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Institute
Taphonomy and palaeoecology of Laetoli as well as Makuyuni, Arusha region in northern Tanzania
(2004)
This thesis is the result of the Hominid Corridor research Project in Tanzania since 1993 to 1995 that include Pliocene and Pleistocene localities. The localities under study include Laetoli and Manyara area in Arusha Region, northern Tanzania. The thesis has the following specific objectives: firstly, to identify taxa recovered from the studied assemblages; secondly, to underpin taphonomic history of the assemblages under study; thirdly, to elucidate further palaeoecological reconstruction of the assemblages; and finally, to examine surface fossil fauna modifications including agents of modifications either hominids or carnivores.
The Upper Laetolil Beds are dated at 3.5 million years ago (Ma) and the Ndolanya Beds are bracketed in age between 3.5 and 2.41 Ma. The Naibadad Beds, also from Laetoli area, are date to be between 2.2 to 2.1 Ma. The Naibadad Beds are correlated with the base of Bed I at Olduvai Gorge. There are so far no absolute dates for Manyara assemblages. Based on biostratigraphic correlation, the younger overlying unit, the Upper Manyara Beds are estimated to belong to Later Pleistocene and the Lower Manyara Beds are estimated to belong to Early Pleistocene. The Upper Manyara Beds are correlated to the age of Bed III at Olduvai Gorge, while the Lower Manyara Beds are interpreted to span the same contemporaneity with the upper part of Bed II at Olduvai Gorge.
At Laetoli localities, terrestrial mammals while localities from Manyara besides terrestrial mammals dominate fauna; they include aquatic species such as fish, crocodiles and hippopotamus. The main families recovered from Upper Laetolil Beds complement those already recovered from former research works by other workers. This is also true for the younger overlying stratigraphic horizon, the Upper Ndolanya Beds. Thus, mammalian families recovered from Upper Laetolil Beds include Bovidae, Carnivora, Elephantidae, Equidae, Lagomorpha, Suidae, Rodentia, Hominoidea and Rhenocerotidae. Remains of an invertebrate, Gastropoda were also recovered. For Upper Ndolanya Beds include almost the same families recovered from Upper Laetolil Beds, but based on former recovery of fossil fauna, these Beds outnumber greatly the Upper Laetolil Beds in bovid composition by 20 per cent. Such a change in species composition is noticed also from South African localities and East African localities such as the East Turkana. This is interpreted to be due to climatic change drier environments that included species adapted to such palaeoclimates.
For the first time, our team has been able to retrieve specimens identifiable to taxa, a pattern that not possible from previous workers who claimed to have recovered too sparse specimens to be identifiable to any taxon.
The Upper Manyara Beds as well as Lower Manyara taxonomic composition include aquatic species besides the large terrestrial mammalian fauna retrieved from there. In due regard, the former horizon is attributed to have affinity with Olduvai Bed III components and the latter, older horizon, is attributed to have affinity with upper parts of Bed II times at Olduvai Gorge. The Lower Manyara Beds can be said to have, in relative terms, affinity to species recovered from site RC 11 of the Chiwondo Beds, Malema region in northern Malawi, although the former site may be equable to the terminal age of the latter locality.
Fossil hominid remains; attributable to genus Homo and possibly species Homo erectus have been recovered from two localities, Mk 2 and Mk, along Lower Manyara Beds. On the other hand, stone tools, identified to belong to the Acheulian industrial technocomplex, were recovered from site Mk 4.
All of fossil fauna from Laetoli sites were mostly exfoliated and there shows to be little effect in terms of hydrodynamic sorting of the fossil bones. However, intense carnivore activity is witnessed due to the almost one to one ratio of proximal to distal ends. This is also true for the Lower Manyara Beds locality. Through examination of surface modifications of the fossil fauna, it has been established that there was carnivore consumption of ungulates. There is no evidence of hominid involvement that has to be testified by stone tools.
Highlights
• Protocol for extracting and analyzing pollen grains from fossil insects
• Individual fossil grains can be analyzed using a combined approach
• Simple and fast TEM embedding and sectioning protocol
• Protocol enables a taxonomic assignment of pollen
Summary
This protocol explains how to extract pollen from fossil insects with subsequent descriptions of pollen treatment. We also describe how to document morphological and ultrastructural features with light-microscopy and electron microscopy. It enables a taxonomic assignment of pollen that can be used to interpret flower-insect interactions, foraging and feeding behavior of insects, and the paleoenvironment. The protocol is limited by the state of the fossil, the presence/absence of pollen on fossil specimens, and the availability of extant pollen for comparison.
Fossil dental remains are an archive of unique information for paleobiological studies. Computed microtomography based on X-ray microfocus sources (X-μCT) and Synchrotron Radiation (SR-μCT) allow subtle quantification at the micron and sub-micron scale of the meso- and microstructural signature imprinted in the mineralized tissues, such as enamel and dentine, through high-resolution “virtual histology”. Nonetheless, depending on the degree of alterations undergone during fossilization, X-ray analyses of tooth tissues do not always provide distinct imaging contrasts, thus preventing the extraction of essential morphological and anatomical details. We illustrate here by three examples the successful application of neutron microtomography (n-μCT) in cases where X-rays have previously failed to deliver contrasts between dental tissues of fossilized specimen.
Significance
Identifying the earliest members of the genus Homo is crucial for understanding when and where selective pressures resulted in its emergence from a Plio-Pleistocene hominin taxon. Our revision of a large part of the dental fossil record from southern Africa provides evidence suggesting a paucity of Homo remains and indicates increased levels of dental variation in australopith taxa. Results of the Ba/Ca, Sr/Ca, and elemental mapping of enamel and dentine also indicate that some of the purported Homo specimens show a paleoecological signal similar to that of the australopiths.
Abstract
The origins of Homo, as well as the diversity and biogeographic distribution of early Homo species, remain critical outstanding issues in paleoanthropology. Debates about the recognition of early Homo, first appearance dates, and taxonomic diversity within Homo are particularly important for determining the role that southern African taxa may have played in the origins of the genus. The correct identification of Homo remains also has implications for reconstructing phylogenetic relationships between species of Australopithecus and Paranthropus, and the links between early Homo species and Homo erectus. We use microcomputed tomography and landmark-free deformation-based three-dimensional geometric morphometrics to extract taxonomically informative data from the internal structure of postcanine teeth attributed to Early Pleistocene Homo in the southern African hominin-bearing sites of Sterkfontein, Swartkrans, Drimolen, and Kromdraai B. Our results indicate that, from our sample of 23 specimens, only 4 are unambiguously attributed to Homo, 3 of them coming from Swartkrans member 1 (SK 27, SK 847, and SKX 21204) and 1 from Sterkfontein (Sts 9). Three other specimens from Sterkfontein (StW 80 and 81, SE 1508, and StW 669) approximate the Homo condition in terms of overall enamel–dentine junction shape, but retain Australopithecus-like dental traits, and their generic status remains unclear. The other specimens, including SK 15, present a dominant australopith dental signature. In light of these results, previous dietary and ecological interpretations can be reevaluated, showing that the geochemical signal of one tooth from Kromdraai (KB 5223) and two from Swartkrans (SK 96 and SKX 268) is consistent with that of australopiths.
Neanderthal diet has been on the spotlight of paleoanthropological research for many years. The majority of studies that tried to reconstruct the diet of Neanderthals were based on the analysis of zooarchaeological remains, stable isotopes, dental calculus and dental microwear patterns. In the past few years, there have been a few studies that linked dental macrowear patterns of Neanderthals and modern humans to diet and cultural habits. However, they mostly focused on maxillary molars. Although mandibular molars have been widely used in microwear dietary research, little is known about their usage at the macroscopic scale to detect information about human subsistence strategies. In this study, we compare the macrowear patterns of Neanderthal (NEA), fossil Homo sapiens (FHS), modern hunter-gatherers (MHG), pastoralists, early farmers and Australian Aborigines from Yuendumu mandibular molars in order to assess their utility in collecting any possible information about dietary and cultural habits among diverse human groups. We use the occlusal fingerprint analysis method, a quantitative digital approach that has been successfully employed to reconstruct the diet of living non-human primates and past human populations. Our results show macrowear pattern differences between meat-eater MHG and EF groups. Moreover, while we did not find eco-geographical differences in the macrowear patterns of the fossil sample, we found statistically significant differences between NEA and FHS inhabiting steppe/coniferous forest. This latter result could be associated with the use of distinct technological complexes in these two species, which ultimately could have allowed modern humans to exploit natural resources in a different way compared to NEA.
Palaeosphryon menatensis gen. et sp. nov., first unambiguous representative of the longhorn beetle subfamily Prioninae from the Paleocene of Menat (France), is described and illustrated. The new fossil is placed into the tribe Prionini, showing some similarities with some species of the extant genera Osphryon (Papua New Guinea) and Titanus (Brazil, Colombia, Guianas, Ecuador, Peru), viz. in general body shape, antennomere 3 as long as first and second together but shorter than the length of fourth plus fifth, elongate elytra, and small spines on the lateral margin of the pronotum disposed in a relatively similar way as in Osphryon. Nevertheless, the exact affinities of the new fossil within the Prionini remain uncertain because of the lack of a recent phylogenetic analysis in which it could be integrated. This fossil beetle is exceptional for its very large size, with a body 70 mm long. Some other large longhorn beetles have been found in the same outcrop, and are awaiting description. The positions of the previously described Cerambycidae from Menat are also discussed. This exceptional fauna of Cerambycidae is in accordance with the current palaeoenvironmental reconstruction for the Menat Konservat-Lagerstätte, as a small maar lake surrounded by a warm and humid, probably evergreen forest.
The ammonoids of the family Maenioceratidae from Givetian sedimentary rocks of the Anti-Atlas (Morocco) are investigated. The study is based on new collections stored in the Museum für Naturkunde, Berlin. The genera Maenioceras Schindewolf, 1933 and Afromaenioceras Göddertz, 1987 are revised; the genus Trimaenioceras is newly described. The species Maenioceras afroterebratum sp. nov., Maenioceras mzerrebense sp. nov., Maenioceras oufranense sp. nov., Maenioceras beckeri sp. nov., Afromaenioceras sulcatostriatum (Bensaïd, 1974), Afromaenioceras hiemale sp. nov., Afromaenioceras bensaidi sp. nov., Afromaenioceras brumale sp. nov., Afromaenioceras crassum (Bensaïd, 1974), Trimaenioceras klugi gen. et sp. nov., Trimaenioceras eculeus gen. et sp. nov., Trimaenioceras fuscina gen. et sp. nov. and Trimaenioceras paucum gen. et sp. nov. are described in detail.
The first sawfly from the Oligocene of Céreste (Southern France (Hymenoptera: Tenthredinidae)
(2024)
Luberotenthredo cerestensis gen. et sp. nov. is the first record of the sawfly family Tenthredinidae from the Oligocene of Céreste (Southern France). This taxon is described and illustrated based on a well-preserved specimen. This genus resembles the extant genus Perineura (subfamily Tenthredininae, tribe Perineurini) with which it shares forewing venation similarities and numerous morphological characters. This new taxon is the first fossil representative of the tribe Perineurini and can be used as a calibration point for future investigation of the diversification of the family Tenthredinidae.
Here, we report on 33 molluscan species from Miocene ’Calcari a Lucina’ hydrocarbon-seep deposits in northern Italy. Three new species are described: the chilodontaid gastropod Putzeysia diversii sp. nov., the lucinid bivalve Miltha (sensu lato) romaniae sp. nov., and Sisonia ultimoi sp. nov., a heterodont bivalve of uncertain taxonomic affinity. Fourteen species are described in open nomenclature. The common but enigmatic gastropod species Phasianema taurocrassa is here suggested to belong to the seguenzioid genus Cataegis. Most gastropod species are inhabitants of the deep-sea floor in general, and are not restricted (obligate) to sites of hydrocarbon-seepage. The gastropod Putzeysia diversii sp. nov. and the bivalve Sisonia ultimoi sp. nov. are the geologically oldest members of their genera known to date. While the genus Putzeysia is geographically restricted to the NE Atlantic Ocean and the Mediterranean Sea, Sisonia ultimoi sp. nov. represents another link of the Miocene Mediterranean seep fauna to that of the central Indo-West Pacific Ocean.
Lacewing larvae in the Cretaceous were more diverse in appearance than they are today, best documented by numerous fossils preserved in amber. One morphotype of an unusual larva from about 100 Ma old Kachin amber (Myanmar) was formally recognised as a distinct group called Ankyloleon. The original description erected a single formal species, Ankyloleon caudatus. Yet, it was indicated that among the five original specimens, more species were represented. We here report five new specimens. Among these is the so far largest as well as the so far smallest specimen. Based on this expanded material we can estimate certain aspects of the ontogenetic sequence and are able to recognise a second discrete species, Ankyloleon caroluspetrus sp. nov. We discuss aspects of the biology of Ankyloleon based on newly observed details such as serrations on the mandibles. Long and slender mouthparts, legs and body together with a weakly expressed outer trunk segmentation provide indications for a lifestyle hunting for prey in more confined spaces. Still many aspects of the biology of these larvae must remain unclear due to a lack of a well comparable modern counterpart, emphasising how different the fauna of the Cretaceous was.
The genus Plateosaurus is one of the most challenging aspects of early-diverging sauropodomorph taxonomy, with a total of 29 names and a century of revisions. As a result, nomina dubia have been treated as synonyms of Plateosaurus trossingensis, adding to the confusion about the extent of morphological variability. Here, we provide a thorough revision of the taxonomy proposed by von Huene as well as the subsequent taxonomic interpretations of Plateosaurus through a complete inventory of the specimens stored in the Palaeontological Collection of Tübingen. We reassess the status of the specimens in the collection and propose that the sauropodomorph-bearing layers are not necessarily monospecific. Most of the original fieldwork documentation has been lost, but we use the taxonomy established by von Huene as a historical reference point to reconstruct what was known at the time. This revised taxonomy of Plateosaurus narrows the genus to three species: Pl. trossingensis, Pl. longiceps, and Pl. gracilis (as a metataxon), and restricts the genera Gresslyosaurus and Pachysaurus to large and robust individuals, as pragmatic decisions aimed to test their affinities to other Late Triassic sauropodomorphs. Future studies should consider not only morphological variability, but also stratigraphy, palaeogeography, and environmental data when delineating species within and outside the Plateosaurus plexus.
The Miocene is a key time in the evolution of African mammals and their ecosystems witnessing the origin of the African apes and the isolation of eastern coastal forests through an expanding biogeographic arid corridor. Until recently, however, Miocene sites from the southeastern regions of the continent were unknown. Here we report discovery of the first Miocene fossil teeth from the shoulders of the Urema Rift in Gorongosa National Park, Mozambique, at the southern East African Rift System. We provide the first 1) radiometric age determinations of the fossiliferous Mazamba Formation, 2) reconstructions of past vegetation in the region based on pedogenic carbonates and fossil wood, and 3) description of fossil teeth from the southern rift. Gorongosa is unique in the East African Rift System in combining marine invertebrates, marine vertebrates, terrestrial mammals, and fossil woods in coastal paleoenvironments. The Gorongosa fossil sites offer the first evidence of persistent woodlands and forests on the coastal margins of southeastern Africa during the Miocene, and an exceptional assemblage of fossil vertebrates including new species. Further work will allow the testing of hypotheses positing the formation of a northeast-southwest arid corridor isolating species on the eastern coastal forests from those elsewhere in Africa.
Brief The Miocene is a key time in the evolution of African mammals and their ecosystems encompassing hominine origins and the establishment of an arid corridor that isolated eastern Africa’s coastal forests. Until now, however, Miocene sites from southeastern Africa have been unknown. We report the discovery of the first Miocene fossil sites from Gorongosa National Park, Mozambique, and show that these sites formed in coastal settings. We provide radiometric ages for the fossiliferous sediments, reconstructions of past vegetation based on stable isotopes and fossil wood, and a description of the first fossil teeth from the region. Gorongosa is the only paleontological site in the East African Rift that combines fossil woods, marine invertebrates, marine vertebrates, and terrestrial mammals. Gorongosa offers the first evidence of persistent woodlands and forests on the coastal margins of southeastern Africa during the Miocene.
The Miocene was a key time in the evolution of African ecosystems witnessing the origin of the African apes and the isolation of eastern coastal forests through an expanding arid corridor. Until recently, however, Miocene sites from the southeastern regions of the continent were unknown. Here, we report the first Miocene fossil teeth from the shoulders of the Urema Rift in Gorongosa National Park, Mozambique. We provide the first 1) radiometric ages of the Mazamba Formation, 2) reconstructions of paleovegetation in the region based on pedogenic carbonates and fossil wood, and 3) descriptions of fossil teeth. Gorongosa is unique in the East African Rift in combining marine invertebrates, marine vertebrates, reptiles, terrestrial mammals, and fossil woods in coastal paleoenvironments. The Gorongosa fossil sites offer the first evidence of woodlands and forests on the coastal margins of southeastern Africa during the Miocene, and an exceptional assemblage of fossils including new species.
Thylacocephalans are enigmatic euarthropods, known at least from the Silurian to the Cretaceous. Despite remaining uncertainties concerning their anatomy, key features can be recognised such as a shield enveloping most of the body, hypertrophied compound eyes, three pairs of raptorial appendages and a posterior trunk consisting of eight up to 22 segments bearing appendages and eight pairs of gills. Well-known for its euarthropod diversity, the La Voulte-sur-Rhône Lagerstätte (Callovian, Middle Jurassic, France) has provided many remains of four thylacocephalan species so far: Dollocaris ingens, Kilianicaris lerichei, Paraostenia voultensis and Clausocaris ribeti. In this paper, we study the type material as well as undescribed material. The re-description of La Voulte thylacocephalans reveals an unexpected diversity, with the description of two new species, Austriocaris secretanae sp. nov. and Paraclausocaris harpa gen. et sp. nov., and of specimens of Mayrocaris, a taxon originally described from Solnhofen Lagerstätten. We also reassign Clausocaris ribeti to Ostenocaris. The reappraisal of La Voulte thylacocephalans also provides important insight into the palaeobiology of Thylacocephala. New key anatomical features are described, such as an oval structure or a putative statocyst, which indicate a nektonic or nektobenthic lifestyle. Finally, we document a juvenile stage for Paraostenia voultensis.
Oligoptilomera luberonensis gen. et sp. nov., first fossil representative of the gerrid subfamily Ptilomerinae, is described and figured from the Oligocene of Murs (Vaucluse, Southern France). Extant Ptilomerinae live in streams in warm climates, of the Indo-Malaysian, eastern Palaearctic, and Papouan regions. The discovery of this Oligocene French Ptilomerinae is in accordance with the putative age of the subfamily, at least older than the Eocene, and with the Indo-Malaysian affinities previously recorded for some other insects from the Oligocene of France. The two insect assemblages of Murs and Céreste are compared and the differences discussed. Although of similar ages, that from Murs was possibly corresponding to a more shallow water paleolake than that of Céreste.
This work provides the first revision and illustration of the type material of the ammonite species Parahoplites abichi Anthula [Type species of Protacanthoplites Tovbina] and Parahoplites aschiltaensis Anthula [Type species of Acanthohoplites Sinzow] from the upper Aptian (Lower Cretaceous) of Dagestan, Russia. The close affinities and synonymy between these two species are confirmed, and Acanthohoplites aschiltaensis is here retained as the senior valid name by its long quoting history and its historical use as a zonal index of the upper Aptian. The genus Protacanthoplites should be thus synonymised with Acanthohoplites by priority in the date of publication. Comparison with, and distinction from, closely allied Acanthohoplitidae is provided.
The railway cutting near Oberrödinghausen at the northern margin of the Rhenish Mountains is the cardinal section for the investigation of Early Tournaisian (Early Carboniferous; Mississippian) ammonoids. The ammonoids from the Hangenberg Limestone (= Gattendorfia Limestone) of this and neighbouring outcrops are revised here, using the historical collections as well as undescribed new material. The ammonoid assemblages are composed of a total of 67 species, which occur in four successive ammonoid zones. The assemblages are composed of predominant prionoceratids (Order Goniatitina) with the twenty genera Mimimitoceras (two species), Globimitoceras (one species), Paragattendorfia (two species), Kornia (three species), Stockumites (eleven species), Acutimitoceras (two species), Costimitoceras (one species), Nicimitoceras (four species), Imitoceras (one species), Voehringerites (one species), Gattendorfia (eight species), Zadelsdorfia (two species), Kazakhstania (one species), Gattenpleura (one species), Weyerella (three species), Hasselbachia (three species), Paprothites (five species), Pseudarietites (three species), Rodingites (two species), Paralytoceras (one species) as well as subordinate eocanitids (Order Prolecanitida) with the genera Eocanites (eight species) and Nomismocanites (one species). The new genera Rodingites gen. nov. and Nomismocanites gen. nov. as well as the new species Mimimitoceras perditum sp. nov., Kornia fibula sp. nov., Kornia acia sp. nov., Stockumites parallelus sp. nov., Stockumites voehringeri sp. nov., Acutimitoceras ucatum sp. nov., Acutimitoceras paracutum sp. nov., Imitoceras initium sp. nov., Gattendorfia rhenana sp. nov., Gattendorfia bella sp. nov., Gattendorfia valdevoluta sp. nov., Gattendorfia schmidti sp. nov., Gattendorfia corpulenta sp. nov., Gattendorfia immodica sp. nov., Zadelsdorfia oblita sp. nov., Weyerella lenis sp. nov., Hasselbachia erronea sp. nov., Paprothites beckeri sp. nov., Paprothites kullmanni sp. nov., Eocanites delicatus sp. nov. and Nomismocanites raritas gen. et sp. nov. are described from Oberrödinghausen. Mimimitoceras mina sp. nov., Stockumites marocensis sp. nov., Zadelsdorfia zana sp. nov. and Kazakhstania kana sp. nov. are newly named for material from the Anti-Atlas of Morocco.
The early Tournaisian (Early Carboniferous; Mississippian) ammonoids from the classical abandoned limestone quarry of Gattendorf (Upper Franconia) are revised, using the historical collections as well as so far undescribed material. The ammonoid assemblage is composed of prionoceratid ammonoids of the six genera Mimimitoceras, Paragattendorfia, Stockumites, Acutimitoceras, Gattendorfia and Gattenpleura, which indicate a stratigraphic position near the Devonian–Carboniferous boundary in the earliest Carboniferous. The new species Stockumites hofensis sp. nov. and S. nonaginta sp. nov. are described.
The evolution and interrelationships of carnivorous squamates (mosasaurs, snakes, monitor lizards, Gila Monsters) are a contentious part of reptile systematics and go to the heart of conflict between morphological and molecular data in inferring evolutionary history. One of the best-preserved fossils in this motley grouping is “Saniwa” feisti Stritzke, 1983, represented by complete skeletons from the early-middle Eocene of Messel, Germany. We re-describe it on the basis of superficial examination, stereoradiography, and high-resolution X-ray computed tomography of new and published specimens. The scalation of the lizard is unique, consisting of small, keeled scales on the head (including a row of enlarged medial supraorbitals) and large, rhomboidal, keeled scales (invested by osteoderms) that covered the rest of the body. Two paired longitudinal rows of enlarged scales ran down the neck. The head was laterally compressed and box-shaped due to the presence of a strong canthal-temporal ridge; the limbs and tail were very long. Notable osteological features include: a toothed, strap-like vomer; septomaxilla with a long posterior process; palpebral with a long posterolateral process; a lacrimal boss and a single lacrimal foramen; a well-developed cultriform process of the parabasisphenoid; two hypoglossal (XII) foramina in addition to the vagus; a lack of resorption pits for replacement teeth; and possibly the presence of more than one wave of developing replacement teeth per locus. There are no osteological modifications suggestive of an intramandibular hinge, but postmortem displacement of the angular-prearticular-surangular complex in multiple specimens suggests that there might have been some degree of mobility in the lower jaw based on soft-tissue modifications. Using phylogenetic analyses on a data-set comprising 473 morphological characters and 46 DNA loci, we infer that a monophyletic Palaeovaranidae Georgalis, 2017, including Eosaniwa Haubold, 1977, lies on the stem of Varanidae Merrem, 1820, basal to various Cretaceous Mongolian taxa. We transfer feisti to the new genus Paranecrosaurus n. gen. Analysis of gut contents reveals only the second known specimen of the cryptozoic lizard Cryptolacerta hassiaca Müller, Hipsley, Head, Kardjilov, Hilger, Wuttke & Reisz, 2011, confirming a diet that was at least partly carnivorous; the preservation of the teeth of C. hassiaca suggests that the gastric physiology of Paranecrosaurus feisti (Stritzke, 1983) n. comb. had high acidity but low enzyme activity. Based on the foregoing and linear discriminant function analysis, we reconstruct P. feisti n. comb., as a powerful, widely roaming, faunivorous-carnivorous stem monitor lizard with a sensitive snout. If the molecular phylogeny of anguimorphs is correct, then many of the features shared by Helodermatidae Gray, 1837 and Varanidae must have arisen convergently, partly associated with diet. In that case, a reconciliation of morphological and molecular data would require the discovery of equally primitive fossils on the helodermatid stem.