Cold Fauna Research Articles

Stable isotopes confirm the Banwell Bone Cave Mammal Assemblage Zone represents an MIS 5 fauna

Abstract The position of the Banwell Bone Cave mammal assemblage zone (MAZ) in the mammalian biostratigraphy of the British Isles has been the focus of debate for decades. Dominated by fauna typical of cold environments it was originally linked to the marine oxygen isotope stage (MIS) 4 stadial (ca. 72–59 ka). Subsequently it was argued that the Banwell Bone Cave MAZ more likely relates to the temperate interstadial of MIS 5a (ca. 86–72 ka). It is envisioned that “cold fauna” such as bison and reindeer moved into Britain during stadial MIS 5b (ca. 90 ka) and were subsequently isolated by the rising sea level during MIS 5a. Here we investigate environmental conditions during the Banwell Bone Cave MAZ using bone collagen δ13C and δ15N and tooth enamel δ18O and δ13C isotope analysis. We analyse bison and reindeer from the MAZ type-site, Banwell Bone Cave. Our results show unusually high δ15N values, which we ascribe to arid conditions within a temperate environment. Palaeotemperature estimates derived from enamel δ18O indicate warm temperatures, similar to present day. These results confirm that the Banwell Bone Cave MAZ relates to a temperate interstadial and supports its correlation to MIS 5a rather than MIS 4.

Revision of the occurrence of muskox (Ovibos moschatus Zimmermann 1780) from the Gravettian of Arbreda Cave (Serinyà, northeastern Iberian Peninsula): new insights for the study of Iberian cold‐adapted faunas

During the Last Glacial cycle (MIS 4 to MIS 2), the isthmic Pyrenees mountain range acted as a biogeographical barrier, effectively restricting faunal exchanges to its western and eastern terminations. The study of the composition of megaherbivore communities has revealed the transitional character of the Cantabrian region (northwest Iberian Peninsula) during the MIS 3, between western European cold faunas and southern Iberian temperate faunas. In the Catalan area (northeastern Iberian Peninsula), cold‐adapted faunas are frequently recorded during the cold pulses of MIS 3 and MIS 2, albeit to a lesser extent. In the present paper, we provide the first direct evidence of a muskox record in the Iberian Peninsula, together with a morphometric study and the results of collagen δ13C and δ15N analysis. The results reveal that the Arbreda specimen fits biometrically within the Late Holocene Umingmak specimens (Banks Island, Canada). Moreover, its isotopic values fall into the range of Late Pleistocene Mammoth steppe muskoxen, which indicates a typical ecological behaviour for muskox. Indeed, it suggests a niche partitioning of this cold‐adapted specimen in relation to other Gravettian Iberian fauna. Chronologically, this occurrence is interpreted as an isolated southward expansion of the biogeographical limits of muskox during the Greenland Stadial 4, in the latest MIS 3.

Latitudinal gradients and indicator species in ungulate paleoassemblages during the MIS 3 in W Europe

Studies of mammal assemblages for the European MIS 3 have focused on a regionalization using presence–absence data and traditional cluster analysis. Although this approach has revealed mammal assemblages with no modern analogs, it can easily miss two important dynamical aspects of the MIS 3 assemblages, namely, transitional regions and faunal mixtures. We revisited this regionalization using a complementary approach based on the study of quantitative data for ungulate species. The main aim of this work is to evaluate whether the MIS 3 climatic oscillations led to a mixture of faunas or a latitudinal gradient in faunal composition, as well as to identify indicator species for different areas in ungulate mammal paleoassemblages of Western Europe.Ungulate faunal compositions of 86 fossil assemblages from 63 Western European sites of MIS 3 age were gathered from published sources. The distribution area was divided into four regions: central W Europe, S France, Cantabrian region and rest of the Iberian Peninsula. Chronologies of the assemblages were collected and calibrated. Multivariate ordination and indicator value analysis were carried out.Our study provided four main results: (i) there was a latitudinal faunal turnover during the MIS 3 period consistent with a temperature gradient; (ii) the Cantabrian region hosted a transitional faunal assemblage between the northern cold faunas and the southern temperate faunas; (iii) it was possible to find indicator species of large herbivores for different geographical areas along that latitudinal gradient, although not all of them were linked to climate but to orography (mainly in Cantabrian and other Iberian sites); and (iv) within those indicator species, a signal of mixed temperate and cold faunas was discernible. Together, this evidence supports a scenario in which the succession of cold and warm episodes typical from MIS 3 left signals of mixed cold and temperate faunal assemblages but without totally blurring the latitudinal gradient in climatic conditions.

Recent TIMS dating results from British Late Pleistocene vertebrate faunal localities: context and interpretation

AbstractNew TIMS dates from British sites underpin and extend the growing model of mammalian faunal history during the Late Pleistocene. The sequence of British faunas during the three successive warm phases of MIS 5 presents a challenging model in which the mammal faunas of the first two temperate phases, 5e and 5c is dominated by species of a classically interglacial character, while that of the final phase, sub‐stage 5a, is dominated by bison and reindeer. This we see as the ‘cold fauna’ appearing in Britain during the cool isotope sub‐stage 5b and subsequently isolated by the rising sea level of isotope sub‐stage 5a. All these three faunas are true island faunas. Copyright © 2007 John Wiley & Sons, Ltd.

Art paléolithique, saisons et climats

Palaeolithic art, seasons and climates. During the glaciation of the Würm, the rock and movable art of the hunters – gatherers of Cro-Magnon (approximately 35 000 to 8000 years ago) is made of animals, human beings and geometrical signs. The everyday life is given rhythm by the seasons. They are rather often reflected in the morphology and/or the behaviour of the big herbivores represented. The presence of certain sorts indicates cold episodes: mammoths, woolly rhinoceros, ovibos, reindeers, megaceros. Others testify of reheating (as the interstadium of Lascaux): horses, bisons and especially aurochs, deer and hinds, ibexes. At the end the glaciation, the cold fauna is not any more represented. The great big animal art of the Cro-Magnons disappears, being replaced by an outdoor rock art, simplistic and often narrative. To cite this article: B. Delluc, G. Delluc, C. R. Palevol 5 (2006) .

Le Quaternaire Marin du Cap Achakar (Tanger, Maroc): nótectonique et lithostratigraphie

The lithostratigraphy of the Quaternary deposits of Cap Ashakar (Tangier, Morocco) is made up of three marine Pleistocene formations: 1. i)the oldest indicates an upper shoreface depositional environment and has suggested a “cold fauna” with two common Mediterranean species so far unknown on the Moroccan Atlantic coast: Buccinum undatum and Neptunea contraria; 2. ii) the youngest indicates a foreshore depositional environment and is dated from the last interglacial period; 3. iii) between them, a very thick marine Precambrian formation took place, which indicates an upper shoreface environment followed by a foreshore depositional environment. The morphostructural analysis illustrates the role played by the recurrent epeirogenic phenomena in the morphogenesis of the north Moroccan Atlantic coast and underlines the occurrence of areas with differential motions compared to the general uplift of the Gibraltar Strait.

The Plio-Pleistocene boundary in Southeast Spain: A review

The Plio-Pleistocene boundary in the Mediterranean region has been defined by the entry of the so-called 'northern guests'. The absence of this characteristic cold fauna in the sedimentary record of littoral marine basins in SE Spain has led to the local establishment of this boundary by other means. Most outcropping upper Pliocene and Pleistocene marine sediments in this area have a very shallow character, resulting in the absence of the planktonic foraminiferal assemblages that characterize the upper Pliocene. As a result the only means of determining this boundary has been through regional correlation, detailed mapping and sedimentological analysis, as well as palaeomagnetism. After studies were carried out in different basins, and accepting the age of 1.64 (now recalibrated to 1.8) Ma for the Plio-Pleistocene boundary (Aguirre and Pasini, 1985), we conclude that this boundary does not bear any climatic, environmental or tectonic signature. The changes in most of these basins seem to occur either before or after this date.

Le Quaternaire marin du secteur littoral Cap Achakar -Cap Spartel (Tanger, Maroc) : lithostratigraphie et néotectonique

Marine Quaternary of Cap Ashakar (Tangiers, Morocco) : lithostratigraphy and neotec tonics. Lithostratigraphy of the marine quaternary deposits of Cap Ashakar (Tangiers, Morocco) allows us to reveal three marine pleistocene formations : • the oldest indicates an upper shoreface depositional environment and has given up a "cold fauna" with two common mediterranean species so far unknown on the Moroccan atlantic coast : Buccinurn undatum and Neptunea contraria ; • the youngest indicates a foreshore depositional environment and dates from the last Interglacial ; • between them, takes place a very thick marine azoic formation, which indicates an upper shoreface followed by a foreshore depositional environment. The morphostructural analysis allows to show the part of the recurrent epeirogenic phenomenas in the morphogenesis of the north Moroccan atlantic coast.

Intermediate steps of southwards faunal shifts testified by last glacial submerged thanatocoenoses in the Atlantic ocean

Several southwards faunal shifts (SFS) of “boreal” species (BGs: Boreal Guests of authors) are known to have occurred in the colder periods of the E. Atlantic Pleistocene. The latest invasion of BGs took place during the last glacial (75,000–10,000 yr B.P.) and is recorded all over the Mediterranean by a number of submerged thanatocoenoses containing sublittoral to bathyal faunas. On the other hand, these SFS are quite poorly documented within the Atlantic Ocean. Pre-Modern Quaternary thanatocoenoses have been discovered at many sites along the eastern Atlantic margin between ca. 35° and 48°N. These assemblages contain typical BGs ( Modiolus modiolus, Chlamys islandica, Pseudamussium septemradiatum, Arctica islandica, Trichotropis borealis, Buccinum undatum) and/or scleractinian corals ( Lophelia pertusa, Madrepora oculata, Caryophyllia smithi). Samples of representative species were radiocarbon dated showing four age clusters. Cluster 1 (ca. 27,000 yr B.P.) is relative to the pleniglacial and is documented by C. islandica assemblages in the Bay of Biscay and by coral thanatocoenoses in the Ibero-Moroccan Gulf; cluster 2 (ca. 12,000–13,500 yr B.P.) represents latest glacial faunas containing BG-assemblages of milder-climatic significance ( M. modiolus) and corals dredged in the Ibero-Moroccan Gulf; cluster 3 (7500–10,000 yr B.P.) may represent retreating time-delayed and/or postglacial acclimatization of mild-BGs ( P. septemradiatum) in the Ibero-Moroccan Gulf; finally, cluster 4 is relative to quite recent assemblages (late Holocene) containing potential BGs ( B. humphreysianum) the paleoclimatic significance of which has to be revised. Our results show that intermediate steps (missing links) of SFS during the Quaternary between the NE Atlantic BG-reservoir and the Mediterranean are more frequent than previously supposed calling for their careful survey in order to shed light on expansion and contraction of cold faunas during the last glacial.

Radiolarian responses to recent and neogene California El Niño and anti-El Niño events

Plankton tows from 1983 and 1964 California El Niños suggest that high standing crops of both warm and cold radiolarian faunas are characteristic of these conditions over the southern California continental borderland. High standing crops of both warm and cold radiolarians are suggested to occur during California El Niños because the California Current diminishes in intesity allowing warm faunas to invade from the west (central-gyre waters) and south (gyre and eastern tropical Pacific waters); and, a strongly developed California Countercurrent aids in creating a geostrophic dome that allows cold faunas to invade from the north. These same radiolarian faunas characteristic of California El Niños are found in the Holocene varved sediments of the Santa Barbara Basin and can be correlated to the known California El Niño events of 1957–1958 and 1964. El Niño like events determined via the same radiolarian criteria are found at 6 and 8 Ma in Monterey laminated (presumably varved) sedimentary rocks. It is suggested that warm and cold periods in Monterey may be related to the frequency and magnitude of El Niño like events. It is also suggested that during Neogene warm periods (such as at 8 and 10 Ma) the paleo-California Current was narrower, closer to the coast, and did not extend as far south as during anti-El Niño periods (such as at 5 Ma).

The western border of the Indian plate: implications for Himalayan geology

The study of regions situated beyond the western margin of the present-day Indian plate (Afghanistan principally) point to the following facts: 1. (1) During the Late Precambrian—Early Paleozoic, stratigraphical continuity existed between western and central Iran, Central Afghanistan, Salt Range and western Pakistan. 2. (2) During the Paleozoic a similar epicontinental cover existed in central Afghanistan, Kashmir and Tibet, with Gondwana tillites and associated cold fauna, such as in India (Umaria); however, a so-called Hercynian zone exists also in northern Iran—Hindu Kush and northern Pamir: it exibits a Middle Paleozoic unconformity (Upper DevonianCarboniferous) on metamorphic Early Paleozoic. 3. (3) The end of the Paleozoic, is marked by: a fracturation of the basement of the Hercynian zone, with powerful volcanic eruptions at the northern part of Hindu Kush, Kashmir (Panjal trap) and also Nepal (Nar valley) the formation of a geosynclinal zone at the southern part of Hercynian zone (Turkman, Penjaw). 4. (4) During the Jurassic: the geosynclinal evolution of the Turkman—Penjaw furrow accelerated, with the accumulation of flysch, radiolarites, ophiolites, olistolites and incipient HP metamorphism. A general subduction took place followed by a Neocimmerian orogenic phase with overthrusting of the central Afghanistan ranges on the scar of the geosynclinal furrows. 5. (5) During the Cretaceous: the geosynclinal evolution ended: Lower Cretaceous lies unconformably on the folded Jurassic flysch. In eastern Afghanistan and northern Pakistan, during the Middle (?) or Upper Cretaceous, a new geosynclinal zone was created. 6. (6) During the Cenozoic, central Afghanistan was emerged; northwards, sedimentary basins were created along the Herat fault, with volcanic and magmatic activity. A southeastern geosynclinal furrow evolved with accumulation of flysch, ophiolites and finally molasse deposits (Katawas—Soleimans). Its western border began overthrusting, but this movement changed into a left lateral fault i.e., the presentday Chaman Arghandeh fault. Conclusion: Two major phases of dislocation took place during the geological history of Gondwana: the first one began during the Permian and ended in the Jurassic; the second one began during the Cretaceous and is still active. The important Eocimmerian orogenic phenomena, existing in the Central Afghanistan and northern Pakistan, took place at the edge of a Gondwana continental fragment, which was larger than the presentday Indian plate. Coeval phenomena may exist in the Himalayan region and perhaps in one of the ophiolitic sutures of Tibet.

Late Devensian periglacial phenomena in the upper Cam Valley, north Essex

Detailed stratigraphic investigations in the upper Cam Valley around Newport, Essex, have indicated the presence of discontinuous till-like deposits (diamictons) and distinct Taele Gravel ridges intimately associated with an irregular non-terrace gravel formation (valley gravels). The diamictons are shown by particle size analysis and fabric analysis to be remnants of plateau till (Chalky Boulder Clay) derived and reworked by mass movement (earthflow and debris sliding). In places the diamictons interdigitate with the valley gravels which contain a cold fauna and appear to have been aggraded by accelerated mass movement into mounds and ridges. This irregular relief has impounded several small mere-like lakes whose basal sediments are assigned by palynology and 14 C dating to Late Glacial Zone III. A Late Devensian age is thus inferred for the processes of mass movement whose initiation was probably related to intense permafrost conditions.

Discussion

D r. R. G. W est said that the paper brought forward two main points; first, the glacial sequence of East Anglia and the reality of the Gipping Till and secondly, the correlation of this sequence with that of the continent. These two issues should be considered separately. Taking the definition that a cold stage in the Pleistocene is characterized by the presence of glacial deposits, periglacial deposits or a cold fauna or flora, and a temperate stage (interglacial) by evidence for temperate conditions, then there is evidence in East Anglia for a cold stage between the Hoxnian and Ipswichian interglacials, in the form of periglacial floras above the Hoxnian and late-glacial floras below the Ipswichian. The simplest interpretation of these observations is that there was a single cold stage between the Hoxnian and Ipswichian. There might be more. To this cold stage have been ascribed tills (Gipping Till) and fluvioglacial gravels of north Norfolk. The authors must be right in saying that the widespread chalky boulder clay of Norfolk and Suffolk was pre-Hoxnian. The problem was of the extent of a post-Hoxnian till. It appeared that there was no evidence for a widespread occurrence of post-Hoxnian till. Perhaps some tills in the Breckland might belong here. The conclusion was that it yet remained to be shown what the extent of glaciation was in the post-Hoxnian cold stage. It seemed probable to the speaker that while, as Dr. Bristow and Dr. Cox point out, there is little evidence for a widespread till

La repartition actuelle des mollusques marins envisagee comme base d'appreciation pour les geologues

Abstract Discusses various features of the present-day distribution of marine mollusks, such as habitat, migration, and warm and cold faunas, which are of geologic significance.

A Comparison of the Pleistocene of East Anglia with that of Germany

For many years, Professor P. G. H. Boswell has studied the Pleistocene deposits of East Anglia and, from time to time, published most illuminating reviews summarising the progress of work and discussing the possible relations to the corresponding deposits of other districts (especially 1931, 1932, 1936). In his Presidential Address to the Prehistoric Society last year, he paid particular attention to a problem which has often been attacked but not yet solved satisfactorily, namely the correlation of East Anglia with the Continent; and he suggested, as a possible way out of the difficulties, the correlation of the Hunstanton Boulder Clay with Würm 2, the Upper Chalky Drift with Würm 1, the Great Chalky Boulder Clay with Riss, the North Sea Drift with Mindel, and the later Crag deposits containing a cold fauna, with Günz. He admitted, however, that such a correlation would ‘bring other difficulties in its train.’

Societies and Academies

LONDON. Geological Society, June 25.—Dr. J. W. Evans, president, in the chair.—Miss M. E. Tomlinson: The river-deposits of the lower valley of the Warwickshire Avon; with an appendix by A. S. Kennard and W. J. Woodward. Fluviatile sands and gravels occur as river-terraces between Stratford and Tewkesbury. The surfaces of these terraces lie on curves parallel to the present thalweg of the Avon. Five such terraces have been identified. No. 5 Terrace, the highest above river-level, has yielded no contemporaneous fossils. No. 4 and No. 3 both contain a warm fauna, with Hippopotamus and Belgrandia marginata occurring solely at No. 3 level, and Corbicula fluminalis at No. 4 level. No. 2 contains a cold fauna, with numerous remains of mammoth and Rhinoceros tichorhinus. No. 1 probably underlies the alluvium, and may be connected with the infilling of the buried channel which has been identified at Fladbury and Tewkesbury. The sole evidence of human industry found is a fresh flint of Mousterian type from No. 2 Terrace.—F. Raw: The development of Leptoplastus salteri Callaway, and of other trilobites (Olenidæ, Ptycho-paridæ, Conocoryphidæ, Paradoxidae, Phacopidae, and Mesonacidae). Leptoplastus salteri occurs in the Upper Tremadoc horizon of Shineton Brook (Salop). All stages of development are noted, except the protaspis. The development is divided into successive stages. The earliest stage has a long, simply segmented glabella; long pleural spines occur throughout the body, including three pairs on the head and four on the pygidium; while axial spines occur from the occipital segment throughout. In the last stage the glabella is short, conical, and smooth; of pleural spines, only the genal remain on the head, those in the thorax are reduced, and none remain on the pygidium, where axial spines are also- wanting. With certain reservations, these four stages are also claimed to represent successive stages in the phylogeny. This ontogeny sheds much light on that of other trilobites. In the ontogenies of several families, it is the primitively posterior head-spines that at first are dominant to be superseded later by the middle pair. In Mesonacidæ these again are succeeded by the anterior pair, which, in their lateral revolution, carried before them the anterior branches of the facial sutures, so that these cut the posterior border; they also stretched out the antero-marginal suture to the genal angle. These changes led to the supersession of the dorsal facial suture by the marginal suture, with consequent ankylosis along the former. The family is hereby claimed to have a head-structure not hitherto recognised. It illustrates a general principle—that those only of the cephalic sutures are retained as were necessary for ecdysis. The combined study of trilobite ontogeny and morphology strongly suggests, as Beecher claimed, that here the ontogeny extensively recapitulates the phylogeny; and therefore best indicates relationships. This indicates an early divergence in glabella-form, antedating divergences in cephalic spines and sutures. Members of a “natural “order may, therefore, be expected to agree much more in glabella-form than in the cephalic suture, which has latterlv been made the basis of classification.

Vertebrate Remains in the Caverns of Grimaldi

PROF. MARCELLIN BOULE has completed his studies of the remains of vertebrate animals found with primitive man in the caverns of Grimaldi, and the final results of his work have just been published as the concluding part (fasc. iv.) of the first volume of the Prince of Monaco's “Les Grottes de Grimaldi (Baousse-Rousse).” The new instalment deals with the Carnivora, Insectivora, Cheiroptera, and Rodentia, and various fragments of birds and lower vertebrates, and ends with a valuable general summary. Besides the technical descriptions of the fossils, illustrated by beautiful plates in heliogravure, Prof. Boule continually introduces short discussions of the relationships and distribution of the various animals with which he deals, adding- several maps and some genealogical diagrams. He has therefore produced a most interesting and readable treatise on the European Pleistocene vertebrate fauna, which we commend to the notice of both geologists and zoologists. He specially emphasises the importance of the discovery that in the low latitude of the south coast of France there is the same succession of Pleistocene mammalian faunas that has already been observed throughout the rest of Central and Western Europe. Jn the bottom layers on the floor of the caverns of Grimaldi there are the animals of the warm Chellean episode (Elephas antiquus, Rhinoceros Mercki, hippopotamus, etc.); in the next layers is the cold fauna of the Acheulian and Mousterian (glutton, ermine, marmot, reindeer, etc.); in the upper layers are the ordinary mammals of historic times. Among the animals now described Prof. Boule considers he can recognise every gradation between the Pliocene bears and the modern brown bear; he also sees some approach to a Pliocene species in the Pleistocene leopard. He agrees with other observers that the wild cat most nearly approaches that of Africa, now named Felis ocreata. Equally interesting is his account of the fossil lynx, which proves to be exactly intermediate between the northern and the Spanish races of the lynx at the present day.

Paul Henri Fischer

THE Museum of Natural History of Paris has suffered a great loss in the person of Dr. Paul Henri Fischer, the well-known zoologist and palæontologist, who died on November 29, after a long and painful illness. Born at Paris, on July 7, 1835, he received his early classical and medical education at Bordeaux. He became Interne des Hôpitaux of Paris in 1859, and obtained his degree of Doctor of Medicine in 1863. The study of medicine did not prevent him from devoting himself also to that of the natural sciences; for in 1861 he entered as Demonstrator in the Laboratory of Palæontology of the Museum of Paris, under the direction of M. d'Archiac. His researches chiefly concerned the living and fossil mollusca, and from 1856 he edited the Journal de Conchyliologie in collaboration with M. Crosse. From the position of Demonstrator he rose to be aide-naturaliste (assistant), and studied with great success the marine animals of the coasts of France, their geographical and bathymetric distribution. He indicated the depths at which a large number of foraminifera, cœlenterata, echinodemata, mollusca, bryozoa, &c. can be collected on the coasts of the west of France. In collaboration with the Marquis de Folin he undertook the study of the animals dredged in the extremely interesting region of the Gulf of Gascony, to which the name “Fosse du Cap Breton” has been given. The two savants discovered a large number of forms hitherto unknown, and many which recalled species only known in the fossil condition. With M. Delesse he made researches on the submarine sediments of the French shores. He was elected member of the Commission of Dredging, and took part from 1880 to 1883, on board the Travailleur and the Talisman, in the celebrated expeditions directed by Prof. Milne Edwards. In the course of these expeditions he noted the enormous extension of a cold fauna characterised by boreal and arctic species, and reaching as far as Senegal, where it lives beneath a superficial fauna with intertropical characters. Among the writings of Dr. Fischer, which number not less than 300 titles, including books, pamphlets and memoirs, we may cite: “Paléontologie de I'Asie mineure” (in collaboration with MM. d'Archiac and de Verneuil); “Mollusques de Mexique et de I'Amerique Centrale”; “Species général et iconographie des coquilles vivantes”; “Animaux fossiles du Mont Léberon” (in collaboration with MM. Gaudry and Tournouer); “Paléontologie de I'île de Rhodes”; “Cétacés du Sud-Ouest de la France”; “Catalogue et distribution géographique des mollusques terrestres, fluviatiles et marins d'une partie de I'Indo-Chine; “Sur les caractères de la faune conchyliologique terrestre et fluviatile recemment éteinte du Sahara”; “Sur la faune conchyliologique de I'île d'Haïnan”; numerous memoirs on the malacological fauna of Lord Hudson Island (Pacific Ocean), of Cambodge, of the islands of the Caledonian Archipelago, of Aleutian islands, of the Bay of Suez, &c. In collaboration with M. E. L. Bouvier he published papers on the anatomical peculiarities of certain groups of molluscs. Finally, he wrote a remarkable treatise on conchology which has become classical (“Manuel de Conchyliologie et de paléontologie conchyliologique ou histoire naturelle des mollusques vivants et fossiles, suivi d'un appendice sur les Brachiopodes par Œhlert.” In this manual the author showed that the classification of molluscs ought to be based not alone on the form of the shell, but primarily on the anatomical characters.