A late Pleistocene deposit of 60 species of vertebrates and 12 of invertebrates is described from Robinson Cave, Overton County, Tennessee, U.S.A. Forty-eight species of mammals are represented by at least 2,483 individuals; 10 % are extinct, 10 % occur in the state only as boreal relicts in the Great Smoky Mountains; 23 % no longer occur as far south as Tennessee; 57 % occur at or near the site today. Nínety-one percent of the Recent mammal species can be found living today in the Minnesota-Wisconsin area, approximately 10 degrees farther north. Fluorine analysis suggests a long period of accumulation. The following 10 mammalian species are recorded from Tennessee for the first time. Sorex arcticus, Microsorex hoyi, Citellus tridecemlineatus, Clethrionomys gapperi, Microtus pennsylvanicus, Synaptomys cooperi, Synaptomys borealis, Zapus nudsonius, Napaeozapus insignis, Martes americana. Six additional species are present as boreal relicts in the Great Smoky Mountains of eastern Tennessee but not at the site today : Sorex cinereus, Sorex dispar, Sorex palustris, Parascalops breweri, Glaucomys sabrinus, Mustela nivalis. Six forms are extinct: Canis dirus, Ursus americanus amplidens, Sangamona furtiva, Dasypus bellus, Mammut americanus,Megalonyx jeffersoni. Twenty-six additional species of mammals, all of the snails, birds, reptiles, and amphibians recovered from the fauna still inhabit the area today: The fauna is indicative of a cold-temperate climatic episode associated with the Wisconsin glaciation, but may be chronologically mixed. 

The presence at Silveirinha of one of the earliest, ? Late Paleocene or Lowermost Eocene, european representatives of the genus Diplocynodon is based mostly on isolated bones and teeth (often from juveniles). This small-sized form is the only crocodilian so far recognized in this site. The longevity of Diplocynodon in Portugal becomes much extended; the genus survived there until the Middle Miocene at least. A discussion on the possible affinities with other eocene Díplocynodon and especially those from Cubillos-Valdegallina (Zamora, Spain) is presented. On the other hand, differences have been detected in comparison with: Díplocynodon tormis, from the middle Eocene of the Douro basin in Spain, which may belong to another phyletic line; and the aff. Diplocynodon from Dormaal (Belgium) and Le Quesnoy (France), nearly contemporaneous of Silveirinha. The Silveirinha Diplocynodon and many other data strongly suggest moist, subtropical, quite limited in space environments related to an alluvial plain crossed by small, meandering channels. 

Until now Aplodontidae of the European Oligocene have been documented by four species only. The phylogenetic relations remained obscure. as the distribution of only one species has been known in some detail. New material made it possible to define the stratigraphic range of two of the already existing species (Plesispermophilus angustidens, Sciurodon cadurcense) and to follow their development during the Oligocene beginning with the event of the « Grande Coupure ››. Sciurodon remained nearly without change until the end of the Middle Oligocene. Plesispermophilus angustidens split into two distinct phyletic lines, one of which (P. macrodon n. sp.) reaching considerable size, is represented till the beginning of the Upper Oligocene (Pech de Fraysse, Gaimersheim). The other line leads to Plesispermophilus ernii (basal Upper Oligocene of Burgmagerbein 1. terminal Upper Oligocene of Coderet). Besides the already known forms a new small-sized species (P. atavus n. sp.) is described, which by its primitive features closely resembles the genus Plesispermophilus. Two other small-sized species already known from the Upper Oligocene (? P. argoviensis) and Lower Miocene (? P. descedens) seem to be closely related to the new species. It cannot be decided whether they are descendents of this line or have developed independently, because of their poor fossil record.
Comparison of the evolutionary modalities in the different phylogenetic lines reveals general trends. the most striking of which is the complication of the tooth pattern by the development of additional crests. In the lower molars the cusps diminuate in size and are more and more transformed into ridges. ln addition new connection between the crests appear. in the upper molars, the « selenodont » shape of the teeth becomes more and more dominant, and in the two main evolutionary lines of Plesispermophilus the metaconulus becomes duplicated. A further evolutionary trend is the size increase of the premolars compared to the molars, which is even more pronounced in the Miocene Aplodontidae.
Phylogenetic relations between the primitive Plesispermophilus and certain « prociurines ›› of Northern America as well as between Plesispermophilus (P. angustidens) and more progressive forms of the Upper Oligocene (P. ernii, P. macrodon n. sp.) can be documented. In this light, the taxonomic distinction between Prosciurinae (bunodont) and Allomyinae (selenodont) sensu Rensberger 1976 can be shown to be artificial, because it separates forms from each other, which are evidently closely related. Consequently the separation into two subfamilies has been abolished. 

The comparative morphology and biometry of the lower cheek teeth of modern Equus are studied on approximately 300 mandibles belonging to the 10 usually recognised species : Equus grevyi, E. burchelli, E. quagga, E. zebra, E. africanus, E. asinus, E. hemionus, E. klang, E. przewalskii, E. caballus. The studied parameters comprise : occlusal length and width, postflexid length and index ; shape of the double knot (metaconid + metastylid + lingual groove) ; depth of the vestibular groove on the molars ; frequency of the pli caballinid, protostylid and other enamel plications or islets ; frequency of the dP/l.
The same methods of study are applied to a number of North American, Eurasian and African species. For the sake of comparison, some Hemphillian equids were observed (Dinohippus interpolatus, Dinohippus leidyanus, Astrohippus ansae, Phiohippus mexicanus) but most of the discussed material belongs to Pliocene or Pleistocene species of Equus : the « stenonine ›› E. stenonis, E. simplicidens, E. sanmeniensis and E. teilhardi; the « caballine ›› E. scotti, E. lambei, E. Iaurentius, E. mosbachensis, E. germanicus, E. gallicus, E. taubachensis and the Liakhov horse. The relationships of other species, in particular the North American E. calobatus, E. occidentalis, E. cf mexicanus are not clear for the moment. ln Africa, the Plio-Pleistocene species from Koobi Fora (Kenya) show some stenonine and perhaps asinine affinities. The relationships of E. numidicus and E. tabeti are uncertain but these species are probably related to the East African ones. E. mauritanicus is most certainly related to the Quagga group.
The biometrical data are gathered in 32 tables ; 4 photographie plates and 19 figures illustrate the next. The whole is a complement of the previously published studies of the skulls, upper cheek teeth, incisors and metapodials of modern and fossil Equus.

  

Les Pistes de Vertébrés du Stormberg Supérieur ("Trias terminal à Rhétien"), ou Quthingien

Si les zones du Stormberg inférieur se sont révélées contenir de nombreuses traces, surtout dans les faciès dits "Molteno moyen et supérieur", représentant apparemment la base du Keuper, il est frappant de voir pratiquement l'ensemble de cette grosse faune "Molteno" disparaître avec la fin de cette période, que nous avons appelée le "Maphutsengien".

Dès les premières zones du Stormberg supérieur, que nous nommons le"Quthingien" la zoocénose et la phytocénose, en même temps que les données d'ensemble manifestées par l'environnement, sont modifiées. Nous ne verrons plus guère de dépôts marécageux à flore riche et variée, parfois même luxuriante. Les fougères elles-mêmes ont disparu. Elles sont remplacées par de maigres plantes, aux feuilles très souvent filiformes qui paraissent témoigner d'un climat continental. Le sol est devenu de plus en plus rouge, avec des variations latérales beaucoup plus accusées. Les fleuves amenant des galets des monts du "Grand Sud" ont tari. La faune va en subir les conséquences. Certaines des espèces se révèleront sautillantes ou coureuses, pour un grand nombre plus légères et pour la quasi-totalité d'apparence carnivore ou entomophages, les phytophages devant se contenter d'un régime ingrat,difficile ou à tout le moins irrégulier,les dépôts le montrent.

C'est dans ces conditions que s'inaugure notre Etage nouveau,quelque peu discordant sur les zones A/5, A/6 ou A/7 du Stormberg inférieur (Maphutsengien). Le Stormberg supérieur (ou Quthingien) commence avec le paléopaysage remarquable dit de Moyeni, que nous allons maintenant étudier, typologiquement, avec ses homologues du même âge. Quelques 38 types d'animaux tous nouveaux vont défiler à nos yeux lors de la zone de base de cet Etage, ou zone B/1.

L'on nous avait proposé d'intituler ce Ile Tome de la série : "La grande Dalle de Moyeni et ses homologues. Paléo-spectacles, scènes et paysages animaux au Lesotho à l'approche du Trias finissant". Nous avons préféré garder le sous-titre plus haut, peut-être plus prosaïque.

Un llle Tome est en préparation : "Les développements ultérieurs et terminaux de la faune du Gondwana".

  

The locality of Montredon has provided 8 species of camivores, often little documented, that are discribed and depicted. All the remains are fragmentary and generally badly preserved which lowers very much the possible precision of the taxonomic study; this one has however allowed the creation of a new subspecies. The most richlypdocumented forms are an ursid (Indarctos) and a felid (Machairodus). Issuing often from a westem Europe evolution, the carnivores are well inserted between those of the better known faunas of the "Classical Pontian" and of the lower Vallesian and this situation fits also well with their stratigraphic level, that cannot however be more accurately defined with them alone.
  

A recent campaign of excavations (1965-68) undertaken by the Laboratoire de Paléontologie of Montpellier in pockets of the Quercy phosphorites, has permitted the dating of several localities thanks to the analysis of their micromammalian fauna.
The rodents of localities belonging to the middle Oligocene (La Sauvetat zone) are the object of this study. This rodent fauna has been compared to the forms coming from three stratified localities belonging to the biochronologic zone of La Sauvetat :  Balm, Montalban and Lovagny.
The abundance of material, notably of theridomyids and cricetids, permitted a summary statistical study to be effected and gave some precise details on the variability of the species (Theridomys varians in particular). Other details were provided on certain groups, notably on the upper teeth of Scíuromys cayluxi and on the milk teeth of theridomyids. Some specimens of «Sciurus» sp. and of Plesispermophilus angustidens were collected. By this means, it has been possible to establish, for the first time, a precise upper limit to the epoch of their appearance in Europe. 

The Laprade fauna is chronologically situated between those from Egerkingen and Lissieu and consequently, is close to the MP 14 reference-level of the European mammalian biochronological scale (Symposium of Mainz, 1987).
This new fauna is presently the oldest known in the Quercy phosphorites, formerly the Le Bretou fauna (MP 16) was considered as the oldest one. The Laprade fauna includes 21 species which belong in 7 mammalian orders (Marsupialia: Amphiperatherium bastbergense, Amphiperatherium goethei; Apatotheria: Heterohyus (Gervaisyus) pygmaeus nov. subgen., nov. sp.; Lipotyphla: Saturninia cf. mamertensis, Saturninia cf. intermedia; Chiroptera: Vespertiliavus lapradei nov. sp.; Rodentia: Protadelomys cf. lugdunensis, Elfomys nov. sp.; Primates: Nannopithex cf. filholi, cf. Pseudoloris or Pivetonia; Perissodactyla: ?Palaeotherium ?castrense, small-sized Palaeotherium sp., Plagiolophus sp., Anchilophus sp.; Artiodactyla: Dichobune cf. robertiana, Mouillacitherium cartieri, Tapirulus cf. depereti, Mixtotherium priscum, Pseudamphimeryx schlosseri, and Artiodactyla indet). Sixteen of these species are mentioned for the first time from the Quercy faunas.
The recognition of a new apatemyid, Heterohyus (Gervaisyus) pygmaeus nov. subgen., nov. sp., attests to an early origin of a lineage known in the Late Eocene. The emballonurid bat Vespertiliavus lapradei nov. sp. is presently the earliest record of this genus and family. This Auversian fauna leads to discuss the age of taxa showing archaic features. These taxa were defined on specimens collected in the Quercy during the last century and have never been found in the Quercy localities recently investigated. This fauna bears also evidence of a karstic filling episode older than those previously dated by fossils in the Quercy Jurassic. 

The study of the bovids from Al Jadidah (Hofuf Formation, Saudi Arabia) confirms that the fauna comes from a pre-Hipparion level. The Al Jadidah age is close to that of Fort Ternan (14 m.y.) and Beni Mellal, but cannot be older than that of Fort Ternan. The age of the Hofuf Formation is close to but slightly older than the oldest deposits of the Ngorora Formation (Kenya). 7 to 9 species have been recorded, of which 2 to 4 remain indeterminate. If the great specific diversity of te bovids from this locality gives evidence of immigrations from anterior Asia (Turkey) (e.g. Pachytragus Iigabuei sp. nov.), the bovid assemblage of Al Jadidah results in fact from a double influence: from the anterior Asia and mainly from Africa (e.g. the Caprotragoides lineage and the Neotragini? Homoiodorcas). The Al Jadidah bovids reflect, on the whole, the predominant character of open to very open environment, which supports the conclusions drawn from our two preliminary studies. 

We present here a new updated counting of the perissodactyls of Robiac, the type locality of the MP 16 level of the biochronological scale of paleogene mammals and that of the Robiacian stage of Eocene Land Mammals Ages in Western Europe.
The outcrop of Robiac consists actually of two 500m apart loci, Robiac-Nord and Robiac-Sud, considered of the same age according to the current discriminating power, and is dated from -38,7 MA after the last faunal, magnetostratigraphic and climatic calibrations.
It has yielded a very abundant and rich of 21 taxa perissodactyl fauna, topped by the giant Lophiodon lautricense, last representative of the family Lophiodontidae, of which it is the last proved deposit. The Palaeotheriidae are much diversified with 5 genera and 9 species of "Pachynolophinae", 3 genera and 10 species of Palaeotheriinae. Nine taxa have been defined from Robiac: Chasmotherium depereti n. sp., Palaeotherium castrense robiacense Franzen, 1968, the genus Leptolophus Remy, 1965 with the species L. stehlini Remy, 1965 and L. magnus Remy, 1998, Anchilophus (Paranchilophus) jeanteti Remy, 2012, Metanchilophus chaubeti Remy, 2012, Lophiotherium robiacense Depéret, 1917 and Pachynolophus gaytei n. sp.
The faunal Robiac cenogram with the associated flora testify to a hot, wet and forestal environment, likely corresponding to a short warming climatic phase; the broken up fossil bones should have been carried away and then gathered in swamp areas along the banks of a meandering river.
The swarm of mammals of Robiac, the richest of contemporaneous deposits, has been followed by a drastic drop in perissodactyl diversity at the MP 17A level; a crisis which could have originated in a renewal of the global Eocene cooling. Fons 4, the type-locality of this level, is largely scarcer in perissodactyls and its cenogram testifies to a less diversified fauna, with on the whole smaller species, that likely means a cooler and drier climatic environment; a new perissodactyl diversification occurred but later.
  

The Late Miocene site of Gritsev (MN 9, Ukraine) has yielded a very rich bat fauna, the remains of which are well preserved. Compared to other Neogene bat assemblages of Europe, the Gritsev bat community is unique in preserving plecotine bats, which are rare from Neogene sites. Some peculiar and new bat species, including a large plecotin Otonycteris, already were described from the Gritsev mammal site. Here we report new records of small plecotin bats from Gritsev, including a new taxon, Barbastella maxima nov. sp. This is the earliest reliable fossil record of this genus and it differs from more recent species of Barbastella in being considerably larger. The evolutionary patterns in the odontology within the tribe Plecotini, supported by biostratigraphical distribution of fossil records of Plecotus are discussed. The morphological peculiarities of the new fossils of plecotine bats from Gritsev are discussed in connection with its possible taxonomical affinity. 

Confident attribution of bipedal tridactyl dinosaur tracks to theropods or ornithopods can be challenging. Here we describe trackways produced by tetanuran dinosaurs, previously attributed to hadrosaurs, from Coahuila State, northeastern Mexico. Multiple trackways headed in the same direction suggest gregarious behaviour in these late Campanian theropods.  

The late Pliocene site (MN 16) of Balaruc II (Hérault, France) has provided remains of the following amphibians and reptiles: Chelotriton pliocenicus nov. sp. and Triturus marmoratus (Salamandridae), cf. Rana (Ranidae), cf. Blanus (Amphisbaenidae), cf. Agama (Agamidae),  Gekkonidae indet., Lacerta s.l. (Lacertidae), "Ophisaurus" sp. (Anguidae), Michauxophis occitanus (Aniliidae), Erycinae indet. (Boidae), Elaphe cf. E. longissima and Malpolon sp. (Colubridae), cf. Naja (Elapidae) and Vipera sp. (Viperidae). The salamandrid Chelotriton pliocenicus and the aniliid Michauxophis occitanus constitute, up to now, the only records of these groups in the European Pliocene. The fauna is indicative of a warm, dry
subtropical climate. 

A new Devonian vertebrste locality has been discovered in 1976 in the basal part of the Khush-Yeilagh Formation in the eastern Alborz Mountains of Iran. The fossils occur in a band one centimeter thick which is identifiable at other outcrops in the area. A preliminary study of the remains has yielded the following faunal list : Placodermata (Phlyctaeniida indet.. Groenlandaspididae indet.. Coccosteidae indet., Holonema sp., Ptyctodontida indet., Antiarcha indet., Bothriolepis cf. kwangtungensis), Elasmobranchii (Ctenacanthidae indet.), Acanthodii (Gyracanthus sp., « Onchus » overathensis, Ischnacanthiforme indet.), Dipnoi (? Dipteridae indet.), « Crossopterygii ›› (Onychodus cf. sigmoides, Holoptychiidae indet., Osteolepididae indet.). This fauna is older than the Middle Givetian and probably of Middle or Lower Eifelian age. Owing to its diversity, it may prove possible to use it in the future as a reference fauna for the study of the Devonian vertebrates in Central Asia and the Middle East. 

The Las Águilas site near Porvenir de Jalpa, Coahuila, Mexico, is extremely rich in tetrapod remains comprising both bones and trackways of several dinosaur taxa of late Campanian age. Within a 50 m thick section we identified at least nine layers with dinosaur bone assemblages. In one of these the dinosaur bones are associated with remnants of eusuchian crocodilians, turtles, plesiosaurs, pterosaurs, tyrannosaurids, dromaeosaurids, parksosaurid, hadrosaurids, ceratopsids, and ankylosaurs. This layer is also rich in coprolites of turtles, crocodilians and likely theropods, thus providing evidence for the wealth of Late Cretaceous vertebrate life in the area. 

 A new genus of hystricognath rodent with two new species, Soriamys gaimanensis and Soriamys ganganensis, from the Colhuehuapian Age (Early Miocene), Sarmiento Formation, of Patagonia is described. The first species comes from the south barranca of the valley of Rio Chubut, near Gaiman locality (Chubut Province). It is known through a great number of dental remains in different stages of wear. The teeth are protohypsodont, with cement in the principal valleys. P4 are simpler, more oval inoutline and without a differentiated hypocone-hypoflexus. Upper molars are pentalophodont in early stages of wear and become bilobated in middle and advanced stages; M3 have a temporary third posterior lobe due to the maintenance of the braquiflexus on the posterior wall of the tooth. Lower teeth have a completely asymrnetrical pattern in relation to the uppers. The molars are trilophodont, with only one complex crest in the trigonid composed of the fusion of the anterolophid and mesolophid. The dp4 have a derived pentalophodont pattern due to the loss or fusion of the mesolophid and the development of an accessory transverse crest between the anterolophid and metalophid. The second species is known through a complete skull and jaw and other skull fragments with the dentition, proceeding from Pampa de Gan Gan, Chubut province. Respect to the first species, the teeth are higher crowned, with an earlier occlusal simplification and a third posterior lobe of M3 more developed and persistent. The skull and jaws show many chinchilloid characters, such as jugal bone with an ascending process, concave palate, very prominent dorsal shelf of the masseteric fossa, nasolachrymal duct opened laterally on the maxillary and very developed lateral mandibular fossa. Moreover, Soriamys is closely related to Cephalomys by sharing a similar asymmetric dental pattern and other dental and mandibular characters. These two genera constitute a natural group, the Cephalomyidae, with peculiar dental characters, like the asymmetric pattem of the upper and lower teeth; Cephalomyopsis, and probably Litadontomys, can be considered part of this group. Scotamys and Perimys constitute a distinct lineage (the Neopiblemidae or Perimyidae), more closely related to the chinchillids. Many characters shared by cephalomyids, eocardiids and caviids suggest a closer affinity between the Chinchilloidea and the Cavioiidea as a whole. Abrocoma shows also an asymmetric dental pattern and other chinchilloid features that suggest some degree of relationship with the cephalomyids, still not well determined. 

This study describes four taxa of Gliridae from the Oligocene mammal locality Gröben 3: Gliravus tenuis BAI-ILO, 1975, Bransatoglis micio (MISONNE, 1957), B. planus (BAHLO, 1975) and B. heissigi n. sp. Gliravus tenuis from Gröben 3 is somewhat more advanced than the type population found in Heimersheim. This confirms previous research suggesting that Gröben 3 should be dated earlier than Heimersheim (MP 24). The first documented occurrence of B. mício around level MP 24 was found in Gröben 3. An abundance of tooth material from B. planus in Gröben 3 makes it possible, for the first time, to observe evolutionary stages within this species from MP 21 until MP 28. B. heissigi n. sp. is restricted to level MP 24. This species is located between B. mísonnei (MP 20 - 23) and Microdyromys praemurinus (MP 25 - 28). Within the lineage Bransatoglis bahloi - B. misonnei - B. heissigi, a decrease in size is noticeable.

  

The Montian locality of Hainin (Hainaut, Belgium) yielded about twenty teeth of Multituberculates. They are very peculiar forms, showing no affinities, at the generic level, with those hitherto known from North America, Asia and Europe. They are referred to the new taxa Boffius splendidus nov. gen., nov. sp., Hainina belgica nov. gen., nov. sp., and H. godfriauxi nov. gen., nov. sp. They expose some common features, such as the advanced type of first upper molar. possessing at least three complete rows of cusps. Because of this, and also of the upper premolar reduction, Boffius splendidus appears as the most specialized form within the Ptilodontoidea suborder.
Several other characters of Hainina seem to be less advanced, such as the great number of upper premolars and the simple cusp-formula of the first lower molar.
Till now, only H. godfriauxi has been recovered within the Thanetian fauna from Cemay-les-Reims, where it is very poorly documented. 

The Oligocene deposits of the Ulantatal area in Inner Mongolia (China) contain among the richest mammalian faunas from Asia. To date, only some parts of the rodent faunas have been described. Here, we propose to review the rodent faunal lists for each site, including the description of a few new rodent specimens. We describe three additional rodent species: the Cylindrodontidae Anomoemys lohiculus, the Eomyidae Asianeomys sp., and the Dipodidae Litodonomys huangheensis. This study allows us to constrain the stratigraphic range of Anomoemys lohiculus, which ranged from the late Early Oligocene to the early Late Oligocene in this area. Asianeomys sp. and Litodonomys huangheensis are dated from the latest Oligocene. These Oligocene deposits consist now of more than 70 species of mammals if we include the fauna from Kekeamu. This latter corresponds to the basal part of the Ulantatal Formation and could be dated biochronologically from the earliest Oligocene. When compared to the faunas from the Valley of Lakes in Central Mongolia, the Ulantatal faunas present a great majority of rodents, and this difference can be partly explained by sampling and description biases regarding macro-mammals. This study also shows that variations existed between Inner and Central Mongolia, especially regarding the composition of the rodent paleocommunities. However, the assessment of their evolutionary history in this part of Asia with respect to the important climate and environment changes, require further precisions and more material than current data allow. 

Revision of the hipparion material from Montredon, including newly excavated and other unpublished specimens brings evidence of specific heterogeneity.
A fragmentary very small MC III seems very close to H. macedonicum from the upper Vallesian of Ravin de la Pluie, Greece. In that same site was also found a large hipparion.
Most of the Montredon material is referred to H. depereti. This species associates characters usually found in Vallesian hipparions (highly plicated upper cheek teeth, deep vestíbular grooves on the lower cheek teeth, robust metapodials) with characters more frequent in Turolian forms (middle size, lack of confluence in the upper premolar fossettes, lack of ectostylids on the adult lower cheek teeth, well developped keel on the MC III, facette for the 2nd cuneiform present on all MT III). H. depereti shares some of these characters with the Spanish and Portuguese hipparions transítional between the Vallesian and the Turolian (Masia del Barbo, Azambujeira) but is not identícal to any of them. The upper Vallesian hipparion material from Diavata, Greece, probably belongs to H. depereti but not the large hipparion rests from Ravin de la Pluie.
Thus, Montredon and Ravin de la Pluie may well share the same small species, H. macedonicum, but they differ in the associated one: middle-sized H. depereti at Montredon, large-sized H. primigenium at Ravin de la Pluie. Both sites, however, give evidence of small hipparions during the Vallesian, coexisting with other larger species.
The present paper also proposes an adaptation of the Kiesewalter's indices that calculates the height at the withers using the metapodial length, and discusses indices proposed by Gromova and by Sen et al. to express the relative development of the metapodial keel.