Vertebrate Palaeontology

A comprehensive analysis of early dinosaur relationships raised the possibility that the group may have originated in Laurasia (Northern Hemisphere), rather than Gondwana (Southern Hemisphere) as often thought. However, that study focused solely on morphology and phylogenetic relationships and did not quantitatively evaluate this issue. Here, we investigate dinosaur origins using a novel Bayesian framework uniting tip-dated phylogenetics with dynamic, time-sliced biogeographic methods, which explicitly account for the age and locality of fossils and the changing interconnections of areas through time due to tectonic and eustatic change. Our analysis finds strong support for a Gondwanan origin of Dinosauria, with 99 % probability for South America (83 % for southern South America). Parsimony analysis gives concordant results. Inclusion of time-sliced biogeographic information affects ancestral state reconstructions (e.g., high connectivity between two regions increases uncertainty over which is the ancestral area) and influences tree topology (disfavouring uniting fossil taxa from localities that were widely separated during the relevant time slice). Our approach directly integrates plate tectonics with phylogenetics and divergence dating, and in doing so reaffirms southern South America as the most likely area for the geographic origin of Dinosauria. Dinosaurs are one of the most charismatic and heavily studied groups of vertebrates (e.g., Sereno 1999), but despite ongoing work and many recent discoveries, certain aspects of early dinosaurian evolution remain unclear. A recent study of early dinosaur relationships presented a new phylogeny that departs radically from existing models, but forms a broader framework for inferring dinosaur evolutionary history than those that were previously available (Baron et al. 2017a). While that study was focused primarily on early dinosaurian anatomy and relationships, a general observation was made: based upon the geographic locations of key taxa in their phylogeny, a Laurasian setting for dinosaur origin may have been plausible. This proposal ran contrary to an established consensus based on earlier phylogenies that favoured a Gondwanan and, more specifically, South American setting for this event (Nesbitt et al. 2009; Brusatte et al. 2010; Langer et al. 2010; Sues et al. 2011; Boyd 2015). In particular, the European presence of silesaurids, a close outgroup to dinosaurs, and some early dinosaur taxa (e.g., Saltopus) hinted that northern Pangaea may have played an important role in the earliest stages of dinosaur diversification. However, given the space limitations associated with the original article, no quantitative analyses were carried out to evaluate this alternative hypothesis rigorously. Here, we test the alternative hypothesis implicit in Baron et al. (2017a) by analysing their early dinosaur dataset using a novel integration of time-sliced Bayesian biogeographic models (Landis 2016) and tip-dated Bayesian phylogenetics (Drummond et al. 2012; Bielejec et al. 2014; Gavryushkina et al. 2017). The current method simultaneously estimates phylogenetic relationships, divergence dates, evolutionary rates and biogeographic patterns. Biogeographical changes are inferred using models that account for the changing relationships of continents (and thus dispersal abilities) over time-a 'chronobiogeographical' paradigm (Hunn & Upchurch 2001)-and this information provides potentially critical information for improving estimates of tree topology and divergence dates. For comparison, we also perform analogous analyses using traditional biogeographic parsimony-based methods. Previous studies have advocated a similar approach to integrating phylogeny, stratigraphy, morphology and bio-geography in a parsimony framework (reviewed in Rossie & Seiffert 2006). However, that implementation was problematic: as parsimony trees lack an explicit time element, only very simple time-sliced biogeographic models were possible (e.g., three areas and three time slices) and tree topology and divergence dates had to be assessed manually. The analyses attempted here tackle this complex multidimensional problem more efficiently , using explicit Bayesian computational approaches.

Archaeological sites at Affad (Sudan) are the only ones in northeastern Africa providing ostological remains of both African aurochs (Bos primigenius), dated to 50 kya, and domestic cattle, dated to 7–6 kya. The evidence enables studies of behavioral diversity between taurids. Strontium isotope analyses of the tooth enamel of both Pleistocene and Holocene ruminants suggest the local origin of these animals. The archaeozoological analysis reveals the temporal variability of environmental conditions was linked to a humid climate during the Marine Isotope Stage 3 (MIS3), which was more humid than the dry fluctuations during the African Humid Period. In addition, changes in the phenotype of cattle indicate that humans influenced domesticated animal behavior in the sub-Saharan region.

The rostrum fragment of an ichthyosaur from Northland in New Zealand is described. The specimen appears to belong to the genus Platypterygius. It is the first Mesozoic marine vertebrate fossil reported from Northland and the first cranial element of a Cretaceous ichthyosaur from New Zealand. The fragment is, unfortunately, of uncertain provenance and could have come from the widespread Northland Allochthon (Early Cretaceous–Oligocene) or the allochthonous Houhora Complex (late Early Cretaceous) or Whatuwhiwhi Formation (latest Early to Late Cretaceous), perhaps by way of initial erivation from the Early Miocene Omapere Conglomerate.

Since chimpanzee and bonobo ancestors were evolutionarily closer to the Homo/Pan last common ancestor than extant members of Pan, they probably had more humanlike traits than extant chimps or bonobos. This implies that the presence of humanlike traits in australopithecines (such as a vertical spine, low ilia, smaller canines, thick enamel etc.) does not necessarily indicate human ancestry.

Most paleo-anthropologists conventionally assume that the australopithecines were ancestors of humans (but not of Pan or Gorilla) because they were bipedal. Different lines of evidence (e.g. centrally-placed spines in hominoids, versus dorsally-placed spines in monkeys), however, suggest that not only humans, gibbons and siamangs (who still have habitually vertical spines) had more bipedal ancestors, but all hominoids. This implies that the presence of "bipedal traits" in fossil hominoids does not define human ancestry.

There are a lot of fossil apes in Europe (dryopithecines) as well as in Asia (fossil pongids), but it is traditionally assumed that virtually all fossil hominid fossils in Africa are closer relatives of humans than of gorillas, chimps or bonobos. This is statistically impossible: why should there be plenty of fossil apes in Europe as well as in Asia, but virtually no fossil apes in Africa? Why are all hominid fossils in Africa traditionally assumed to be closer relatives of Homo than of Pan or Gorilla?

This presentation shows that the traditional views that australopithecines are human relatives to the exclusion of Pan and Gorilla, and that human ancestors became bipedal when they moved from the forest to the open plane, are exclusively based on anthropocentric preassumptions, and are contradicted by the comparative anatomy and all other available evidence. We propose alternative hypotheses, fully supported by comparative biology.

Diprotodon Owen, was one of the first fossil mammals described from Australia and has the distinction of being the largest ever marsupial. However, until recently its taxonomy was unclear and knowledge of its continental distribution, palaeoecology and evolution was poorly known. This paper describes cranial elements from an unusually small-bodied Diprotodon from the Early Pleistocene Nelson Bay Formation, Portland, Australia. It is intermediate in size between the smaller-bodied Pliocene ancestor, Euryzygoma dunense de Vis 1895 and the larger-bodied Late Pleistocene D. optatum Owen, . However, it is morphologically inseparable from Late Pleistocene Diprotodon and is here referred to as D. ?optatum. A temporal morphocline most parsimoniously explains the medium body size of the Nelson Bay taxon; thus we provide the first evidence of a transitional form within Diprotodon. The cause(s) of gigantism of the E. dunense-D. ?optatum-D. optatum lineage through the late Cainozoic is unclear but most likely involved the physiological response to changes in the physical and/or biotic environment. Although the size changes within the lineage may have been initially advantageous, they were most likely disadvantageous when the optimum body size was obtained. Our results suggest that Diprotodon was at its largest ever body size during the Late Pleistocene - the time period that coincided with its extinction.

Course name: Introduction to the Paleontology and Paleoanthropology of Spain

Saint Louis University, Madrid Campus, Faculty: Science and Engineering
Semester:  Spring 2005.

Course description: This course gives an introduction into the Paleontology and Paleoanthropology of Spain. Basic principles and methods of Paleontology and Paleoanthropology will be outlined. Especially presented will be the origins of life and human beings, evolution, and extinctions. The paleontological part will concentrate on some interesting Spanish case studies (reefs of the Devonian period, swamp forests of the Carboniferous period, shallow sea of the Jurassic period, dinosaurs from the Upper Jurassic and Lower Cretaceous, and mammals and insects of the Tertiary period). In the center of the paleoanthropological part will be the famous findings of Homo antecessor and Homo heidelbergensis in the caves of Atapuerca and their importance for our knowledge about Human evolution.

Course Calendar:
Monday, Jan. 17: 1. Introduction – What are Fossils?
Wednesday, Jan. 19: 2. The Geologic Time Scale
Monday, Jan. 24, & Wednesday, Jan. 26: 3. Evolution and Taxonomy – Biology and Paleontology in Dialogue (I)
Thursday, Jan. 27: End of DROP/ADD period
Monday, Jan. 31, & Wednesday, Feb. 2: 3. Evolution and Taxonomy – Biology and Paleontology in Dialogue (II) & 4. Biogeography and Mass Extinctions (I)
Monday, Feb. 7: 4. Biogeography and Mass Extinctions (II)
Wednesday, Feb. 9: 5. From the Origins of Life to the Silurian
Monday, Feb. 14: Review & Preparation of first mid-term exam
Monday, Feb. 14: Deadline for the submission of the title of the paper
Wednesday, Feb. 16: First mid-term exam.
Monday, Feb. 21: 6. Reefs of the Devonian Period
Wednesday, Feb. 23: 7. Swamp Forests of the Carboniferous Period (I)
Monday, Feb. 28: 7. Swamp Forests of the Carboniferous Period (II) & 8. Reptiles of the Triassic Period
Wednesday, Mar. 2: 9. Life of the Shallow Sea of the Jurassic Period (I)
Saturday, Mar. 5, to Sunday, Mar. 13: SPRING BREAK
Monday, Mar. 14: 9. Life of the Shallow Sea of the Jurassic Period (II)
Wednesday, Mar. 16: 10. Dinosaurs from the Upper Jurassic and Cretaceous (I)
Monday, Mar. 21: 10. Dinosaurs from the Upper Jurassic and Cretaceous (II)
Wednesday, Mar. 23: 11. Fossils of the Tertiary Period
Thursday, Mar. 24, to Sunday, Mar. 27: Easter
Monday, Mar. 28: Review & Preparation of second mid-term exam
Wednesday, Mar. 30: Second mid-term exam
Monday, Apr. 4: Deadline for the submission of papers and PowerPoint presentations
Monday, Apr. 4, & Wednesday, Apr. 6: 12. Outline of Human evolution.
Friday, Apr. 8: Last day to drop a class and receive a grade of "W"
Monday, Apr. 11, & Wednesday, Apr. 13: 13. The Earliest Man in Spain – the Homo from Orce. & 14. Homo antecessor and Homo heidelbergensis from the Caves of Atapuerca (Prov. Burgos) (I)
Saturday, Apr. 16: Field trip to Guadalajara
Monday, Apr. 18: 14. Homo antecessor and Homo heidelbergensis from the Caves of Atapuerca (Prov. Burgos) (II)
Wednesday, Apr. 20: 15. The Humans after Homo heidelbergensis in Spain.
Monday, Apr. 25, & Wednesday, Apr. 27: Presentation of papers

Course name: Introduction to the Paleontology and Paleoanthropology of Spain

Saint Louis University, Madrid Campus, Faculty: Science and Engineering
Semester:  Spring 2005.

Course description: This course gives an introduction into the Paleontology and Paleoanthropology of Spain. Basic principles and methods of Paleontology and Paleoanthropology will be outlined. Especially presented will be the origins of life and human beings, evolution, and extinctions. The paleontological part will concentrate on some interesting Spanish case studies (reefs of the Devonian period, swamp forests of the Carboniferous period, shallow sea of the Jurassic period, dinosaurs from the Upper Jurassic and Lower Cretaceous, and mammals and insects of the Tertiary period). In the center of the paleoanthropological part will be the famous findings of Homo antecessor and Homo heidelbergensis in the caves of Atapuerca and their importance for our knowledge about Human evolution.

Course Calendar:
Monday, Jan. 17: 1. Introduction – What are Fossils?
Wednesday, Jan. 19: 2. The Geologic Time Scale
Monday, Jan. 24, & Wednesday, Jan. 26: 3. Evolution and Taxonomy – Biology and Paleontology in Dialogue (I)
Thursday, Jan. 27: End of DROP/ADD period
Monday, Jan. 31, & Wednesday, Feb. 2: 3. Evolution and Taxonomy – Biology and Paleontology in Dialogue (II) & 4. Biogeography and Mass Extinctions (I)
Monday, Feb. 7: 4. Biogeography and Mass Extinctions (II)
Wednesday, Feb. 9: 5. From the Origins of Life to the Silurian
Monday, Feb. 14: Review & Preparation of first mid-term exam
Monday, Feb. 14: Deadline for the submission of the title of the paper
Wednesday, Feb. 16: First mid-term exam.
Monday, Feb. 21: 6. Reefs of the Devonian Period
Wednesday, Feb. 23: 7. Swamp Forests of the Carboniferous Period (I)
Monday, Feb. 28: 7. Swamp Forests of the Carboniferous Period (II) & 8. Reptiles of the Triassic Period
Wednesday, Mar. 2: 9. Life of the Shallow Sea of the Jurassic Period (I)
Saturday, Mar. 5, to Sunday, Mar. 13: SPRING BREAK
Monday, Mar. 14: 9. Life of the Shallow Sea of the Jurassic Period (II)
Wednesday, Mar. 16: 10. Dinosaurs from the Upper Jurassic and Cretaceous (I)
Monday, Mar. 21: 10. Dinosaurs from the Upper Jurassic and Cretaceous (II)
Wednesday, Mar. 23: 11. Fossils of the Tertiary Period
Thursday, Mar. 24, to Sunday, Mar. 27: Easter
Monday, Mar. 28: Review & Preparation of second mid-term exam
Wednesday, Mar. 30: Second mid-term exam
Monday, Apr. 4: Deadline for the submission of papers and PowerPoint presentations
Monday, Apr. 4, & Wednesday, Apr. 6: 12. Outline of Human evolution.
Friday, Apr. 8: Last day to drop a class and receive a grade of "W"
Monday, Apr. 11, & Wednesday, Apr. 13: 13. The Earliest Man in Spain – the Homo from Orce. & 14. Homo antecessor and Homo heidelbergensis from the Caves of Atapuerca (Prov. Burgos) (I)
Saturday, Apr. 16: Field trip to Guadalajara
Monday, Apr. 18: 14. Homo antecessor and Homo heidelbergensis from the Caves of Atapuerca (Prov. Burgos) (II)
Wednesday, Apr. 20: 15. The Humans after Homo heidelbergensis in Spain.
Monday, Apr. 25, & Wednesday, Apr. 27: Presentation of papers

The modern Australian koala (Phascolarctos cinereus) is commonly regarded as a dwarf descendent of a Late Pleistocene giant koala (Ph. stirtoni). The implication of that hypothesis is that the giant koala survived the Late Pleistocene megafaunal extinction ‘‘event’’, albeit as a smaller body-sized form. It is important to be able to constrain rates of Late Pleistocene faunal turnover, an aspect reliant on having accurate taxonomic information of extinct species. The koala dwarfing hypothesis is tested here by using a temporally-constrained biogeographical record of fossil koalas, and a morphological character analysis. The contemporary occurrence of both taxa in pre-Late Pleistocene deposits and significant differences in dental morphologies between those forms suggests that the modern koala is not a derived dwarf of the Pleistocene giant koala. Thus, the giant-form was among a number of other giant mammals, lizards and birds that suffered extinction sometime during the Late Pleistocene. The potential phenomenon of dwarfing of other Late Pleistocene and Recent faunas, such as grey kangaroos, is commonly used as a test for or against various megafaunal extinction hypotheses. However, the results of this study also demonstrate that the dwarfing hypothesis has not been adequately tested for a suite of other taxa. Thus, until the dwarfing hypothesis can be more fully tested, a clear understanding of the fate of Late Pleistocene faunas that apparently survived the extinction ‘‘event’’, and the origins of many extant forms will remain elusive.

Leopard remains are rare in the European fossil record, probably a consequence of its solitary and elusive habits. Equi, dating back to Marine Isotope Stage 3 (MIS3), represents a rich and outstanding exception. Historical excavations (from 1911 to about 1917) at Equi allowed  more than 200 remains to be recovered of this large felid, a sample attesting the richest leopard-bearing site in the Pleistocene of Europe. The Equi site testifies the survival of a prosperous leopard population in northwestern Italy (the Apuan Alps area) during the pre Last Glacial Maximum Late Pleistocene. Almost all skeletal elements are represented in the collection (except ribs, and most of axial skeleton). At Equi, leopard is represented by both young and adult animals  and, up to now, it seems the first and unique Pleistocene evidence of cave usage for cubs raised in Europe.
This study focuses on the whole record of leopard from Equi, housed in the Museum of Natural History of Florence, and is aimed to analyze the taphonomical assemblage, to provide an anatomical description of the remains and the comparison to the European evidences. Moreover, we infer the paleoecology of leopards, and their relationships with other large carnivores from the Equi guild (Ursus spelaeus, Canis lupus, and Panthera leo spelaea). The site was probably an occasional large carnivores den, used over time by different large carnivore taxa, perhaps in different seasons or times (mutual avoiding).
Our morphological analysis allows us to recognize sexual dimorphic characters in the cranium, and a large morphological variability in body size.

Although in the past fifty years Aguascalientes has been characterized by important paleontological discoveries, is currently observed a marked deterioration in the sites of
fossiliferous outcrops that occur naturally, due to rapid urban growth, the lack of information and lack of given value of this important heritage of our state by the government officials and lay people.
This document is intended to raise awareness of the paleontological richness that exists in our state and in this sense is also a starting point for further research. In its
development, the authors consulted various sources on the subject and conducted paleontological activities both in field and laboratory.

Pliosaurier gehören zu den größten Meeresreptilien, die je gelebt haben. Vom Mittleren Jura bis in die obere Unterkreide erreichten einige Gattungen gigantische Ausmaße und standen meist an der Spitze der Nahrungskette. In den letzten Jahren haben besonders zwei Pliosaurier-Funde das Interesse der Öffentlichkeit erregt. Einer davon ist als „Monster von Aramberri“ bekannt geworden. Es handelt sich dabei um ein unvollständiges Skelett aus dem Kimmeridgium von Mexiko, dessen fossile Überreste von Buchy et al. (2003) als nicht näher bestimmbarer Pliosauride beschrieben wurden. Das Tier erreichte vermutlich eine Gesamtlänge von 15 m. Der zweite populär gewordene Fund, gleichfalls ein unvollständiges Skelett, stammt aus dem Tithonium von Svalbard (Spitzbergen) und erhielt den Spitznamen „Predator X“. Das Exemplar wurde von Knutsen et al. (2012) als neue Art der Gattung Pliosaurus, Pliosaurus funkei beschrieben und erreichte vermutlich eine Länge von zwischen 10 und 13 m. Keiner der uns bekannten Funde von Pliosauriern mit Längen von mehr als 10 m ist vollständig. Meist werden lediglich fragmentarische Reste von Skeletten entdeckt, die oft auch eher schlecht erhalten sind.
Keywords: Pliosauridae, Liopleurodon, Pliosaurus giganteus, Thaumatosaurus, Jurassic, Germany

Abrupt collapse of the tropical rainforest biome (Coal Forests) drove rapid diversification of Carboniferous tetrapods (amphibians and reptiles) in Euramerica. This finding is based on analysis of global and alpha diversity databases in a precise geologic context. From Visean to Moscovian time, both diversity measures steadily increased, but following rainforest collapse in earliest Kasimovian time (ca. 305 Ma), tetrapod extinction rate peaked, alpha diversity imploded, and endemism developed for the first time. Analysis of ecological diversity shows that rainforest collapse was also accompanied by acquisition of new feeding strategies (predators, herbivores), consistent with tetrapod adaptation to the effects of habitat fragmentation and resource restriction. Effects on amphibians were particularly devastating, while amniotes ('reptiles') fared better, being ecologically adapted to the drier conditions that followed. Our results demonstrate, for the first time, that Coal Forest fragmentation influenced profoundly the ecology and evolution of terrestrial fauna in tropical Euramerica, and illustrate the tight coupling that existed between vegetation, climate, and trophic webs.

Gigantopithecus blacki is the largest hominoid that ever lived. The consensus view is that it is a specialized pongine and late-surviving member of the Sivapithecus-Indopithecus lineage. It is known primarily from Early and Middle Pleistocene cave sites in southern China, dating from 2.0 Ma to almost 300 ka. The cause of its extinction in the late Middle Pleistocene is unknown, but ecological change or the arrival of Homo erectus may have been contributing factors. Gigantopithecus is highly specialized in its dentognathic anatomy, with a unique combination of features that distinguish it from all other hominoids. Based on the size of its dentition and mandible, a reasonable estimate of its body mass would be 200–300 kg. There was a progressive increase in dental size from the Early Pleistocene to the Middle Pleistocene, and possibly a shift towards greater complexity of the cheek teeth. Gigantopithecus exhibits a relatively high degree of sexual dimorphism, implying a high level of male-male competition, but the relatively small canines in both sexes suggest that these teeth were not important in agonistic behaviors. The species inhabited a subtropical monsoon forest with a closed canopy and dense understory. Foraging was focused on the forest floor and its diet included a broad range of C 3 plants, including fruits, leaves and stems, and possibly tubers. The cheek teeth and jaws were adapted for processing a wide variety of bulky, fibrous, and abrasive food items, but the small incisors indicate that incisal preparation was not an important part of its feeding repertoire.

The actual abstract (summary) is 2 printed pages.

This monograph is based on my PhD dissertation and describes microvertebrate faunas from seven localities in the lower Chinle Group of Texas, New Mexico, and Arizona. This includes Trilophosaurus quarry 1 (Colorado City Formation, TX), the lower and upper Kalgary localities (Tecovas Formation, TX), the Ojo Huelos locality (Ojo Huelos, NM), L-2739 (Sixmile Canyon, NM), Krzyzanowski bonebed (St. Johns, AZ), and Dying Grounds (Petrified Forest, Arizona). There's a few new taxa here---Diplolonchidion murryi, Palaeollanosaurus fraseri, Planocephalosaurus lucasi, Crosbysaurus harrisae, and Protecovasaurus lucasi. They are a shark, two sphenodontians, and two archosauriforms that may or may not be convergent with early ornithischians or pterosaurs (subperpendicular, offset denticles). I particularly like Crosbysaurus because it has compound denticles and would appear to represent an intermediate stage between fine denticles (serrations) and coarse denticles typical of both Revueltosaurus and many ornithischian dinosaurs.

The status of current research about the distribution of large game during the Upper Paleolithic in Middle and Western Europe and its history can be only investigated and presented through intensive literary research and contact with colleagues locally and abroad. In spite of the navoidable simplification about the history of this research, this portion of the work was necessary, in order to be able to critically consider the data. The marked difference in the number of
established research positions for the study of the Paleolithic in Spain, Italy, France, and Germany is not only a consequence of differing research systems and research institutions, it reflects the importance given to Paleolithic research in each of the different countries and also permits predictions about future reserches (pp. 5–23).

Likewise, a detailed overview of the topography and the climate of Europe, as well as the environnmental changes that occurred, serve as a basis to critically evaluate the distribution of large game during the Upper Paleolithic in Middle and Western Europe from an ecological point of view (pp. 25–37). The importance of the coastal region and its sea level changes have been mapped and emphasized.
For at least 400 kya, especially after the appearance of wooden lances and throwing spears used for hunting, human beings have been considered hunters. During the Upper Paleolithic, the spear thrower was developed and later was followed by the bow and arrow. Thus, there then existed the ability to kill any animal. Many drawings and engrawings from the Upper Paleolithic show scenes in which almost all
species of animals are pierced by projectiles. In addition almost all species have been depicted on spear throwers. This place the Upper Palaeolithic man at the top of the ecological pyramid (pp. 39–47).

For the first time, many of the published depictions of uncommon animal species are brought together: Equus hydruntinus, Sus scrofa, Dama dama, Capreolus capreolus, Alces alces, Megaloceros giganteus, Ovibos moschatus, Rupicapra rupicapra, Saiga tatarica, Phocidae, Cetacea and Pinguinus impennis.
The analysis of the distribution of large game in Middle and Western Europe, on the basis of faunal remains, as well as artistic depictions, provides for numerous hypotheses (especially 49–145). The distribution of specific species, in comparison to that of other species is discussed, such as the significance of animals in overlapping regions that belong to the same family and, sometimes, to the same genus. These
animals exhibited competition-driven ecological adaptations as seen between the Equus ferus and Equus hydruntinus, Coelodonta antiquitatis and Stephanorhinus kirchbergensis and Stephanorhinus hemitoechus, Mammuthus primigenius and Elephas antiquus, Cervus elaphus and Rangifer tarandus,
Alces alces and Megaloceros giganteus, Bison priscus and Bos primigenius, Capra pyrenaica and Capra ibex and Rupicapra rupicapra.
It is especially significant that during OIS4 (oxygen isotope stage), the first glacial maximum of the last ice age, the distribution of species such as mammoth and woolly rhino extended as for south as southern Spain and southern Italy, without the extinction of the forest elephant, forest rhino, and
steppe rhino, which ware living to these temperate regions. Aside from a few definitive pictures of reindeer, researchers have made only a few ambiguous identifications from the „mammuthus-coelodonta“ faunal complex during the time of the Upper Paleolithic, south of Cantabria. There are few Upper Paleolithic remains that have been discovered in Italy and no representations of Mammuthus primigenius,  coelodonta antiquitatis or Rangifer tarandus are reported.
The extinction of the forest elephant, as well as the forest and steppe rhino, did not occur until shortly after 30 kya. At about the same time, the European bison and the giant deer disappeared from southern Italy. Beginning around this period, and no later than the second glacial maximum (OIS2) of the last ice age, the number of fallow deer declined sharply and even may have completely died out in southwestern Europe. This sugeste a strong impoverishment of the larger game in southern Europe. Though not analyzed
in this study, one could also add Homo neanderthalensis to the list of these extinct animals.

The analysis of the few indications of the larger animals in the coastal tract shows that this region was not only later submersed by the sea (sea level has risen more than 100 m) but also that researchers have often forgetten about them. The few sites with elements from the coast, must only represente a small number of such original sites that currently lie underwater (pp.147–165).

In conclusion, the interpretation of the data shows that a direct relationship exists between the natural appearance of a species and its artistic representation. Therefore, both the presence of the species and their representations are more dependent on environmental components than from any cultural components (pp. 167–175).
Finally, this hypothesis would be formulated as follows, that the destiny of Homo sapiens and Homo neanderthalensis, as well as other linked species, is strongly influenced by factors such as climatic and ecological changes during the period between 40 and 25 kya.