Quaternary palaeontology

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.

More than 800 isolated teeth of fossil Pongo have been recovered from cave sites in the vicinity of Chongzuo in Guangxi, southern China, ranging from the Early to Late Pleistocene (2.0e0.1 Ma). These collections provide a unique regional window into the evolutionary history of orangutans over a twomillion-year period at the northernmost extent of their former geographic range. Here we investigate the nature and timing of the evolutionary change in the dental size of fossil orangutans from Chongzuo. Fossil tooth size (mesiodistal length*buccolingual breadth) was compared against an extant Pongo pygmaeus standard (n ¼ 106 individuals). During the course of the Pleistocene, orangutans from southern China exhibited a progressive reduction in overall dental size. Early Pleistocene Pongo has cheek teeth with occlusal areas that are 38.1% larger than those of extant P. pygmaeus. Those from the Middle and Late Pleistocene are 25.2% and 18.9% larger, respectively. Previously, the size difference in dentition between the Early to Middle Pleistocene and Middle to Late Pleistocene samples was used to differentiate time-successive species of Pongo, namely Pongo weidenreichi and Pongo devosi. However, with access to larger samples and better representation of populations through time, the evidence in support of this taxonomic arrangement requires reconsideration. Diminution of the teeth now appears to be a gradual evolutionary transformation rather than a punctuated event. Moreover, the morphological features that distinguish the Chongzuo fossil orangutans from extant Pongo spp. remain uniform throughout the Pleistocene. Retaining P. weidenreichi and P. devosi as anagenetic species remains an option, but, given the current evidence, we consider it preferable to assign all of the fossil orangutans from Chongzuo to P. weidenreichi. Beyond resolving questions of alpha taxonomy, the study of fossil orangutan dental size provides a basis for estimating body mass, which has implications for interpreting the paleobiology of Pleistocene Pongo in southern China.

ABSTRACT

Geographic and Geological setting

The cave of Eirós is situated at the East of Galicia, on the north face of the Serra do Ouribio, in the foothills of the eastern ranges of Galicia (see page 44).

The entrance to the cave is halfway up a steep slope on the North side of the mount Costa do Val. The topography of the cave is markedly longitudinal, following a NNW direction. It is a small karstic cave 100 m long. It has an altitude of 780 m above sea level, and 25 m above the present level of a small river, Bezcas (see page 45).

It opens out in the limestone of Cándana, of the Lower Cambrian age. This is grey limestone with parallel lamination, with intercalations of green slates. At this point they reach a thickness of 70 m.

The Cova Eirós deposit

At the Cova Eirós deposit of Ursus spelaeus, there is not evidence to suggest that the type of dental and bone remains discovered has been due in any way to anthropical action or to that of other animals (scavengers). Neither has it been affected by any geological process. These conclusions are based on the following:

- In the material studied, no one part of the skeleton is found to be more abundant than any other (see page 59).
- The well-preserved state of the remains, their position within the sediment and the absence of any size-distribution indicate that the dinamic of the flow which acted upon the deposit was not very great (see pages 57, 58).

The fact that the bones are not always connected anatomically can be explained by displacement on a small scale as well as the movement caused by the bears themselves during their long occupation of the cave.

The Ursus spelaeus population of Cova Eirós

In the bones and dental remains studied, we observe a slight predominance of females over males. The age range is similar to that found at other Cave Bear deposits, and indicates an extremely high incidence of young and neonate mortality (70%) and the presence of a large percentage of senile specimens, recognizable by the extensive erosion of their teeth (see pages 60, 61).

We have not found significant differences to a taxonomic level between the bone and dental morphologies of the Cova Eirós Ursus spelaeus and those of European populations described by other authors. In any case, intrapopulational variability is high, as is usual with this species, which makes it difficult to interpret the polymorphism observed here in interpopulational terms.

We have been able to separate the variability caused by size differences (due principally to sexual dimorphism) from that resulting from morphological differences using the Principal Component Analysis. This allows us to determine the sex of the bone and dental remains, when the size of the sample is significant. The first principal component would explain the variability produced by sexual dimorphism, whilst the remaining principal components would explain purely morphological differences.

Metric characterization of the Cave Bear population of the Cova Eirós has enabled us to quantify the sexual dimorphism in the various remains studied.

As far as the skeleton is concerned, the maximum dimensions of each bone are not only greater in males than in females, but the latter also indicate a more delicate skeleton, mainly reflected in a lesser development in the bone superestructures of the cranium and in he dimensions of the proximal and distal epiphyses of the limb bones, also proportionally smaller.
In the case of dental specimens, the sex difference is clearly ascertained from the canines and (though less evidently) from the molariforms. In the latter case, the sexual dimorphism is hardly discernible using basic statistic techniques, but becomes clearer when a principal component analysis is carried out. Not all dental remains reflect the intersexual difference to the same degree.

On the whole, the first principal components are general components, and deal with the size of the teeth and, finally, with sexual dimorphism. The second principal components seem to explain the shape of the teeth, that we call oclusal line. Finally, the third principal components are related to the development of the cusps and their higher or lower degree of convergence (see page 237).

The lower third molar tends to display a high morphometric variability, even if a large percentage of the variance present (78.2%) is explained by the first principal component. The upper and lower fourth premolars show high percentages of variance produced by sexual dimorphism (71.4% and 63.5% respectively), followed by the upper first molar (58.3%), the lower second molar (41.3%), the upper second molar (39.2%) and the lower first molar (35.6%).

In the three last items, the high percentage of variance present is explained by the following principal components: In the upper second molar, the PC II (related to the morphology of the heel) explains a very high percentage of variance (21.6%), which indicates that it is that area of the tooth that is mainly responsible for the existing high variability, almost equalling the differences produced by sexual dimorphism.

In the lower second molar, the variance explained by the first three principal components is only 64.6%. Its variability will be explained, therefore, by the other principal components, which deal with very specific parts of the tooth topography.
This same phenomenon occurs, to a main degree, in the lower first molar, given that the first three principal components explain only 56.3% of the total variance.

Population comparison

A comparative study of the dental morphologies of the six populations studied using Cluster analysis displays a variable tendency in each tooth. However, an analysis grouping together all the morphological data of all the molariforms from the population of Eirós, Odessa, Troskaeta, Ekain, El Toll y Arrikrutz, relates the various populations according to their geographical situations. If, in the Cluster analysis, we include the populations of El Reguerillo (Madrid) and Cueva Mayor (Burgos), the latter is Ursus deningeri, this process becomes even more apparent. All the peninsular populations of Ursus spelaeus are grouped together, where Eirós is associated with Ekain and El Toll with El Reguerillo. The peninsular group is connected to Ursus deningeri of Cueva Mayor, and then, finally, with Ursus spelaeus of Odessa (see page 210).

The sexual dimorphism which exist in Ursus spelaeus would appear to recommend a sexing of the material before carrying out any metric interpopulational study. However, it is not always possible to do the sexing.

This paper demonstrates that  the problem can be avoided by carrying out a principal component analysis and using the data obtained, except the first principal component, for the interpopulational study. Consequently, the variability produced by the differences in size between sexes disappears and the study becomes coherent.

The interpopulational variability which exist in those populations studied manifests itself to a varying degree in each molariform. The results of the variance and discriminant analyses indicate the existence of a very high interpopulational difference in the lower first molar (with a Wilks l value of 0.042), followed by the upper first molar (0.295), lower second molar (0.358) and upper second molar (0.420) whilst in the upper fourth premolar and above all in the lower one, the capacity to differenciate between populations is lower (0.583 and 0.830 respectively) (see page 238).

The overall interpretation of the principal component analysis and the discriminant analysis indicates that  certain dental items (the lower second molar, the upper first molar and principally the lower carnassial) display a high variability produced by specific morphological differences (such as the development of each cusp) and not only by sexual dimorphism. These differences, given the high percentages of correct reclassifications obtained, allow us to distinguish between the populations studied.

Other items, on the other hand, owe most of their variability to interpopulational differences, mainly sexual dimorphism. This is the case of the upper and lower fourth premolars, where an interpopulational study using a discriminant analysis does not show up a clear morphological difference between the populations studied (see page 239).

When studying an extinct species such as the cave bear (Ursus spelaeus ROSENMÜLLER 1794), it is possible to apply a variety of molecular biology techniques such as the study of stable isotopes or mitochondrial DNA (mDNA) to infer patterns of behaviour or physiology that would otherwise remain concealed. Throughout Europe and along time, differences in the isotopic values (δ 13 C and δ 15 N) of cave bears arise from environmental differences and the Pleistocene climatic evolution. The climate determines the hibernation length, during which the cave bears undergo a particular physiology that can be related to an increase in δ 15 N during climate cooling. In order to verify whether hibernation affected the isotopic values, we compared cave bears in different ontogenetic stages. The results show that perinatal values reflect the values for mothers during hibernation, while juveniles show differences in maternal investment. A previous study in the literature based on complete mitochondrial DNA sequences of several individuals collected from closely situated caves showed that each cave housed, almost exclusively, a single lineage of haplotypes. This pattern suggests extreme fidelity to the birth site, or homing behaviour, and that cave bears formed stable maternal social groups, at least for the purpose of hibernation. Studies of this type offer unexpected data on the palaeobiology of this extinct animal.

The mollusc  fauna  of  the  Brabant  Loam  (Liempde Deposit,  Formation  of  Boxtel)  is  described on  the basis of a  review of  research of  the  last one hundred years. Although much research was undertaken, hardly anything of it can be traced in publications. This contribution  tries  to  fill  this gap  in our knowledge. Because  the very varying methods of  research, results are difficult to compare with each other and therefore only a review has been made with presence or absence of taxa. For the same reason, conclusions are only possible to a limited degree. Only for one location, an outcrop near Acht, do the data lend themselves for a graphical representation and an analysis of the palaeoecological developments in the section.
The associations point here to a becoming wetter with a subsequent becoming dried up of a shallow pool in an open landscape with a vegetation of grasses and herbs. Seventeen locations are know  from the Brabant Loam  from which 35 taxa are known, 29 of which could be identified to species level. The most common species are Succinea schumacheri, Anisus leucostomus auct., Succinea oblonga s.l., Pupilla muscorum, Vertigo genesii, Gyraulus rossmaessleri and Columella  columella. All associations  found  indicate moist  to wet biotopes  in an open landscape during a subarctic climate. The average number of taxa per  location  is 10 which is a normal number for mollusc associations indicative of cold non-marine conditions. The mollusc species present give no indication of a specific stratigraphical age other than that it concerns a cold period. Earlier published data show that the Brabant Loam dates from a late phase of Marine Isotope Stage 3.

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.

Palinološka istraaivanja kvartarnih sedimenata u Petnièkoj peaeini (nastavak istraaivanja) Tokom 2014. godine zapoèeta su palinološka istraaivanja kvartarnih sedimenata u Petnièkoj peaeini. Pošto kvalitet analize polena zavisi od velièine ispitivanih uzoraka, ovaj rad pred-stavlja nastavak palinoloških istraaivanja ple-istocenskog sloja u Petnièkoj peaeini radi što preciznije rekonstrukcije paleoekoloških uslova. Uzorci su uzeti iz istog sloja na rastojanju od 10 cm od mesta prošlogodišnjeg uzorkovanja, dok je ekstrakcija polena vršena na isti naèin kao tokom prošlogodišnjih istraaivanja. Analizom je konstantovano 12 zrna polena. Ustanovljen je manji broj familija (sedam), u odnosu na pro-šlogodišnje istraaivanje (trinaest). Identifiko-vana su po tri polenova zrna rodova Solanum (Solanaceae) i Schoenoplectus (Cyperaceae), dva iz familije Primulaceae i po jedno zrno rodova Andropogon (Poaceae), Alnus (Betula-ceae) i Quercus (Fagaceae) i familije Astera-ceae.

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.

The southern part of the North Sea is well known for its rich fossil-bearing localities. Abundant Pleistocene mammal remains are retrieved by fishing and during large-scale sandextraction projects. Archaeological artefacts are found regularly as well, albeit in lower quantities. Little is known about the geological background and stratigraphical context of these finds and that hampers the reconstruction of the natural palaeoenvironment as well as the accurate determination of the associated age. In general the older units, located deeper under the seabed, are sampled only by destructive coring techniques, preserving little of the
geological and stratigraphical context recorded in the sedimentary succession.
Here we present preliminary results of a unique multidisciplinary study in and around the offshore sand-extraction pit created for the extension of the Maasvlakte harbour area southwest of Rotterdam. A series of minimally disturbed vibrocores collected perpendicular to the slope of this 20-m-deep (300 million m3) pit shows a main stratigraphical sequence of fluvial Rhine-Meuse deposits dating back to the Middle Pleistocene, preserved between 25 and 46 m below NAP. As part of on-going work, phases of reworking resulting in mixing and concentration of fossils and artefacts are identified by analysing the coarse residue of core samples. Ages of individual units are constrained by a variety of laboratory analyses, including OSL dating. Acoustic data are used to show the morphology of the extraction pit, coarse clasts on the seabed and details of the sediment architecture. The results are linked to the well-know stratigraphy of the adjacent onshore area, to the result of ‘bone’ trawling surveys, and to analyses of the dumped sediment at the Maasvlakte extension. All data will be used to reconstruct the Middle- to Late-Pleistocene palaeolandscapes that characterized this part of the Southern Bight.

The lower levels of the archaeological sequence of Kozarnika cave (north-western Bulgaria) provide levels with non-Acheulian core-and-flake (as opposed to pebble-core) industries. Associations with numerous taxa of large mammals indicate that the date of the lower layers of the cave falls between MNQ17 and MNQ19. These layers produced several bones showing anthropic traces. Its date and geographical position place Kozarnika cave as a landmark site in the primary phase of the settlement of Europe.

La fin du Pliocène et le début du Pléistocène représentent pour l’Europe occidentale et centrale une période charnière au cours de laquelle le climat se modifie, passant d’un stade chaud et stable à un stade beaucoup plus instable et froid. Ce changement a impliqué de profondes modifications dans la composition de la faune de grands mammifères européens. Depuis longtemps on a supposé que toute apparition d’une nouvelle espèce se faisait par migration venant d’Asie. Un recensement quasi exhaustif des sites à faunes de grands mammifères ainsi qu’un calage précis de ceux-ci dans l’espace et dans le temps a permis d’identifier les grands courants de migration successifs mais aussi de proposer reconstitution hypothétique relativement précise des différents milieux écologiques.
Mémoire de maîtrise sous la direction d'Armel Coudé. Jury: Jean-Pierre Lautridou. Expertise: Marie-Françoise Bonifay.
The late Pliocene and early Pleistocene represent for Western and Central Europe, a pivotal period during which the climate changes, going from a warm and stable stage to a much more unstable and cold stage. This change has involved major changes in the composition of the fauna of large European mammals. For a long time it was assumed that any appearance of a new species was by migration from Asia. An almost exhaustive inventory of sites with large mammals and precise timing of these in space and in time identified the major successive migration flows but also provided relatively accurate hypothetical reconstruction of the different ecological environments.
Master's thesis under the direction of Armel Coudé. Jury: Jean-Pierre Lautridou. Expertise: Marie-Francoise Bonifay.

Este artículo tiene por objetivo el análisis de la distribución biogeográfica de felinos
(Felis, Lynx y Panthera) en la Península Ibérica durante el Pleistoceno Superior. Este periodo
comprende fases climáticas rigurosas, última fase glacial, y templadas. Estas cambiantes condiciones
afectaron en la distribución geográfica de la fauna, propiciando la presencia de especies
adaptadas a medios fríos en el tercio norte. La Península Ibérica es el territorio europeo situado
más al Suroeste, su situación hace que en este territorio existan diversas áreas climáticas, esto
unido a un complejo relieve afectaron en la distribución biogeográfica de las especies durante el
Pleistoceno Superior.
En la Península Ibérica han sido identificados tres géneros de felino, desde un pequeño carnívoro
como el gato montés, al lince de tamaño medio, hasta grandes carnívoros como el leopardo
y el león. El análisis de esta familia de carnívoros permite realizar un repaso diacrónico y geográfico
a las adaptaciones de los carnívoros al cambiante medio ambiente del Pleistoceno Superior.
Sin embargo, esta investigación ha de considerarse como un trabajo preliminar, futuros trabajos
podrían modificar la actual distribución biogeográfica, especialmente en aquellas regiones en las
que ciertas especies están ausentes.

A diverse Late Pleistocene fossil assemblage was recovered from a sea cliff locality near Gopnath in Gujarat, northwestern India. These remains are the first large sample of Pleistocene faunal materials from arid northwestern India. Several taxa known primarily from coarse alluvial deposits of central India are
documented for the first time from an undisturbed open-air site adjoining the Great Indian Desert. The sample includes a new species of antelope from a lineage considered extinct outside of Africa since the Early Pleistocene. The paleoenvironmental context, faunal composition and type of fossil preservation
reported here is unique. The Gopnath fauna accumulated in a pond within a carbonate dune field that formed part of a larger coastal oasis ecosystem. This paleoscape occupied the Cambay Gulf during hyperacid glacial low stands. The Gopnath fossils are correlated to Late Acheulean lithics from a coastal cliff locality (<8 km) at Madhuban. These finds provide the first vertebrate evidence of glacial low stands and their influence on Late Pleistocene paleobiogeography within the dynamic dry coastal corridor linking India to Africa. They offer a rare glimpse of a lost landscape and an obscure fossil community that are critical to understanding the paleobiogeography of the hinterland along the Arabian seashore and informing models of early human dispersals.

Abstract: This paper presents a peer-reviewed and updated/corrected list oif >100 publicly known late Quaternary proboscidean sites that have certain or possible traces of hominin utilization in Africa, Europe, and Asia, along with a sample of references, chronometric or estimated ages, and brief descriptions of the associated materials and bone modifications. Summary discussions of important sites are also presented. Lower Palaeolithic/Early Stone Age hominins created far fewer proboscidean site assemblages than hominins in later Palaeolithic phases, in spite of the time span being many times longer. Middle Palaeolithic/Middle Stone Age hominins created assemblages at eight times the earlier hominin rate. Upper Palaeolithic/Later Stone Age hominins created site assemblages at >90 times the rate of Lower Palaeolithic hominins. Palaeoloxodon spp. occur in nearly one third of the sites with an identified or probable proboscidean taxon and Mammuthus species are in nearly one half of the sites with identified or probable taxon. Other identified proboscidean genera, such as Elephas, Loxodonta, and Stegodon, occur in few sites. The sites show variability in the intensity of carcass utilization, the quantity of lithics bedded with bones, the extent of bone surface modifications, such as cut marks, the diversity of associated fauna, and mortality profiles.

A small but significant assemblage of Late Pleistocene mammals was recovered from an eroding shoreline at Paw Paw Cove, located on the Chesapeake Bay side of Tilghman Island, Talbot County, Maryland. Additionally, Clovis-age (11,050–10,800 radiocarbon [14C] years before present) artifacts were found in a lag deposit beneath a loess deposit at the site. An accelerator mass spectrometry date obtained from an organic stratum below the loess at the same location suggests that this small fossil assemblage is approximately 21,000 years old. Identifiable taxa include four extinct species, Castoroides sp. (giant beaver), Equus sp. (horse), Tapirus veroensis (Vero tapir), and Mammut americanum (American mastodon), along with Canis cf. C. latrans (coyote), and Cervus elaphus (wapiti). Significantly, the tapir and giant beaver are the first records for Maryland, and the tapir is the first record from the Coastal Plain of the Chesapeake region of Delaware, Maryland, and Virginia. The provisional identification of Canis cf. C. latrans may represent the first Late Pleistocene record from the Coastal Plain of the Chesapeake region of Delaware, Maryland, and Virginia.  See:  https://scholarlypress.si.edu/store/new-releases/late-pleistocene-rancholabrean-mammalian-assemblag/.

Non-marine molluscan faunas from sites of late Lower and early Middle Pleistocene age in Britain, The Netherlands, France, Germany and Denmark are reviewed. The fauna from the Cromerian stratotype at West Runton shares two extinct prosobranchs (Valvata goldfussiana and Tanousia runtoniana) with that of the Bavel Interglacial at its type locality. Conversely, two other prosobranchs, Lithoglyphus jahni and Fagotia wuesti, characteristic of the Bavelian and earlier periods, are absent at West Runton. These facts suggest that the two sites are close in age but that West Runton is the younger. Moreover, the stratigraphical range of V. goldfussiana in the Netherlands (Middle Tiglian - Cromerian Interglacial I) suggests that West Runton should be placed early in the 'Cromerian Complex'.
British sites assigned to the 'Cromerian' Stage fall loosely into three groups. Group I is characterized by V. goldfussiana, T. runtoniana, Bithynia troscheli and Viviparus viviparus gibbus, and includes West Runton, Kessingland and Sugworth. The fauna of Group 2 with Valvata naticina, B. troscheli, Tanousia cf. stenostoma and Belgrandia marginata is known only from Little Oakley. Group 3 faunas, such as those from Sidestrand and Trimingham, with V. naticina, Bithynia tentaculata and B. marginata, lack distinctive 'Cromerian' elements and have close affinities with certain Hoxnian assemblages. The extinct water vole Mimomys savini has been recorded from sites assigned to each of these three Groups. At Ostend, Norfolk, an undiagnostic molluscan fauna has been recovered from beds that have yielded the vole Arvicola t. cantiana. It would appear from the malacological evidence that two, or more likely several, interglacials, currently assigned to the British Cromerian, are represented.
It is not possible to assign with confidence any of the continental sites discussed to these groups but the faunas from Noordbergum and Exmorra have closest affinities with Group 3, whereas those from IJsselstein and from oNrth Sea borehole E8-4, both with V. goldfussiana, are closest to Group I. The malacological data do not support correlation of Noordbergum ('Interglacial IV') with West Runton but would support a recent suggestion, based on vertebrate evidence, that West Runton is older. It is not possible to correlate West Runton with any known parts of the four Dutch 'Cromerian' interglacials.

In an unpublished 1899 handwritten letter addressed by Agustin de Hornedo to González de Linares, the sender informs about a trip to
and describes the selected paleontological materials at the site. Other documents, in 1903, are archaeological notes written by González de
Linares about painted caves discovered that year in Cantabria and the first photographs of Covalanas and El Haza. Decontextualized phrases
in Les cavernes de la région cantabrique (Monaco, 1911) caused the misunderstanding that led several authors to think of González de Linares as the discoverer of the group of caves in the Miera valley: Puntida, Salitre and Sapo. According to the known documentation, the discovery  of La Puntida must be attributed to the engineer Agustin de Hornedo, and Salitre to professor Lorenzo Sierra.