomical differ-ences within Laricola are, for the most part, of allometric nature.Many postcranial bones are only known for the very abundantLaricola elegans and, unless mentioned otherwise, we base thegenus 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 assignedto this species by their sheer number and gull-like morphology.For the same reasons and unless there existed a conflict in size,we considered it most likely that other gull-like bones that wererepresented by smaller numbers (e.g., pelvis) belong to thisabundant species rather than one of the rarer species present inthe material.

Description and Comparisons

Skull—The skull of Laricola has so far only been incompletelyknown (Milne-Edwards, 1867–1868:pl. 57, fig. 1), with several fea-tures in need of a thorough description. In our material thereare two skulls, an exceptionally well-preserved and almost com-plete one (NMB s.g.18810; Fig. 3A, E–F, H) and a cranium (NMBs.g.18812; Fig. 3P, R–T), which differ marginally in size and mi-nor morphological features. Overall, the complete skull resem-bles Larus minutus in proportions and shape. The beak is straightand intermediate between L. minutus and Sterna paradisaea. Thetip, contrary to the condition found in Laridae, is barely de-curved, though not as straight as in Sternidae s.s., reflecting bestthe condition found in Anous stolidus. The distal portion of the

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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 ele-gans and the extant taxa Stercorarius parasiticus, Anous stolidus, Sterna paradisaea, Larosterna inca, Larus ridibundus, Rissa tridactyla, and Larusminutus. 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 Gohlichand 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 areonly slightly longer than those of S. paradisaea. These tend to beshorter in “Sternidae,” with a proportionally longer symphysisof the rostrum maxillare, whereas they are very long, extendingthroughout most of the length of the beak in Laridae and Sterco-rariidae (Fig. 3F–G). Unlike in Laridae, the fossae glandularumnasales are shallow.

The os lacrimale (Fig. 3A) is not as wide as in Sternidae s.s. andRynchopidae and is thus in agreement with the condition in Lar-idae and Anous. The small anterior process (see Cracraft, 1968)that is characteristic of the “Sternidae” is not present in this spec-imen, although due to its fragility it is unlikely that it would havebeen preserved.

The os ectethmoidale of NMB s.g.18810 (Fig. 3M) has a distinc-tive shape with a pointed tip and concave ventral border, match-ing the condition in A. stolidus and other terns (e.g., S. paradis-aea). In NMB s.g.18812 (Fig. 3P), the ectethmoid is broader andlacks the concave ventral margin, as in Stercorariidae.

The processus maxillopalatinus forms a wide and deep, later-ally concave trough, as in A. stolidus and some species of the Ster-corariidae (Fig. 3H–J). In Laricola, the maxillopalatine processis divided by a relatively wide bar of bone connecting its dorsaland ventral margins, and a similar condition is found in Sterco-rarius parasiticus (Fig. 3J). In Anous (Fig. 3I) the maxillopalatineprocess is laterally closed except for a round caudal foramen. InSternidae S.S., Laridae, and Rynchopidae, the overall shape ofthe maxillopalatine process is less like that of a deep trough, be-ing much flatter. In Laridae (Fig. 3L), the maxillopalatine pro-cess is widely open with one or more thin bars of bone extending

along the bottom of the trough. In Sternidae and Rynchopidae, itis ‘fenestrated’ by thin bars of bone (Fig. 3K).

In NMB s.g.18810, the processus postorbitales are worn, butthey show no indication of having been well developed. InNMB s.g.18812, they project laterally and caudally and areabout the size of those of Larus minutus, albeit broader (Fig.3R–T). A maxillopalatine strut A in the sense of Zusi and Jehl(1970) is absent in Laricola. The processus laterales parasphe-noidales (Fig. 3T) are well developed and ventrally protruding,and the processus suprameaticus is not longer than the proces-sus zygomaticus (Mayr, 2011), both features in accordance withthe condition in Laridae, Rynchopidae, and “Sternidae.” As inother Lari except the Turnicidae, processus basipterygoidei areabsent.

The fossae temporales are well marked, although not as en-hanced as in some terns (e.g., Thalasseus bergii and S. paradisaea)and Laridae (e.g., Larus ridibundus). In these taxa, they almostreach 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. 3V–W) assignable to Laricolaelegans (MNHN Av.4847) is present in the MNHN collection.It lacks the well-developed fenestra caudalis, a condition sharedwith stercorariids and Anous, which indicates that its presence ingulls, skimmers, and some terns is plesiomorphic for Laromor-phae. Also, a fossa caudalis, which is absent in skuas, is presentin Laromorphae.

Vertebrae—Vertebrae of Laricola have not been previouslydescribed. In our material, only one cervical vertebra is likelyto belong in Laricola, and does not differ from those within

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Laromorphae. A couple of well-preserved thoracic vertebrae arealso present within the material, which are indeed opisthocoelousand pleurocoelous. Opisthocoelous thoracic vertebrae are alsopresent in Alcidae, Stercorariidae, and Laromorphae, whereaspleurocoelous vertebrae are absent in Stercorariidae and Alcidae(e.g., Mayr, 2011).

Coracoid—Our material includes numerous complete cora-coids, two of which possess an intact processus lateralis(Fig. 5A–B). In Laricola elegans and L. totanoides, this bonebears a close resemblance to the coracoid of the small gullLarus minutus. The pneumatic recess under the facies articularisclavicularis, characteristic of most gulls (Fig. 5E), is neitherpresent in Laricola elegans nor L. totanoides. A coracoid of analleged Miocene gull lacking this feature was described by Olsonand Rasmussen (2001), who hypothesized that this might be aplesiomorphic trait of gulls. Indeed, this pneumaticity is absentfrom most, but not all, specimens of Laricola, and is thus avariable trait (it is present in Laricola desnoyersii, for example).

Furcula—In Alcidae, Sternidae, Rynchopidae, and Laridae,the extremitas omalis of the furcula exhibits a distinct subovatefacies articularis acrocoracoidea, almost extending over the en-tire width of the scapus claviculae (Mayr, 2011). This feature isalso present in Laricola (Fig. 5G–H). Nonetheless, the articula-tion facet is much shallower in the fossil than in the examinedrecent species, which is likely to be the plesiomorphic conditionfor Laromorphae. The scapula of Laricola matches thoroughlythat of recent Laromorphae.

Sternum—There are several fragments of the sternum of Lari-cola present in the collection. Among these is a corpus sterni witha preserved rostral portion, whose preservation is so poor thatbarely any details are recognizable. One rostral portion of thesternum clearly shows the shape of the apex carinae (Fig. 5L),which is pointed and cranially projecting, as in Anous and someLaridae.

Humerus—A large number of complete and partial humeriin the collection can be referred to Laricola. A second fossapneumotricipitalis is absent (Fig. 4A, C, H, K, N), a condi-tion that within Laromorphae is only also present in Anous(Fig. 4X). Strauch (1978) mentioned that the absence of a sec-ond fossa pneumotricipitalis is a variable trait both in Anous andRynchops, but we are unable to support this statement. The sul-cus nervi coracobrachialis is very well marked in Laricola, a traitthat is variable within Laridae and Sternidae. In agreement withrecent 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-markedtubercle on the cranial surface of the distal end, between thebase of the tuberculum supracondylare dorsale and the fossa m.brachialis (Fig. 4V), for the attachment of m. supinator, whichis less marked in the rest of the Laromorphae. The humerus ofLaricola otherwise matches that of other Laromorphae (for moredetails see Mayr [2011], Chu [1998], and Strauch [1978]).

Ulna—Ulnae are fairly uniform throughout Laromorphae.The tuberculum ligamenti collateralis ventralis (Fig. 5M) is welldeveloped throughout this group, but in most Laridae (exceptLarus hyperboreus) and Laricola it is more pronounced ventrally

than in the examined Anous and Stercorariidae; in both thesetaxa the tuberculum is also slightly more elongated. In Rynchops,the tuberculum ligamenti collateralis ventralis is not developed atall.

Carpometacarpus—Unlike in recent Laridae, the car-pometacarpus of Laricola bears a small processus inter-metacarpalis (Fig. 5U). Otherwise this bone does not differfrom that of Laromorphae. Milne-Edwards (1867–1868:354)mentioned that the spatium intermetacarpale is narrower inLaricola and Laridae than in terns. However, this is true only forvery small terns, such as Sterna saundersi, and is probably a sizerelated effect.

Other Wing Bones—The radius of Laricola shows no peculiar-ities. In agreement with the condition found in Laromorphae andStercorariidae, the phalanx proximalis digiti majoris is bifenes-trated (Fig. 5V–X).

Pelvis—Although no complete pelvis is known from theNMB collection (Fig. 5Z–AA), a specimen illustrated by Milne-Edwards (1867–1868:pl. 56, fig. 22) shows that the pelvis ofLaricola possesses two rows of foramina transversaria, and thusmatches that of smaller Laridae, such as Larus minutus, Xemasabini, and some Sternidae s.s. The one well-preserved partialpelvis present in the NMB collection shows that the recessus cau-dalis fossae (Fig. 5Y) is present in Laricola, and this recess is alsopresent in Laridae and Anous stolidus (contra Strauch, 1978). Itis absent in Sternidae (except Sterna caspia) and Rynchopidae.

Femur—The femur of Laricola (Fig. 5BB–CC) agrees withthat of extant Laridae and Sternidae. There is a prominence justproximal of the condylus medialis (Fig. 5CC), which is also foundin Sternidae and Rynchopidae, and is variable within Laridae,being present in some small species such as Larus minutus, butabsent in Stercorarius and Anous. However, this prominenceis developed as a ridge in Laricola and Laridae, whereas it isindeed a prominent tubercle in Sternidae and Rynchopidae. Thetuberculum musculi gastrocnemialis lateralis is well developed,as in Laromorphae.

Tibiotarsus—The legs of Laricola are long and slender, longerthan in all examined recent Laromorphae (Fig. 2). The tibio-tarsi of Laromorphae and Stercorariidae are very uniform inappearance, and that of Laricola matches this morphology(Fig. 5DD–FF).

Tarsometatarsus—The hypotarsus of Laricola exhibits a veryprominent crista medialis hypotarsi and two distinct cristae in-termediae hypotarsi (Fig. 5KK–NN). This configuration agreeswith that of Laridae, although the shape of the hypotarsus isvery variable within Laromorphae, and some variation is also ob-served in Laricola. In Sternidae this sulcus is actually a canal, andsome terns such as Thalasseus bergii and Anous stolidus have twoclosed canals, although given the current phylogeny of the group(Baker et al., 2007), these are unlikely to be homologous. Fur-thermore, the hypotarsus of the tern Larosterna resembles thatof gulls and Laricola. One tarsometatarsus (Fig. 5NN) assigned toLaricola elegans bears a closed canal, but unlike in the Sternidae,in which the canal is centrally positioned, the orientation matchesthat of the sulcus in other Laricola specimens. The closed canalcould thus be the result of ossification of cartilage that closes the

← FIGURE 3. Skulls and mandible of Laricola elegans in comparison to extant Lari. A–D, 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). F–G, right lateralview of the skulls of Laricola elegans (F, NMB s.g.18810) and Larus canus (G). H–Q, not to scale, details of the skull of Laricola elegans (H, M, NMBs.g.18810; P, NMB s.g.18812) in comparison to Anous stolidus (I–N), Stercorarius parasiticus (J–O), Sterna paradisaea (K), and Larus canus (L–Q).R–S, skull of Laricola elegans (NMB s.g.18812) in dorsal (R) and ventral (S) views. T–U, occipital view of the skulls of Laricola elegans (T, NMBs.g.18812) and Larus canus (U). V–W, 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 five species of Laricola (A–P) and the two species of Sternalara (Q–U) in comparison to extant Lari (V–AA). A–D, lefthumerus 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).E–G, left humerus of Laricola totanoides (NMB s.g.2180) in caudal view (E), cranial view (F), and distal end in cranial view (G). H–I, right humerusof Laricola robusta, sp. nov. (NMB s.g.18465), in caudal (H) and cranial (I) views. J–K, right humerus of Laricola intermedia, sp. nov. (holotype,MNHN SG 13744), in cranial (J) and caudal (K) views. L–P, humeri of Laricola desnoyersii, left humerus (MNHN Av.4059) in cranial (L) andcaudal (M–N) views, and distal left humerus (NMB MA 1106) in cranial (O) and caudal (P) views. Q–R, right humerus of Sternalara milneedwardsi,sp. nov. (holotype, MNHN SG13742), in cranial (Q) and caudal (R) views. S–U, 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). V–X, right humerusof Anous stolidus in cranial (V) and caudal (W–X) views. Y–Z, 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 desnoy-ersii (see below) also possesses this feature (Fig. 5LL).

The foramen vasculare distale (Fig. 5II) is situated in a widerand better-marked fossa than in extant Laridae. This fossa is alsowell marked in the examined Stercorariidae, “Sternidae,” andRynchopidae. In Laricola the trochlea metatarsi III projects far-ther distally than in all other examined Laromorphae, althoughsome gulls (e.g., Larus marinus) do resemble Laricola in this

aspect. Rynchopidae and Sternidae, on the other hand, have atrochlea metatarsi III that does not project much farther distally.

LARICOLA ELEGANS (Milne-Edwards, 1868)(Figs. 3–5, Table 1)

Larus elegans Milne-Edwards, 1868:350–357, pl. 56, figs. 11–29;pl. 57, figs. 1–11; pl. 58, fig. 1, reconstructed skeleton (originaldescription).

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Laricola elegans (Milne-Edwards, 1868): Mlıkovsky, 2002:136(new combination).

Lectotype—MNHN Av.4134 (left humerus).Referred Specimens—All specimens from NMB: NMB

s.g.18810 (skull, with beak), NMB s.g.18812 (cranium), NMBs.g.21782, NMB s.g.1881, and NMB MA.1379 (partial skulls),MNHN Av.4847 (mandible, fragment of right ramus), NMBs.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), NMBs.g.18816 (left coracoid), NMB s.g.18827 (right coracoid),NMB s.g.13027 (left coracoid), NMB s.g.13030 (right cora-coid), NMB s.g.13028 (left coracoid), NMB s.g.10608 (rightcoracoid), NMB s.g.18824 (left coracoid), NMB s.g.18815 (rightcoracoid), NMB s.g.18825 (left coracoid), NMB s.g.17023 (lefthumerus), NMB s.g.18885 (right humerus), NMB s.g.21935 (lefthumerus), NMB M.A.1091 (right humerus), NMB s.g.21936 (lefthumerus), NMB s.g.18878 (left humerus), NMB s.g.21933 (lefthumerus), NMB s.g.18987 (right ulna), NMB s.g.18983 (leftulna), NMB s.g.18995 (right ulna), NMB s.g.12951 (right ulna),NMB s.g.16139 (left ulna), NMB s.g.16579 (left ulna), NMBs.g.15913 (right ulna), NMB s.g.15665 (left carpometacarpus),NMB MA 3178 (left carpometacarpus), NMB s.g.16183 (left car-pometacarpus), NMB M.A.2589 (left carpometacarpus), NMBM.A.2599 (left carpometacarpus), phalanx proximalis digiti ma-joris (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), NMBs.g.16806 (left femur), NMB s.g.19127 (left tibiotarsus), NMBs.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 tibio-tarsus), 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 tar-sometatarsus).

Locality and Age—Antoingt (late Oligocene, MP 25), France(Mlıkovsky, 2002); Gannat (late Oligocene, MP 30), France(Mlıkovsky, 2002); Saint-Gerand-le-Puy (late Oligocene?–earlyMiocene, MN2), Allier, France.

Measurements—Skull, NMB s.g.18810: 61 mm long; maximumwidth excluding processus postorbitales: 17.24 mm; height in-cluding processus laterales parasphenoidales: 17.0 mm. Cranium,NMB s.g.18812: 30.36 mm long (as is; Fig. 3); maximum width ex-cluding processus postorbitales: 17.90 mm; height including pro-cessus laterales parasphenoidales: 16.1 mm.

For postcranial elements see Table 1.Remarks—Hundreds of postcranial bones are assignable

to Laricola elegans, in the collections of both MNHN andNMB. These constitute a continuum varying greatly in size(Table 1), but show no obvious anatomical differences. Basedon the postcranial skeleton alone it is impossible to discernbetween potential species within these ranges. Because of thelarge amount of fossil material (hundreds of bones), we havereferred 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 havebeen figured by Milne-Edwards (1868:pl. 57, figs. 2–3).

LARICOLA TOTANOIDES (Milne-Edwards, 1868)(Figs. 4–5, Table 1)

Larus totanoides Milne-Edwards, 1868:358–361, pl. 57, figs. 12–17(original description).

Laricola totanoides (Milne-Edwards, 1868): Mlıkovsky, 2002:137(new combination).

Lectotype—MNHN Av.4004 (right humerus). Illustrated byMilne-Edwards, 1867–1868:pl. 57, figs. 16–17.

Referred Specimens—MNHN Av.9455 (right coracoid), NMBs.g18809 (right coracoid), NMB s.g.2180 (left humerus), NMBs.g.17047 (distal right humerus), NMB s.g.12946 (left ulna),NMB s.g.18994 (left ulna), NMB s.g.12947 (right ulna), NMBM.A.1602 (right ulna), NMB M.A.5181 (right carpometacarpus),NMB s.g.19012 (left carpometacarpus), NMB s.g.12959 (rightcarpometacarpus), NMB M.A.1098 (left femur), NMB s.g.21796(right tibiotarsus).

Locality and Age—Saint-Gerand-le-Puy, Allier, France; lateOligocene?–early Miocene, MN2.

Differential Diagnosis—All skeletal elements between 10%and 15% larger than the average for Laricola elegans.

Remarks—The criteria chosen by Milne-Edwards (1867–1868)to distinguish between Laricola elegans and Laricolatotanoides—namely size and thickness of the bones—arelargely unwarranted since these fall within a continuum(Table 1), and are therefore not discriminatory. From theoriginal material examined by Milne-Edwards (1867–1868),only the ulnae and the illustrated humerus are large enough tobe considered a different species from Laricola elegans. Theseclearly fall outside this range (Table 1), and we thus considerthese specimens to be the ‘true’ Laricola totanoides, even thoughwe were unable to find other distinguishing morphologicalfeatures. This reasoning is nevertheless supported by the smallnumber of bones of this size that are present in the collection.Given that there are no perceivable anatomical differences forall other bones falling within this continual size range, we havedecided to unite all of these under Laricola elegans.

We further designate as lectotype of L. totanoides the humerus(MNHN Av.4004), that was figured by Milne-Edwards (1868:pl.57, figs. 16–17), and labeled as type in the MNHN collection. Oneof the coracoids that we refer to this species was originally as-signed to Totanus lartetianus, and it was the specimen illustratedby Milne-Edwards (1867–1868:pl. 63, figs. 12–16) for this taxon.However, the presence of a foramen nervi supracoracoidei, aswell as a dorsoventrally widened medial portion of the faciesarticularis sternalis—both features characteristic of Laromor-phae and Stercorariidae—indicate that this bone is evidently notthat of a scolopacid. Indeed, it matches exactly the coracoid ofLaricola. Because it is marginally above the size range known forLaricola elegans (Table 1), we have referred it to L. totanoides.The carpometacarpus, tibiotarsus, and femur are only tentativelyassigned to this species based on size.

LARICOLA DESNOYERSII (Milne-Edwards, 1863)(Figs. 4–5, Table 1)

Larus desnoyersii Milne-Edwards, 1863:161 (original descrip-tion).

Larus desnoyersii Milne-Edwards, 1868:344–349; pl. 54, figs.15–26; pl. 56, figs. 1–10.

Original Material—MNHN Av.4044 (left coracoid), MNHNAv.4045 (right coracoid), MNHN Av.4046 (right coracoid; fig-ured by Milne-Edwards, 1967: pl. 54, figs. 22–23), MNHNAv.4048 (coracoid), MNHN Av.4050 (humerus, juvenile),MNHN Av.4061-67 (humeri), MNHN Av.4074 (ulna with dis-tal end missing), MNHN Av.4095–98 (phalanx proximalis dig-iti majoris), MNHN Av.4106 (left tibiotarsus; juvenile), MNHNAv.4107 (left tibiotarsus; juvenile), MNHN Av.4108 (right tibio-tarsus, juvenile), MNHN Av.4099 (left tarsometatarsus), MNHNAv.4100 (right tarsometatarsus), MNHN Av.4101 ( right tar-sometatarsus).

Newly Referred Specimens—NMB s.g.19325 (left coracoid, ju-venile), NMB s.g.13024 (left coracoid, juvenile), NMB M.A.1106

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(left humerus, proximal end missing), NMB s.g.7276 (left car-pometacarpus, juvenile), NMB Gn.433 (left tarsometatarsus).

Locality and Age—Saint-Gerand-le-Puy, Allier, France; lateOligocene?–early Miocene, MN2.

Differential Diagnosis—Approximately 30% larger than Lari-cola totanoides. Differs from Laricola elegans and L. totanoides inhaving greater pneumaticity (i.e., a pneumatic recess) under thefacies articularis clavicularis of the coracoid (Fig. 5D).

Remarks—Based on comparisons with the published illustra-tions, Olson (1985:180) hypothesized that the humerus of L.desnoyersii has a pneumatic foramen in the fossa pneumotricipi-talis, and regarded this feature as potential evidence for affinitiesbetween L. desnoyersii and Stercorariidae. The bone is broken inthis region, which, without proper examination of the specimenitself, gives the appearance of a pneumatic foramen, especially inMilne-Edwards’s illustration. Our inspection of the original ma-terial in MNHN showed this hypothesis to be erroneous becausethere is no pneumatic foramen present in the fossa pneumotricip-italis, which is nevertheless very pronounced (Fig. 4N).

We have referred “Larus” desnoyersii to the taxon Laricolabecause we were unable to find any morphological features thatare absent in Laricola. Although considerably larger than theother two species of Laricola previously described, such size dif-ferences within a genus are not unusual within Laromorphae, forinstance in the extant genus Larus, the smallest species (L. min-utus) and one of the largest (L. marinus) differ in size by over60%.

The first 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 didhe figure the specimen, and an illustration of a distal end of ahumerus is also absent in Milne-Edwards (1867–1868). Thereare several distal ends of humeri attributable to L. desnoyersii inMNHN (Av.4062–64), but because of the aforementioned lack ofinformation, no holotype can be determined among these spec-imens. Mlıkovsky (2002) considered the specimen mentionedin Milne-Edwards (1863) to be the lectotype for the species.However, we note that with no catalog number, a designation ofthis bone as a lectotype does not meet the requirements of theInternational Code of Zoological Nomenclature, which demanda type to be specified. If a lectotype were to be chosen, then wepropose specimen MNHN Av.4059, a right humerus, which wasfully illustrated and described by Milne-Edwards 1867–1868 (pl.554, figs. 24–25).

The scapula and the pelvis referred to L. desnoyersii by Milne-Edwards (1867–1868:pl. 56, figs. 1–2 and 5–7) do not belong toCharadriiformes. The single carpometacarpus in MNHN in goodcondition assigned to Laricola desnoyersii is likely to belong tothe Scolopacidae.

Description and Comparisons

Coracoid—We have assigned two juvenile coracoids fromNMB to Laricola desnoyersii based on size. Unfortunately manyfeatures are worn and few details can be recognized. Thespecimens in MNHN are much better preserved and, apartfrom a better-developed recess distal to the extremitas oma-lis (pneumaticity under the facies articularis clavicularis, seeFig. 5D)—which could be size related, are not different fromother Laricola.

Humerus—Although considerably larger, the humerus of Lari-cola desnoyersii matches that of Laricola elegans in proportions.The fossa m. brachialis of L. desnoyersii is not as deep as in mostextant species of Larus, thus resembling Laricola elegans. Thewell-marked tubercle on the cranial surface of the distal end, be-tween the base of the tuberculum supracondylare dorsale and thefossa m. brachialis, is present.

Ulna—Only one ulna present in the Saint-Gerand-le-Puy col-lection in MNHN can be assigned to Laricola desnoyersii. Allother ulnae in this collection either belong possibly to Scolopaci-dae, or to Anseriformes.

Carpometacarpus—Carpometacarpus NMB s.g.7276 (Fig. 5Q)is only tentatively assigned to this taxon based on size. Its edgesare worn and most of the distal end is missing. Although theone complete specimen from MNHN is indeed larger (43.4 mm),MNHN Av.4092 bears a dorsally rounder condylus medialis andmay belong to the Scolopacidae. Also, the synostosis metacarpidistalis is proportionally longer than in stercorariids and Laro-morphae, thus resembling some Scolopacidae. The two otherspecimens from MNHN (Av.4091 and Av.4093) are too fragmen-tary for a reasonable identification. Unfortunately, the distinctivefeatures of the distal end of NMB s.g.7276 are missing. The pro-cessus extensorius, however, is intact and resembles that of Lari-cola.

Other Wing Bones—Only fragmentary specimens of the pha-lanx proximalis digiti majoris are present in the MNHN collec-tion. However, it is clear that this bone was, as in all Laromor-phae and Stercorariidae, bifenestrated. As in Laricola and other

← FIGURE 5. Postcranial skeletal elements of Laricola and Sternalara gen. nov. A–B, right coracoid of Laricola elegans (NMB s.g.18825) inventral (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; G–H, caudal aspectof right scapus claviculae (furcula) of Laricola elegans (NMB s.g.16378); I, caudal aspect of left scapus claviculae of Larus hyperboreus; J, right scapulaof Laricola elegans (NMB s.g.19575) in lateral view; K–L, sternum of Laricola elegans (NMB s.g.15879); M, ventral aspect of left ulna of Laricolatotanoides (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); R–S, right carpometacarpus of Laricola totanoides (NMB M.A.5181) in dorsal (R) and ventral (S) views; T–U, left carpometacarpusof Laricola elegans (NMB M.A.3178) in ventral (T) and dorsal (U) views; V–W, left phalanx proximalis digiti majoris of Laricola elegans (NMBs.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; Z–AA, 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; DD–EE, left tibiotarsus of Laricola elegans (NMB s.g.19127) in cranial (DD), proximal (EE, not to scale), and medial (FF, not toscale) views; GG–HH, left tarsometatarsus of Laricola desnoyersii (Gn.433) in plantar (GG) and dorsal (HH) views; II–JJ, right tarsometatarsus ofLaricola 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); OO–QQ, right carpometacarpus of Laromorphae indet. (NMB s.g.19007) indorsal (OO, QQ, not to scale) and ventral (PP) views; RR–SS, 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, cristamedialis 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 ligamenticollateralis ventralis.

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TABLE 1. Skeletal measurements (in mm) of the long bones of three Laricola species: L. elegans, L. totanoides, and L. desnoyersi.

Laricola elegans Laricola totanoides Laricola desnoyersi

n Range Mean SD n Range Mean n Range Mean

CoracoidMaximum length 14 19.7–22.4 21.0 0.8 1∗ 22.8 — 1 32.0 —

(21.3) 1 23.3 — (30.0)(22.0)

Medial length 14 17.3–19.0 18.4 0.6 1∗ 20.9 — 1 29.1 —1 19.8 —

Distal width 14 6.4–7.9 7.1 0.4 1∗ 8.0 — 1 10.0 —(6.9) 1 7.7 — (11.0)

(7.5)Width of processus acrocoracoideus 14 3.1–3.8 3.5 0.2 1∗ 3.4 — 1 5.5 —

1 4.1 —Humerus

Maximum length 43 47.0–53.9 50.0 2.0 1 ∼57(est) — 2∗ 84.3–85.3 84.8(48.4) (51.0) (85.3)

Proximal width 37 8.9–10.7 9.5 0.4 — — — 2∗ 14.5–15.6 15.1(9.9) (10.0)

Distal width 43 6.4–7.6 6.9 0.3 2 7.6–8.0 7.8 1 11.2 —(7.0) (7.1) 2∗ 10.4–11.3 10.9

Minimum width of shaft 43 2.7–3.3 3.0 0.6 1 3.6 — 1 5.3 —(3.0) (3.0) 2∗ 4.4–4.9 4.7

UlnaMaximum length 51 51.4–62.9 56.4 3.4 4 65.1–66.3(est) 65.5 1∗ ∼94.5(est) —

(56.9) (65.0) (96.3)Proximal width 51 5.1–6.2 5.6 0.3 4 5.6–6.5 6.1 1∗ 9.6 —

(5.9) (6.7)Distal width 51 43.9–5.5 4.7 0.4 4 4.8–5.5 5.2 — — —

(4.3) (4.4)Minimum width of shaft 48 2.4–3.0 2.7 0.2 4 2.8–3.0 2.9 1∗ 4.6 —

(2.3) (2.9)Carpometacarpus

Maximum length 40 27.9–32.0 30.2 1.4 3 33.1–34.6 34.0 1 39.4 —(30.0) (30.0) (44.0)

Proximal depth 40 6.4–7.9 6.9 0.35 3 7.4–7.5 7.4 1 8.9 —(3.2) (3.4) (9.6)

Distal depth 38 3.6–4.5 4.1 0.23 2 4.6–4.9 — — (6.0) —(3.2) (3.4)

FemurMaximum length 34 22.5–27.1 24.4 1.3 1 29.1 — — — —

(22.2) (25)Proximal width 34 4.2–5.2 4.6 0.3 1 5.1 — — — —

(4.0) (4.7)Distal width 34 4.2–5.2 4.7 0.3 1 5.4 — — — —

(4.4) (4.9)Minimum width of shaft 34 1.8–2.3 2.0 1.1 1 2.3 — — — —

(1.8) (1.8)Tibiotarsus

Length from facies articularis 17 45.1–51.9 48.8 2.8 1 54.0 — 1∗ 72.3 —(52.0, maximum

length)(—)

Proximal width 16 3.5–5.1 3.8 0.4 1 5.4 — — — —(4.9) (—)

Distal width 15 3.9–4.4 4.1 0.1 1 4.4 — 2∗ 5.8–6.3 6.1(3.9) (4.5) (6.1)

Minimum width of shaft 18 1.6–2.0 1.8 0.1 1 2.2 — 2∗ 2.9–3.6 3.3(1.9) (1.9)

TarsometatarsusMaximum length 24 31.8–37.7 34.3 2.0 — — — 1 49.1 —

(34.1) (37.2) 2∗ 44.1–45.7 44.9(45.7)

Proximal width 23 4.1–5.0 4.6 0.2 — — — 1 7.1 —(4.7) (5.0) 2∗ 6.4–7.9 7.2

Distal width 23 4.0–4.6 4.3 0.2 — — — 1 6.9 —(4.4) (4.5) 2∗ 6.1–6.4 6.3

Minimum width of shaft 24 1.4–1.9 1.6 0.1 — — — 1 2.6 —(1.7) (2.0) 2∗ 2.6 2.6

In parentheses are the measurements provided by Milne-Edwards (1867–1868). Note that for all bones except the ulna, the measurements listed byMilne-Edwards for Laricola totanoides overlap with those of Laricola elegans. All bones are from NMB, except those marked by an asterisk, whichare from MNHN.

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TABLE 2. Measurements (in mm) of the skeletal elements known for Laricola intermedia, L. robusta, Sternalara minuta, and S. milneedwardsi.

Laricola intermedia Laricola robusta Sternalara minutaSternalara

milneedwardsi

n Range Mean n Range Mean SD n Range Mean SD n Range

HumerusMaximum length 1 75.3 — 9 47.5–50.9 48.8 1.1 6 44.6–47.5 46.2 1.0 1∗ 133.9Proximal width 1 12.9 — 9 6.7–7.4 7.1 0.3 6 8.5–8.8 8.7 0.1 1∗ 25.6Distal width 1 9.7 — 9 9.8–11.0 10.3 0.4 6 6.2–6.7 6.5 0.2 1∗ 18.1Minimum width of shaft 1 4.4 — 9 3.0–3.4 3.2 0.2 6 2.6–3.0 2.8 0.2 1∗ 7.7

UlnaMaximum length — — — 3 53.0–56.0 54.6 1.5 4 49.7–50.7 50.3 0.4 — —Proximal width — — — 3 5.4–6.2 5.2 0.3 4 5.0–5.6 5.2 0.3 — —Distal width — — — 3 4.7–5.0 4.9 0.2 4 4.2–5.0 4.7 0.4 — —Minimum width of shaft — — — 3 2.9–3.1 3.0 0.1 4 2.4–2.9 2.7 0.2 — —

FemurMaximum length 2 31.5–31.6 31.6 — — — — — — — — — —Proximal width 2 7.5–7.8 7.7 — — — — — — — — — —Distal width 2 5.9–6.0 6.0 — — — — — — — — — —Minimum width of shaft 2 2.8–2.9 2.9 — — — — — — — — — —

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

Laromorphae, the processus internus indicis is strongly elongatedcaudally (Fig. 5X).

Tibiotarsus—Unfortunately, the tibiotarsi in MNHN be-longing to L. desnoyersii are juveniles and/or badly preservedspecimens. The distal end closely resembles that of otherLaromorphae, exhibiting a wide sulcus extensorius.

Tarsometatarsus—Corresponding with the condition in Lari-cola, the tarsometatarsus of L. desnoyersii is proportionallyvery long. MNHN Av.4101 was figured by Milne-Edwards(1867–1868:pl. 54, figs. 16–18) and has a hypotarsus similar to thatof Laricola elegans (Fig. 5KK), exhibiting an elongated crista me-dialis hypotarsi, two crista intermedia hypotarsi, and no closedcanal for the tendon of the musculus digitorum longus. The hy-potarsus of MNHN Av.4099 (Fig. 5LL) differs from this specimenin that it bears a closed canal for the tendon of the musculus digi-torum longus, as is the case for one specimen we have assigned toLaricola elegans (Fig. 5NN). It should be noted that slight vari-ations in length and orientation of the cristae of the hypotarsusare seen throughout the genus Larus, and indeed within Laricola(which partly may be due to the preservation of these delicatestructures). NMB G.n.433 (Fig. 5GG–HH) is only slightly largerthan the ones from MNHN, and unfortunately the hypotarsus isworn, but matches Laricola in all other features, as do the speci-mens from MNHN.

LARICOLA INTERMEDIA, sp. nov.(Figs. 4–5, Table 2)

Holotype—MNHN SG.13744 (right humerus).Type Locality and Age—Saint-Gerand-le-Puy, Allier, France;

late Oligocene?–early Miocene, MN2.Differential Diagnosis—Species of Laricola, which is distin-

guished from L. elegans and L. totanoides by its much greatersize, being, however, smaller than L. desnoyersii (see Table 2).

Measurements—See Table 2.Referred Specimens—NMB M.A.5197 (left femur), NMB

M.A.5198 (left femur).Etymology—The species name Laricola intermedia alludes to

the intermediate size between Laricola elegans/L. totanoides andL. desnoyersii.

Remarks—The femora are only tentatively assigned to thistaxon based on size; they are smaller than what would be pre-dicted for Laricola desnoyersii.

Description and Comparisons—The humerus MNHNSG.13744 (Fig. 4J–K) is approximately 15% smaller thanthat of Laricola desnoyersii. A second fossa pneumotricipitalis is

absent and, as in Laricola desnoyersii, the fossa pneumotricipi-talis that is present is very pronounced, but bears no pneumaticforamen. The sulcus nervi coracobrachialis is well marked; otherfeatures of the proximal end do not differ from those of otherLaromorphae. The fossa musculi brachialis is deep, and thewell-marked tubercle on the cranial surface of the distal end,between the base of the tuberculum supracondylare dorsale andthe fossa m. brachialis, is present.

The femora bear a well-marked tuberculum musculi gastrocne-mialis lateralis, but the prominence just proximal of the condylusmedialis that is present in some Laromorphae is not evident be-cause the area is worn.

LARICOLA ROBUSTA, sp. nov.(Figs. 4–5, Table 2)

Larus totanoides Milne-Edwards, 1868:358 (original description).

Holotype—MNHN Av.3777 (right humerus, labeled as Lari-cola totanoides).

Type Locality and Age—Saint-Gerand-le-Puy, Allier, France;late Oligocene?–early Miocene, MN2.

Differential Diagnosis—Member of Laricola with stout bones;differing from other Laricola in: enlarged crista bicipitalis andcrista deltopectoralis, broader and better-developed processussupracondylaris dorsalis, proportionally shorter shaft of humerus(see Fig. 4, compare A–B with H–I).

Measurements—See Table 2.Referred Specimens—NMB s.g.18887 (left humerus), NMB

s.g.16467 (right humerus), NMB s.g.16514 (left humerus), NMBPh.2013 (left humerus), NMB s.g.16554 (left humerus), NMBs.g.7209 (right humerus), NMB s.g.7206 (right humerus), NMBs.g.21 (right humerus), NMB s.g.18465 (right humerus). The fol-lowing ulnae are only tentatively assigned to this taxon: NMBM.A.2539 (left ulna), NMB s.g.16581 (right ulna), NMB s.g.20(right ulna).

Etymology—Laricola robusta is named after its stout and ro-bust skeletal elements.

Remarks—The humeri that we have assigned to this taxonlie approximately in the mid-size range of Laricola elegans (seeTable 1 and 2). The proportions of the shaft are unlike those forother Laricola, in which the shaft is proportionally longer than inthis species (Fig. 4H–I). Because of the wide size ranges (Table1) for all long bones of Laricola elegans, it is difficult to separatematerial based on ‘stoutness,’ and for that reason the referredulnae can only be tentatively allocated to this taxon. In the

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MNHN collection these elements are labeled ‘Larus totanoides,’to which they were assigned by Milne-Edwards (1867–1868), whodescribed Larus totanoides as being much stouter than Laricolaelegans in all bones. As we have shown, much of this differenceis attributed to a wide size range of L. elegans, but in the case ofthe humerus, the different proportions allow for the constructionof a new species. It is, nevertheless, not understandable whyMilne-Edwards would have referred these specimens to “Larus”totanoides when the illustration of this taxon clearly depictsthe much larger incomplete humerus already mentioned in thespecies section for Laricola totanoides.

Description and Comparisons—The humeri (Fig. 4H–I) arestouter than those of Laricola elegans, with a broader and better-developed processus supracondylaris dorsalis. The fossa musculibrachialis is deep, and the bone bears the well-marked tubercleon the cranial surface of the distal end, between the base of thetuberculum 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 betterdeveloped than in the other specimens of Laricola. The shaftis proportionally shorter than in other Laricola (compare withFig. 4A–B). The ulnae (Fig. 5O) do not differ from other ulnaethat we have assigned to this genus, and it is because of theirthickness in relation to their length (Table 1 and 2) that we havereferred these to Laricola robusta.

FAMILY incertae sedisSTERNALARA, gen. nov.

Type Genus—Sternalara.Type Species—Sternalara minuta.Included Species—Sternalara minuta, Sternalara milneed-

wardsi.Differential Diagnosis—Differs from Laricola in: presence of a

second fossa pneumotricipitalis (humerus); differs from Laridaeand Sternidae in: second fossa pneumotricipitalis not as marked,not reaching into the caput humeri.

Etymology—The genus name refers to a closer morphology ofthe humerus to more derived Laromorphae, such as Sterna andLarus.

STERNALARA MINUTA, sp. nov.(Figs. 4–5, Table 2)

Holotype—NMB s.g.12727 (right humerus).Type Locality and Age—Saint-Gerand-le-Puy, Allier, France;

late Oligocene?–early Miocene, MN2.Differential Diagnosis—As for genus; differs from Sternalara

milneedwardsi in: second fossa pneumotricipitalis proportionallywider. Overall much smaller than S. milneedwardsi (around 65%smaller).

Measurements—See Table 2.Referred Specimens—NMB M.A.1582 (left humerus), NMB

s.g.2189 (left humerus), NMB s.g.17520 (right humerus), NMBs.g.5703 (right humerus), NMB M.A.2513 (right humerus). Thefollowing ulnae are only tentatively assigned to this taxon:NMB M.A.1611 (right ulna), NMB M.A.2550 (right ulna), NMBM.A.5161 (right ulna), NMB s.g.16570 (right ulna).

Etymology—The species name minuta refers to the small sizeof the new species.

Remarks—This species is smaller than most individuals ofLaricola elegans, with some bones overlapping with the smallestspecimens. Although the humerus is very close to that of Sternaparadisaea, we refrain at this point from including it in the fam-ily Sternidae, particularly because of the difference in shape ofthe fossa pneumotricipitalis. In any case, a new genus is justi-fied by the amount of time between this record and that of mem-bers of recent Sternidae (putative record dating back to the earlyPliocene, see Olson and Rasmussen, 2001).

Description and Comparisons—The humeri (Fig. 4S–U) aresmaller than those assigned to Laricola elegans. They differ fromLaricola in that a second fossa pneumotricipitalis is present. Thisfossa is, nevertheless, not as deep as that of Laridae and Stern-idae s.s. Also, it does not reach into the caput humeri, and isthus more caudally situated. The sulcus nervi coracobrachialisis distinct. The fossa musculi brachialis is deep, and the well-marked tubercle on the cranial surface of the distal end, be-tween the base of the tuberculum supracondylare dorsale andthe fossa m. brachialis, is evident. Again, we have tentatively as-signed the ulnae to this species based on size. Because Sternalaraminuta is close in size to the smallest specimens of Laricola ele-gans, the smallest ulnae within the range assignable to the lattertaxon may indeed belong to S. minuta. As already mentioned,ulnae within Laromorphae are fairly uniform, and it is thusnot possibly to confidently refer the smallest ulnae (e.g., NMBM.A.2550; 50.4 mm) to S. minuta, but we do acknowledge thispossibility.

STERNALARA MILNEEDWARDSI, sp. nov.(Fig. 4, Table 2)

Holotype—MNHN SG.13742 (right humerus).Type Locality and Age—Saint-Gerand-le-Puy, Allier, France;

late Oligocene?–early Miocene, MN2.Differential Diagnosis—As for genus; differs from Sternalara

minuta in: second fossa pneumotricipitalis not as wide (Fig. 4R).Measurements—See Table 2.Etymology—In honor of palaeontologist Alphonse Milne-

Edwards (1835–1900).Remarks—The humerus belonging to this taxon is markedly

larger than that of Sternalara minuta (Table 2). Still, because ofthe presence of a second fossa pneumotricipitalis, albeit not adeep one, as in extant Laromorphae, we tentatively assign thisspecimen to the same genus.

Description and Comparisons—The second fossa pneu-motricipitalis (Fig. 4R) is not as deep as in extant Laridae orSternidae, also not reaching into the caput humeri. Furthermore,it is not as wide as that of Sternalara minuta. The sulcus nervicoracobrachialis is well defined. As in Stercorariidae, Laricolaand some other Laromorphae, the humerus bears a well-markedtubercle on the cranial surface of the distal end, between thebase of the tuberculum supracondylare dorsale and the fossa m.brachialis (Fig. 4Q). This taxon differs from Sternalara minuta inthat the processus supracondylaris dorsalis does not project ascranially as in the latter species (Fig. 4Q, S), and in that the pro-cessus supracondylare ventrale is distinctly more pronounced inSternalara milneedwardsi.

LAROMORPHAE INDET.

Referred Specimens—NMB s.g.19007 (right carpometacar-pus), NMB s.g.2044 (right carpometacarpus, distal end). Figure4OO–QQ.

Measurements—NMB 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.

Remarks—In size these carpometacarpi match the largestspecimens of Laricola elegans. They differ from Laricola, nev-ertheless, in having a slightly less developed processus inter-metacarpalis and in the presence of a bony bridge at the sulcusinterosseus. Within the Lari, we have found the latter feature tobe present only in Catharacta skua. There are no other featuresdifferentiating it from the carpometacarpus of the Laridae, Stern-idae, and Stercorariidae.

Referred Specimen—MNHN Av.7318 (right humerus), Figure5RR–SS.

Measurements—In mm; greatest length: 72.2; distal width:10.3; minimum width of shaft: 4.3.

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

Remarks—Although close in size to Laricola intermedia, it dif-fers from Laricola in several aspects: the tuberculum supracondy-lare ventrale is situated more laterally in this specimen, makingthe ventral side appear much wider than in Laricola. Laricolaalso has a proportionally longer shaft. The tuberculum ventrale ismuch enlarged in this specimen, and the caput humeri is clearlymore prominent. A second fossa pneumotricipitalis is absent.

PHYLOGENETIC ANALYSIS

Analysis of the character matrix resulted in two most parsi-monious trees (tree length = 61 steps, retention index = 0.67,consistency index = 0.68), which differed only in the positionof Laricola and Anous relative to the clade including Sternidae,Rynchopidae, and Laridae (Fig. 6A–B). The strict consensustree is shown in Figure 6C.

Three characters were optimized as synapomorphies of theclade (Laricolidae + Anous + Sternidae + Rynchopidae + Lari-dae) in both most parsimonious trees: (1) beak, rostrum maxillarevery 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 articu-laris acrocoracoidea that extends over nearly entire width of sca-pus claviculae.

Laricola and Anous fall outside a clade including Sternidae,Rynchopidae, and Laridae. The position of Anous outside thisclade agrees with results obtained in recent molecular phyloge-nies (Baker et al., 2007). The position of Laricola as sister taxonto all other Laromorphae is supported by the presence of a shal-low facies articularis acrocoracoidea in the furcula (21), althoughit would be equally parsimonious to presume Anous in this posi-tion, which is endorsed by the presence of a shallow fossa mus-culi 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 autapomor-phies were recovered for Laricola.

The following characters were optimized as unambiguoussynapomorphies for the clade Sternidae + Rynchopidae +Laridae, but are also present in Limosa: (5) skull, processusmaxillopalatinus 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 distalend, between base of tuberculum supracondylare dorsale andfossa m. brachialis.

A sister-group relationship between Sternidae s.s. and Ryn-chopidae is supported by one ambiguous synapomorphy (36;femur, distal end, crista supracondylaris medialis with distinctcaudally projecting tubercle just proximal of condylus medialis)recovered only with ACCTRAN optimization, and by thefollowing unambiguous synapomorphies: (3) skull, os lacrimalewith maximum craniorostral width at least twice as much asmaximum mediolateral width); (7) skull, processus maxillopalat-inus, concave surface bridged by a pierced sheet of bone; and(34) pelvis, absence of recessus caudalis fossae. Character 3is a unique synapomorphy, whereas characters 7 and 34 showhomoplasy, with the same condition also present in Glareola,and Limosa and Glareola, respectively.

The monophyly of Sternidae is supported by two unambigu-ously optimized characters: (15) mandible, fossa caudalis presentand well-marked, chevron-shaped in ventral view; and (37) femurconsiderably longer than tarsometatarsus.

The following homoplastic characters support Laridae: (1)beak, rostrum maxillare short; occupying less than one third ofthe length of the beak; (2) skull, os frontale, paired fossae glan-dularum nasales meeting at midline of skull; and (6) skull, maxil-lopalatine strut A present.

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DISCUSSION

Phylogenetic Affinities between Laricola and OtherLaromorphae

Laricola trenchantly differs from gulls in several skull features(e.g., length of rostrum maxillare, the shape of the fossae glan-dularum nasales, and processus maxillopalatini), mandible (ab-sence of fenestra caudalis), and in few postcranial features (fur-cula with shallow facies articularis clavicularis, humerus with nosecond fossa pneumotricipitalis, and better marked attachmentsite for m. pronator brevis). Although Laricola is generally re-ferred to gulls (Laridae) (e.g., Mlıkovsky, 2002), this genus hasbeen considered more primitive than modern Laridae by someauthors (e.g., Ballmann, 1976; Olson, 1985), who based this state-ment on the plesiomorphic features of the humerus. As detailedabove, our analysis supports a placement of Laricola outside aclade including Sternidae, Rynchopidae, and Laridae.

Although the aim of our analysis was to place the fossil genusLaricola within a phylogenetic context of recent forms and notto provide an exhaustive assessment of the interrelationships ofrecent Laromorphae based on osteological characters, we notethat our results also constitute the first anatomical evidence fora paraphyletic Sternidae, which has not yet been recovered withanatomical data (e.g., Chu, 1998; Strauch, 1978). The following,presumably plesiomorphic, features set Anous apart from extantgulls, terns, and skimmers: the processus maxillopalatinus (skull)forming a wide and deeply concave trough (the maxillopalatineprocess is less concave in other extant Laromorphae, displayingan entirely different shape, see Fig. 3H–L for comparisons), theabsence of well-developed fenestra caudalis (mandible), the ab-sence of a second fossa pneumotricipitalis (humerus), and thepresence of a very distinct attachment site for m. pronator bre-vis (character 30; humerus).

Although Anous clearly falls outside the clade comprisinggulls, terns, and skimmers, our analysis did not resolve whetherAnous or Laricola are more closely related to this group.Anous shares one plesiomorphic feature with Stercorariidae,namely the shallow fossa m. brachialis, whereas Laricola andStercorariidae both bear a shallow facies articularis clavicularison the furcula. Both morphological and molecular analyses haveyielded conflicting results concerning the position of Rynchops.In his phenetic study of the Lari, Schnell (1970) concludedthat skimmers are overall more similar to terns, something thathad already been speculated upon by Zusi (1962) based on thespecialized feeding habits and corresponding morphologicaladaptations found in both these groups. Livezey and Zusi’s(2007) analysis of modern birds based on anatomical charactersrecovered Rynchopidae as sister group to the Laridae andSternidae, as did the analysis based on nuclear DNA sequencesby Ericson et al. (2003). Baker et al. (2007) recovered a sister-taxon relation between Laridae and Rynchopidae, albeit withlow branch support. Our results are consistent with those ofFain and Houde (2007) who obtained substantial support for asister-taxon relationship between Rynchopidae and Sternidae.We have found this relationship to be overall supported by fouranatomical features (see Results), whereas this grouping wasendorsed by only one character in Mayr (2011).

Diversity of Early Laromorphae

The fossil record indicates that the early Miocene diversityof Laromorphae is far higher than assumed previously. At leastfive 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 insize, whereas L. robusta can be distinguished by its relativelyshorter and stouter shaft of the humerus, as well as by better-developed cristae bicipitalis and deltopectoralis. The sheer num-

ber of species and the combination of features present in all thesespecies warrant the construction of a new family, Laricolidae.Sternalara has a more derived morphology of the humerus, witha second fossa pneumotricipitalis still differing from that of ex-tant Laromorphae in being shallower and not reaching into thecaput humeri, implying that it is unlikely for this taxon to havebelonged in any recent families of Laromorphae. Whether Ster-nalara was more closely related to Laricola or to more-derivedLaromorphae is difficult to say based on the material available,namely humeri, which is also why we considered this taxon un-suitable for inclusion in the phylogenetic analysis.

Not much material from the original description of Laricolatotanoides by Milne-Edwards can still be retained in this species,despite him mentioning several skeletal elements, many of whichare labeled L. totanoides in the MNHN collection. We considerthe depicted humerus and the ulnae with the dimensions speci-fied by him as true L. totanoides because they are explicitly largerthan the range we determined for Laricola elegans. All otherbones fall within the large size spectrum of L. elegans, whereasthe humeri and some ulnae clearly belong to a different speciesaltogether, Laricola robusta.

Concerning Laricola elegans, we acknowledge that the widesize range indicated for this bird may be too broad for asingle species, and indeed this range of variation may encom-pass several species, which is primarily evinced in the variationsfound within the skull and the configuration of the hypotarsus. Inliving Laromorphae, size differences of this magnitude—around20%—are undocumented (see del Hoyo, 1996), but direct com-parisons with the fossil record have not yet been performed. Asexplained in Geology (see above), the fact that the fossil collec-tion from Saint-Gerand-le-Puy potentially spans a period rangingfrom the late Oligocene to the early Miocene may also explainthe large size variability found within the material attributed toLaricola elegans.

Incorporating Laricola into a phylogenetic analysis of extanttaxa has demonstrated that this extinct taxon is an early off-shoot of Laromorphae whose divergence predates the radia-tion of modern gulls, terns, and skimmers. The fossil record ofcrown group Laromorphae is fairly recent. Putative terns andgulls from Miocene deposits (e.g., Miller and Sibley, 1941; Grig-orescu and Kessler, 1977; Mlıkovsky, 1996) are based on frag-mentary records, and cannot be assigned to any modern genus.Pliocene fossil evidence for Sternidae is minor, including car-pometacarpi of two taxa from, most likely, the early PlioceneNorth America (Olson and Rasmussen, 2001). Pliocene fossils ofLaridae appear to be more recurrent, particularly in North Amer-ica: several specimens were described from the early Plioceneof North Carolina (Olson and Rasmussen, 2001); Emslie (1995)described two species (Larus perpetuus and L. lacus) from theLate Pliocene of Florida; and Larus elmorei is known from theearly–middle Pliocene of Florida (Brodkorb, 1967). None ofthese fossils seems to significantly depart from the morphologyof extant gulls. The Quaternary fossil record of Laromorphae isfar more abundant and diverse (e.g., Brodkorb, 1967, Mlıkovsky,2002).

Fossils from Saint-Gerand-le-Puy are difficult to date with pre-cision. The quarry in Montaigu-le-Blin and its fauna yielded fos-sils from biozone MN2 (from 22.5 to 20.5 Ma), but the uncertaintyas to whether the old collections all come from this very placecasts doubt on the precise dating of the fossil remains. Indeed,fossils were collected in the 19th and early 20th centuries from thewhole area in various localities surrounding Saint-Gerand-le-Puyand no exact provenience was recorded. It is therefore possiblethat the different species of laromorphs present in the lake systemduring this time did not co-occur. This lake system was most cer-tainly used as breeding grounds by several taxa of birds, includ-ing Laricola, as reflected by the vast amount of juvenile individ-uals recovered. Whether these charadriiform birds were seasonal

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visitors or permanent residents of the area can, however, only bespeculated upon.

ACKNOWLEDGMENTS

We thank C. Meyer (NMB) for facilitating the study of thefossil specimens. We are grateful to R. Allain (MNHN) for hisassistance 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. We-ick (NMB) assisted with the preparation of fossil specimens. Wethank the Willi Hennig Society for subsidizing the free edition ofTNT. A travel grant from Stiftung zur Forderung des Naturhis-torischen Museums (Basel, Switzerland) to V.L.D. supported thisresearch. Helpful comments by D. Ksepka (University of NorthCarolina, Chapel Hill, U.S.A.), C. Mourer-Chauvire (UniversiteClaude Bernard, Lyon, France), and T. Worthy (University ofNew South Wales, Sydney, Australia) improved the manuscript.

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Submitted March 9, 2011; accepted May 5, 2011.Handling editor: Trevor Worthy.

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A revision of the Lari (Aves, Charadriiformes) from the early Miocene of Saint-Gérand-le-Puy...

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