A new Miocene local fauna from the Sierra Madre Oriental at San Luis Potosí, Central-East Mexico,and its paleontologic significance

Mexico's Late Neogene mammal faunas are largely known from localities in the Trans-Mexican Volcanic Belt; those from other morphotectonic provinces are few and far apart. Thus, the discovery of Late Miocene vertebrates in western Sierra Madre Oriental at San Luis Potosí, the Paso del Águila local fauna, significantly adds to this meager record. The assemblage was collected from the floodplain facies of the San Nicolás Formation, a ∼1100-m thick, dominantly fluviolacustrine and calcilithitic, 15°–20° NE dipping sequence preserved in the Peotillos-Tolentino Graben, between 22°11’–22°19’ N and 100°30’–100°39° W. It includes remains of cf. Trachemys, a small to medium-sized emydid chelonian, a large camelid, a small cervid and a new species of the equini Pliohippus s.s., comparable in size, cranial morphology and odontographic characters to the Clarendonian-Early Hemphillian horses of the Pliohippus clade. Ar-Ar dates from ash-fall tuffs seemingly above and below the fossiliferous strata, bracket the age between 12.33 and 7.41 Ma (i.e., late Middle to Late Miocene), that is, within the Late Clarendonian-Early Hemphillian NALMA interval, making this fauna the first in Mexico from this age. The Paso del Águila local fauna is at least partly correlative with the Hemphillian local faunas from the TMVB and adjacent areas (e.g., Rancho El Ocote, Guanajuato and Tecolotlán, Jalisco), the Central Plateau (e.g., Arroyo Los Fragmentos, Zacatecas), and the Sierra Madre Occidental (e.g., Yepómera). Elsewhere, it is broadly correlative with the Late Clarendonian-Early Hemphillian faunas from the California Coast Ranges (e.g., North Tejon Hills, Ricardo and Dove Springs in the Mohave Desert), and the Gulf Coast Plain, Florida (McGehee Farm and Mixon). The Paso del Águila local fauna was part of a subtropical savannah and pine-oak forest (with a well-developed understory) biome that thrived on a climate regime much more humid than today.     

Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications

Crocodylian remains are collected in 39 fossil-bearing localities but only in seven localities specimens with reliable taxonomic attributions, at least to genus level have been collected. Three species have been reported from the early Lutetian Purga di Bolca site: Pristichampsus cf. Pristichampsus rollinati, Asiatosuchus sp., Hassiacosuchus sp. (=Allognathosuchus sp.). The three crocodilians discovered at Purga di Bolca have been reported also from Geiseltal and Messel (Middle Eocene, Germany). Bolca at that time was part of a Tethysian archipelago and no mammals have been found there till now. Crocodilians and turtles clearly arrived from the European mainland across a marine water barrier. Among the other fossiliferous localities of Veneto, very interesting is the Monte Zuello site, of late Middle Eocene age, yielding a longirostrine crocodilian, Megadontosuchus arduini, a tomistomine species. Tomistomines are known in contemporaneous sediments of both Europe and Africa, but the European forms Dollosuchus and Kentisuchus seem the closest taxa. Remains of Oligocene age have been collected in Veneto and Liguria, but the fossils discovered in the second region are teeth or fragmented bones. The fossil crocodilians of Monteviale (Veneto), of Early Oligocene age, have been assigned to two species but they have been recently all identified as Diplocynodon ratelii, known from several European sites of Late Eocene, Oligocene and Miocene age. This species arrived in the Monteviale area from the European mainland across a narrow sea. Several crocodilian fossils of Miocene age are very fragmentary or represented by isolated teeth. In the Middle and Late Miocene of Sardinia, a well-established species, Tomistoma calaritanum is present. Remains of Tomistoma of the same age have been reported in some localities in Tuscany, Apulia, Sicily and Malta. In the Mediterranean area, the genus is known from European and African sites (of older age). The colonisation of Europe by this genus is the result of a dispersion from Africa (or less probably from Asia). During Late Miocene Sardinia and Tuscany belong to the same palaeobioprovince characterized by the Oreopithecus-Maremmia fauna. In Tuscany, a crocodilian identified as Crocodylus bambolii is present in the late Miocene site of Monte Bamboli. If the generic attribution of this form is correct, its ancestors must have arrived from Africa. Another fossil assemblage of Late Miocene age characterizes the Apulia-Abruzzi palaeobioprovince (Hoplitomeryx-Microtia fauna) and testifies complete isolation between the two palaeobioprovinces. In this last area, remains of Crocodylus sp. have been collected in coastal sandstones at Scontrone (Abruzzi) and in several fissure fillings of Gargano of slightly younger age. The ancestors of this species arrived from Africa while no African elements are present among the mammalian fauna. The dispersion of the genus Crocodylus in the Italian palaeoislands may have taken place once, with allopatric differentiation of the two populations (Tuscany-Sardinia and Apulia-Abruzzi) or twice with independent colonisation of each area.     

Şerefköy-2, a new Late Miocene mammal locality from the Yatağan Formation,Muğla,SW Turkey

Here we report on a new fossil locality, ?erefköy-2, from the Yata?an Basin of southwestern Turkey that preserves a well-sampled, abundant, and diverse mammal fauna. Indeed, after three field seasons, more than 1200 catalogued specimens representing 26 mammal species belonging to 14 genera make the ?erefköy-2 mammalian assemblage one of the richest Late Miocene fauna from Anatolia. Five hipparionines, six bovids, including the rare and enigmatic Urmiatherium rugosifrons and the presence of Pliohyrax graecus, strongly support affinities with Late Miocene faunas from Samos Island, Greece. Through a consideration of the identified material and the subsequent comparison with material from well-known Balkan and Anatolian faunas, a Middle Turolian (MN12) age for ?erefköy-2 is indicated.     

Creodonts and carnivores from the Middle Miocene Muruyur Formation at Kipsaraman and Cheparawa,Baringo District,Kenya

The Middle Miocene Muruyur Formation (ca 14.5 Ma), Tugen Hills, Kenya, has yielded a huge creodont and a variety of carnivores ranging in size from mongoose-sized viverrids and herpestids to lion-sized amphicyonids. The fauna partly fills what used to be a major gap in our knowledge of Neogene African carnivores, spanning the period between the better known Early Miocene assemblages of western Kenya and eastern Uganda, and the Late Miocene and Plio–Pleistocene faunas of East Africa. Present in the deposits are Megistotherium, two species of Hecubides, one species of Agnotherium, Herpestes, Vishnuictis, and one or two undetermined felids.     

New observations on the Late Miocene–Early Pliocene Lutrinae (Mustelidae: Carnivora,Mammalia) from the Middle Awash,Afar Rift,Ethiopia

New observations on the Late Miocene and Earliest Pliocene mustelids from the Middle Awash of Ethiopia are presented. The Middle Awash study area samples the last six million years of African vertebrate evolutionary history. Its Latest Miocene (Asa Koma Member of the Adu-Asa Formation, 5.54–5.77 Ma) and Earliest Pliocene (Kuseralee and Gawto Members of the Sagantole Formation, 5.2 and 4.85 Ma, respectively) deposits sample a number of large and small carnivore taxa among which mustelids are numerically abundant. Among the known Late Miocene and Early Pliocene mustelid genera, the Middle Awash Late Miocene documents the earliest Mellivora in eastern Africa and its likely first appearance in Africa, a new species of Plesiogulo, and a species of Vishnuonyx. The latter possibly represents the last appearance of this genus in Africa. Torolutra ougandensis is known from both the Late Miocene and Early Pliocene deposits of the Middle Awash. The genus Sivaonyx is represented by at least two species: S. ekecaman and S. aff. S. soriae. Most of the lutrine genera documented in the Middle Awash Late Miocene/Early Pliocene are also documented in contemporaneous sites of eastern Africa. The new observations presented here show that mustelids were more diverse in the Middle Awash Late Miocene and Early Pliocene than previously documented.     

Albertona balkanica from Early Miocene of Snegotin (Serbia) – implications for the distribution and evolution of Miocene Ochotonidae

Remains of Albertona balkanica from the Early Miocene clays of eastern Serbia have shifted the range boundary of this species in direction of Central Europe. The peculiarities in tooth morphology were used for comparison and defining of the possible evolutionary connection with other representatives of fossil Ochotonidae. Due to the similarity with fauna of Aliveri in Greece, the association of small mammals of Snegotin (which includes Albertona) was included in the MN4 zone.     

Prostrepsiceros and Protragelaphus (Artiodactyla,Mammalia) from the Late Miocene locality of Chomateri (Attica,Greece)

The Late Miocene fossiliferous locality of Chomateri is located close to the classic locality of Pikermi, but unfortunately its faunal context is not sufficiently known. Some fossil remains of spiral-horned antelopes from Chomateri are assigned to Prostrepsiceros rotundicornis and Protragelaphus skouzesi. The co-existence of P. rotundicornis and P. skouzesi is relatively rare in the Late Miocene, but well-documented in Pikermi. Comparisons of the Chomateri material support a middle Turolian age for the Chomateri locality and suggest a close chronological relation to the Pikermi fauna.     

Evidence for the longest stratigraphic range of a post-Triassic Ichthyosaur: a Leptonectes tenuirostris from the Pliensbachian (Lower Jurassic) of Switzerland

A three-dimensional well-preserved ichthyosaur skull and parts of the postcranial skeleton are attributed to the species Leptonectes tenuirostris (Conybeare, 1822). It was found vertically embedded in Pliensbachian deposits representing three successive biozones (ibex to margaritatus Zone). The find is dated as early Late Pliensbachian (margaritatus Zone) by a rich ammonite and ostracod fauna. It is the first record of the genus Leptonectes from Switzerland and from the Late Pliensbachian. It is so far the best preserved and most complete ichthyosaur from this time interval worldwide. With diagnostic specimens known from the Rhaetian (Late Keuper) up to the early Late Pliensbachian (Middle Liassic), L. tenuirostris (Conybeare, 1822) has the most extensive stratigraphic range documented for any post-Triassic ichthyosaur so far.     

Change in diversity,ecological significance and biogeographical relationships of the Mediterranean Miocene toothed whale fauna

The main evolutionary trend in the Mediterranean Miocene toothed whale fauna is related (1) to the change in diversity and (2) to the turnover in community structure. Diversity increases from Upper Aquitanian–Lower Burdigalian to Burdigalian–Langhian, when it reaches its maximum. Starting from this time, diversity decreases progressively. The Early Miocene (Upper Aquitanian–Lower Burdigalian) Mediterranean toothed whale fauna, as well as the extramediterranean ones, is characterised by a high number of endemic taxa and by the prevalence of longirostral forms living in estuarine-neritic environments. A more diversified fauna spreading in neritic and pelagic environments characterises the Burdigalian–Langhian age, while an increase in pelagic forms and the nearly complete disappearance of some archaic longirostral taxa is typical of the Serravallian–Messinian fauna. Decrease in diversity and disappearance of archaic longirostral taxa are also recorded, at more general scale, in the Late Miocene extramediterranean fossil bearing deposits. These events can be related to the progressive global climatic deterioration, starting from Middle Miocene. From a biogeographic point a view, we can outline some relationships between the Mediterranean and western North Atlantic Miocene faunas. Closer affinities are observed between the Baltringen fauna and the northern Atlantic one, because of the presence of the genera Pomatodelphis and Zarhachis (platanistids) in both areas. In the Miocene Mediterranean and in North Atlantic, the delphinids are apparently absent as well as other extant delphinoid groups even if erroneously recorded in the past.     

Hyaenictitherium minimum, a new ictithere (Mammalia, Carnivora, Hyaenidae) from the Late Miocene of Toros-Menalla, Chad

A new species of Hyaenidae, Hyaenictitherium minimum, is described in the carnivore fauna of the Late Miocene layers of Toros-Menalla (Chad). Its size is similar to that of a jackal and it had probably a similar ecological niche. It is found in several fossil-bearing localities of this area. The genus Hyaenictitherium is known from the early Late Miocene in Eurasia from China to Spain; the Chadian material is, perhaps with some specimens from Sahabi and Lothagam, the earliest occurrence of the genus in Africa. It results certainly from Eurasian migration, which will have to be taken into account for the analysis of the bulk of the fauna. To cite this article: L. de Bonis et al., C. R. Palevol 4 (2005).     

Les otolithes des Téléostéens du Plio-Pleistocène belge

The study of approximately twenty three thousand otoliths from Plio-Pleistocene sites in the harbour region of Antwerp, as well as a critical revision of already published material allowed us to identify a teleostean fauna with fourty seven species (including two subspecies and seven species in open nomenclature). Two of these, Ophidion springeri and Uranoscopus septentrionalis are new to science. The fauna is typical for coastal waters slightly warmer than those of the actual North Sea and caracterized by the predominance of Gadidae. Biostratigraphically this fauna is well individualized with respect to preceeding ones (only seven species in common with the Miocene fauna) and those following twenty-three species in common with the extant fauna). The following biostratigraphical subdivision has been recognized in the Upper Miocene and the Plio-Pleistocene of the Antwerp region: (1) an association with «genus? Macrouridarum labiatus, Trisopterus sculptus, Gadiculus benedeni and Trisopterus luscus spectabilis, in the Sands of Deurne (Upper Miocene); (2) an association with Gadiculus benedeni and Merlangius pseudaeglephinus, in the Formation of Kattendijk (Pliocene); (3) an association with Gadiculus verticalis and Merlangius pseudaeglefinus in the Formation of Lillo (Plio-Pleistocene); (4) an association with Gadus morhua and Merlangius pseudaeglefinus in the so called «Icenian deposits not present in Belgium but found in the Netherlands.     

A juvenile skull of Acerorhinus yuanmouensis (Mammalia: Rhinocerotidae) from the Late Miocene hominoid fauna of the Yuanmou Basin (Yunnan,China)

A unique juvenile skull bearing both milk premolars and unerupted but fully developed permanent premolars and molars (observed using X-ray microcomputed tomography), and some isolated upper cheek teeth, all from the Late Miocene hominoid fauna of the Yuanmou Basin (Yunnan, China), closely resemble craniodental material of Acerorhinus yuanmouensis Zong, 1998 from the same locality, and are referred to this species. A phylogenetic analysis based on 214 craniodental morphological characters scored for 31 terminal taxa reveals that A. yuanmouensis should be assigned to the genus Acerorhinus indeed. The newly discovered specimens improve our understanding of this species, especially with respect to the morphology of the milk premolars and premolars. Two intraspecific variations in the upper premolars are noted: a lingual bridge may be present or absent, and the lingual cingulum continuous or reduced. The analysis also indicates that: the phylogenetic status of Acerorhinus lufengensis Deng and Qi, 2009 should be reconsidered; “Aceratherium” huadeensis Qiu, 1979 does neither belong to Aceratherium nor Acerorhinus, and its phylogenetic status remains debatable.     

A new Miocene fauna of snakes from eastern Siberia,Russia.: Was the snake fauna largely homogenous in Eurasia during the Miocene?

Togay, a locality from Ol’khon Island, Baykal Lake, Russia, has yielded a fauna of snakes of Late Middle or early Late Miocene age. It is located in a broad area from which no Neogene snake has been reported; therefore, it represents an important landmark. The fauna includes a non-erycine boid, two or three colubrids, a viperid of the ‘oriental complex’ of Vipera, and perhaps another viperid. This assemblage is astonishingly reminiscent of the snake faunas from the late Early and early Middle Miocene from western and central Europe, it being understood that Miocene faunas are practically unknown in the geographically intermediary area. It may be entertained whether a homogenous snake fauna inhabited Eurasia (except the southern part of the continent) during the Miocene.     

Early Miocene Amphibians (Caudata, Salientia) from the Mokrá-Western Quarry (Czech Republic) with comments on the evolution of Early Miocene amphibian assemblages in Central Europe

The Mokrá-Western Quarry exhibits the rare occurrence of Early Miocene (MN 4) vertebrate fauna within the area of the eastern part of Central Europe. In addition to a rich fauna of reptiles and mammals, two fossiliferous karst joints (Mokrá-Western Quarry, 1/2001 Turtle Joint and Mokrá-Western Quarry, 2/2003 Reptile Joint) yielded a rich fauna of amphibians including 13 amphibian taxa: Salamandridae: Mioproteus sp., Chelotriton sp., type I, Chelotriton sp., type II, Triturus aff. roehrsi, Triturus cf. marmoratus, Triturus sp. (T. cristatus species group), Chioglossa meini, Mertensiella mera, Salamandridae gen. and sp. indet.; Pelobatidae: Pelobates sanchizi; Ranidae: Rana sp. (synklepton Rana esculenta); Bufonidae: Bufo sp. The first records of the West European species Triturus cf. marmoratus and Chioglossa meini are reported from the eastern part of Central Europe indicating the wide distribution of those taxa throughout the whole of Europe as early as MN 4. The oldest known record of Pelobates sanchizi documents the Early Miocene presence of representatives closely related to the extinct Late Oligocene representatives of Pelobates. The slow evolution of amphibian species is documented by the presence of Triturus cf. marmoratus and the oldest known occurrence of the extinct salamander Mertensiella mera.     

Early Miocene reef- and mudflat-associated gastropods from Makran (SE-Iran)

A new gastropod fauna of Burdigalian (early Miocene) age is described from the Iranian part of Makran. The fauna comprises 19 species and represents three distinct assemblages from turbid water coral reef, shallow subtidal soft-bottom and mangrove-fringed mudflat environments in the northern Indian Ocean. Especially the reef-associated assemblage comprises largely new species. This is explained by the rare occurrence of reefs along the northern margin of the Miocene Indian Ocean and the low number of scientific studies dealing with the region. In terms of paleobiogeography, the fauna corresponds well to coeval faunas from the Pakistani Balochistan and Sindh provinces and the Indian Kathiawar, Kutch and Kerala provinces. During the early Miocene, these constituted a discrete biogeographic unit, the Western Indian Province, which documents the near complete biogeographic isolation from the Proto-Mediterranean Sea. Some mudflat taxa might represent examples of vicariance following the Tethys closure. The fauna also displays little connection with coeval faunas from Indonesia, documenting a strong provincialism within the Indo-West Pacific Region during early Miocene times. Neritopsis gedrosiana sp. nov., Calliostoma irerense sp. nov., Calliostoma mohtatae sp. nov. and Trivellona makranica sp. nov. are described as new species.     

Euroceros bulgaricus gen. nov., sp. nov. from Hadzhidimovo (SW Bulgaria) (Late Miocene) - the first European record of Hornbills (Aves: Coraciiformes)

A new genus and species of ground-hornbill are described from the Late Tertiary (Late Miocene) fluviative deposits in Hadzhidimovo (Hadzhidimovo-1, or Hadzhidimovo-Girizite locality), in southwestern Bulgaria. The cranial half of a scapula and the distal third of a femur have been recovered. The morphology distinguishes the specimen from the recent species of Bucorvus, the closest genus. Most interesting is the morphology of the scapula (the holotype), which differs considerably from the ground-hornbills in some structures (e.g. the clavicular joint) while being surprisingly similar in other structures (e.g. the humeral articular joint). The last peculiarity may indicate that the fossil species was a more primitive form having relatively poorer flying capacity in comparison to other ground-hornbills. This first record of Bucerotidae in Europe proves the occurrence of the family in the Tertiary of the western parts of the Eurasian continent. It came from the richest Bulgarian locality of the Hipparion fauna, from which ca. 36 species of large mammals and birds are now known.     

Palynology and age of the Early Oligocene units in Çardak-Tokça Basin, Southwest Anatolia: Paleoecological implications

In this study, the lignite bearing sediments of Çardak-Tokça basin exposed in southwest Anatolia, were palynologically examined. A well preserved and diverse palynomorph assemblage indicating an Early Oligocene age was recovered from the Hayrettin and Tokça formations. The palynomorph assemblage is dominated by Pinus, Sparganiaceae, Juglandaceae and diverse tricolpate and tricolporate pollen. In addition a few species of marine dinoflagellate cysts were encountered as well. The Early Oligocene age is based primarily on the presence of stratigraphic markers such as: Boehlensipollis hohli, Slowakipollis hippophaëoides, Aglaoreidia cyclops, Dicolpopollis kockeli, Compositoipollenites rhizophorus ssp. burghasungensis, Mediocolpopollis compactus ssp. ellenhausensis, Pentapollenites pentangulus, Subtriporopollenites simplex and Intratriporopollenites instructus. Palynological data indicate a humid subtropical climatic conditions during the deposition of the Çardak-Tokça sediments. Ecological analysis of the palynomorph assemblage identifies several paleo-associations of montana, lowland and slope, swamp and water-edge and freshwater aquatic elements. In this study, Çardak-Tokça, Çank?r?-Çorum, Thrace and southwest Anatolian molasse basins (Kale-Tavas and Denizli) were correlated in accordance with their palynostratigraphic content and the results show that the deposition took place during the Early Oligocene in the Çardak-Tokça basin. This basin is older than Thrace basin and southwest Anatolian molasse basins (Kale-Tavas and Denizli molasse) which were deposited during the Late Oligocene-Early Miocene.     

Late Pleistocene and Holocene chitons (Mollusca, Polyplacophora) from Uruguay: Palaeobiogeography and palaeoenvironmental reconstruction in mid latitudes of the southwestern Atlantic

Here we present a revision of the fossil record of chitons (Mollusca, Polyplacophora) of Late Pleistocene and Holocene marine deposits of Uruguay and discuss their potential for palaeoenvironmental reconstruction. Chitons were recorded as isolated valves in bivalve- and gastropod-rich assemblages. They are represented by the species Chaetopleura angulata (Spengler), C. isabellei (d’Orbigny), C. asperrima (Gould) and Ischnochiton striolatus (Gray). The last two species are recorded for the first time as fossils not only in Uruguay but also in South America. Exclusively recorded for the Late Pleistocene is the warm water species I. striolatus, whose current southern range limit is located in Santa Catarina, Brazil. Higher temperatures than at present are inferred for the Uruguayan coast during the Late Pleistocene, probably driven by the southward influence of the warm Brazilian Current along the eastern margin of South America. Chitons from Holocene deposits show a wider geographic distribution along the Uruguayan coast in comparison to current distributions. They are recorded where an estuarine and eurihaline fauna lives today, which suggests the former existence of marine conditions. This is explained by the westward displacement of the marine front in the Uruguayan coast during the Holocene transgressive events. Chitons have proven to be useful palaeoenvironmental proxies for the reconstruction of salinity and temperature trends in the Uruguayan coast during the climatic oscillations of the Late Pleistocene and Holocene epochs. As such, their patterns of distribution are related to the recent physical evolution of the Southwestern Atlantic Ocean.     

Messinian Lago-Mare mollusc fauna from the Gorgona Island slope, Tyrrhenian Sea

Late Miocene Lago-Mare macrofossiliferous sediments were recovered in the northeastern Tyrrhenian Sea by dredging the continental slope off Gorgona Island, Tuscan Archipelago, at 300-470 m depth. The fossil assemblage consists of a rich lymnocardiid bivalve fauna dominated by Pontalmyra ex gr. P. incerta (Deshayes), associated with Dreissena ex gr. D. rostriformis (Deshayes), Pontalmyra cf. partschi (Mayer), “Limnocardium” sp., the gastropods Melanopsis narzolina (D’Archiac), Melanopsis sp. and cf. Saccoia sp. All bivalve taxa recognized at species level are of Paratethyan (Pontian) affinity and widespread in the Late Miocene of the Mediterranean Basin while M. narzolina has so far only recorded from the Mediterranean Basin. This finding represents the most diverse Lago-Mare macrofauna reported thus far from any submerged location in the Mediterranean Basin and documents that the post-evaporitic Cusercoli Formation contributes to the syn-rift neoautochthonous units of this sector of the Northern Tyrrhenian Sea.     

Sexual and individual morphometric variation in Libycosaurus (Mammalia, Anthracotheriidae) from the Maghreb and Libya

Anthracotheres of late Middle Miocene and Late Miocene age have been described from several localities in northern Africa, all of them currently assigned to the genus Libycosaurus Bonarelli, although in several previous works they were assigned to Merycopotamus Falconer and Cautley, a considerably younger and specialised form from the Indian subcontinent which has quite different dental and cranial morphology. Three species of Libycosaurus have been named, but there has been some doubt about the morphometric variation within the various species, with some authors such as Gaziry (1987) placing the fossils from Sahabi (Latest Miocene, Libya) and Beglia (end Middle Miocene to basal Late Miocene, Tunisia) into the same species despite marked size differences, and others (Ducrocq et al., 2001) creating a species for a restricted sample of small specimens from Nementcha (Late Middle Miocene, Algeria), but which overlaps with the range of size variation of Beglia fossils. The aim of this paper is to examine the available samples in greater depth in order to understand the morphometric variation in these anthracotheres. It is confirmed that the Beglia sample is quite variable (Black, 1972), both morphologically and metrically, but it is concluded that it nevertheless belongs to a single species, because specimens from some of the localities within the Beglia Formation (e.g. Loc. 17 in the lower levels at Beglia) span the entire range of variation (Pickford, 1994). The sample from Nementcha cannot be distinguished from the Beglia sample on any consistent metric or morphological basis, but in general the specimens fall at the low end of the range of variation of the Beglia sample. It is thus likely that L. algeriensis is a synonym of L. anisae. The Sahabi and Chad samples (L. petrocchii), in contrast, fall above the known range of variation of the Beglia material in almost all metric features, but are close to it morphologically, and they are considered to represent a species distinct from the Beglia sample.