Radiolarian faunal turnover across the Oligocene/Miocene boundary in the equatorial Pacific Ocean

The global warming trend of the latest Oligocene was interrupted by several cooling events associated with Antarctic glaciations. These cooling events affected surface water productivity and plankton assemblages. Well-preserved radiolarians were obtained from upper Oligocene to lower Miocene sediments at Ocean Drilling Program (ODP) Leg 199 Sites 1218 and 1219 in the equatorial Pacific, and 110 radiolarian species were identified.Four episodes of significant radiolarian faunal changes were identified: middle late Oligocene (27.5 to 27.3 Ma), latest Oligocene (24.4 Ma), earliest Miocene (23.3 Ma), and middle early Miocene (21.6 Ma). These four episodes approximately coincide with increases and decreases of biogenic silica accumulation rates and increases in δ¹⁸O values coded as “Oi” and “Mi” events. These data indicate that Antarctic glaciations were associated with change of siliceous sedimentation patterns and faunal changes in the equatorial Pacific.Radiolarian fauna was divided into three assemblages based on variations in radiolarian productivity, species richness and the composition of dominant species: a late Oligocene assemblage (27.6 to 24.4 Ma), a transitional assemblage (24.4 to 23.3 Ma) and an early Miocene assemblage (23.3 to 21.2 Ma). The late Oligocene assemblage is characterized by relatively high productivity, low species richness and four dominant species of Tholospyris anthophora, Stichocorys subligata, Lophocyrtis nomas and Lithelius spp. The transitional assemblage represents relatively low values of productivity and species richness, and consists of three dominant species of T. anthophora, S. subligata and L. nomas. The characteristics of the early Miocene assemblage are relatively low productivity, but high species richness. The two dominant species present in this assemblage are T. anthophora and Cyrtocapsella tetrapera. The most significant faunal turnover of radiolarians is marked at the boundary between the transitional/early Miocene assemblages.We also reviewed changes in other microfossil assemblages in the low latitudes during the late Oligocene through early Miocene. The microfossil assemblages of major groups show sequential changes near the Oligocene/Miocene (O/M) boundary (23.8 Ma). Many extinction events and some first occurrences of calcareous nannofossils and many occurrences of radiolarians are found from about 24.8 to 23.3 Ma, and first occurrences of planktic foraminifers and diatoms followed from 23.2 through 22 Ma. Hence, the O/M boundary is identified as a significant level for microfossil evolutions.     

Fish otoliths from the pre-evaporitic (Early Messinian) sediments of northern Italy: their stratigraphic and palaeobiogeographic significance

The study of otolith assemblages from the pre-evaporitic Messinian deposits allows the reconstruction of a fauna of 79 taxa of which 35 could be identified at the specific level. Three of these are new: Diaphus rubus, Myctophum coppa, and Uranoscopus ciabatta. The assemblages reflect mainly a neritic environment influenced by the oceanic realm. Analysis of the global present-day geographic distribution of 42 of the recognised Messinian genera indicates that 88% of these are still living in the Mediterranean, 98% in the Atlantic and 78% in the Indo-Pacific realm. These results are in good agreement with the evolutionary trends documented for the Oligocene and Miocene teleost fauna, specifically an increase in percentage of genera inhabiting the modern Mediterranean, a very high percentage of Atlantic and Indo-Pacific genera, and a slight fall of the importance of present-day Indo-Pacific genera from the Rupelian up to the Late Miocene. Analysing the composition of the Early Messinian fauna at the level of nominal species indicates that about 53% of the species represented in the assemblages are still living in the Recent Mediterranean, and that a significant number of these were already present in the Tortonian. It is interesting that these species are mainly neritic. This seems to confirm that the close affinity of the fossil assemblage with the present-day Mediterranean neritic fauna, which was already recorded at the genus level for the Rupelian fauna, persists during the Neogene and continues until the Pleistocene.     

A new species of fossil Scutus Montfort, 1810 from New Zealand (Mollusca: Gastropoda: Fissurellidae)

ABSTRACT

A new fossil species of the genus Scutus (Scutus mirus n. sp.) is described from five Late Oligocene to Early Miocene (Waitakian to Altonian; 25.2–15.9?Ma) localities in the South Island, New Zealand. It is one of the oldest fossil species of Scutus known and probably inhabited very shallow, sub-tropical waters surrounding Zealandia during this time. The holotype of Scutus petrafixus Finlay, 1930 is re-examined; it is possibly from All Day Bay, Kakanui (Waitakian 25.2–21.7?Ma). The New Zealand species documented herein significantly expand our understanding of the fossil record of this shallow-marine molluscan lineage, and by proxy, also indicate the presence of very shallow coastal marine environments around the late Oligocene and early Miocene in southern Zealandia.     

A new coral‐associated decapod assemblage from the Upper Miocene (Messinian) Upper Coralline Limestone of Malta (Central Mediterranean)

Abstract: A rich coral‐associated decapod assemblage is recorded from the ‘Depiru Beds’ of the upper part of the Upper Coralline Limestone (Messinian, Upper Miocene), from the island of Malta. Nineteen species within 17 genera have been discovered, where 14 genera are new for Malta. Four new species are described, namely Micippa annamariae sp. nov., Pilumnus scaber sp. nov., Panopeus muelleri sp. nov. and Herbstia melitense sp. nov. Herbstia melitense sp. nov. constitutes the first record of the genus from the fossil record in the Mediterranean region. This discovery more than doubles the number of known fossil decapod species from Malta. The fossil bivalve Jouannetia (J.) semicaudata Des Moulins, 1830 and the extant decapod Maja goltziana D’Oliveira, 1888, are also recorded for the first time from Malta. Other Neogene coral‐associated decapod assemblages are investigated and correlated with the new assemblage from Malta. The migration of taxa between the Mediterranean region and the Paratethys, particularly during the Lower Badenian (Langhian), is evidenced by the strong affinity of the Maltese decapod assemblage with that of the Middle Miocene Badenian assemblages from Hungary, Poland and Ukraine. Upper Miocene, Messinian assemblages from Spain, Algeria and Morocco are also similar to that from Malta.     

Small mammal (rodents and lagomorphs) European biogeography from the Late Oligocene to the mid Pliocene

Aim To analyse the fossil species assemblages of rodents and lagomorphs from the European Neogene in order to assess what factors control small mammal biogeography at a deep‐time evolutionary time‐scale. Location Western Europe: 626 fossil‐bearing localities located within 31 regions and distributed among 18 successive biochronological units ranging from c. 27 Ma (million years ago; Late Oligocene) to c. 3 Ma (mid Pliocene). Methods Taxonomically homogenized pooled regional assemblages are compared using the Raup and Crick index of faunal similarity; then, the inferred similarity matrices are visualized as neighbour‐joining trees and by projecting the statistically significant interregional similarities and dissimilarities onto palaeogeographical maps. The inferred biogeographical patterns are analysed and discussed in the light of known palaeogeographical and palaeoclimatic events. Results Successive time intervals with distinct biogeographical contexts are identified. Prior to c. 18 Ma (Late Oligocene and Early Miocene), a relative faunal homogeneity (high interregional connectivity) is observed all over Europe, a time when major geographical barriers and a weak climatic gradient are known. Then, from the beginning of the Middle Miocene onwards, the biogeography is marked by a significant decrease in interregional faunal affinities which matches a drastic global climatic degradation and leads, in the Late Miocene (c. 11 Ma), to a marked latitudinal pattern of small mammal distribution. In spite of a short rehomogenization around the Miocene/Pliocene boundary (6–4 Ma), the biogeography of small mammals in the mid Pliocene (c. 3 Ma) finally closely reflects the extant situation. Main conclusions The resulting biogeographical evolutionary scheme indicates that the extant endemic situation has deep historical roots corresponding to global tectonic and climatic events acting as primary drivers of long‐term changes. The correlation of biogeographical events with climatic changes emphasizes the prevalent role of the climate over geography in generating heterogeneous biogeographical patterns at the continental scale.     

Mammalian biochronology of the Paleogene-Neogene boundary at Aktau Mountain, eastern Kazakhstan

At Aktau Mountain in the Ili depression of eastern Kazakstan, fossil mammals that encompass the Paleogene-Neogene boundary occur at three stratigraphic levels. The lowest level is in the lower Kyzylbulak Formation and produces Brontotheriidae and the hyracodontidArdynia and is tentatively assigned a late Eocene (Ergilian) age. The lower part of the overlying Aktau Formation produces fossils of the giant rhinocerosParaceratherium and is tentatively assigned a late Oligocene (Tabenbulukian) age. The upper part of the Aktau Formation yields a fossil mammal assemblage that includesGomphotherium,Stephanocemas, Brachypotherium andLagomeryx. It is clearly of Miocene age, probably late early Miocene (late Burdigalian), a correlative of European Reference Level MN 5 and the late Shanwangian of China. The Paleogene-Neogene boundary at Aktau Mountain thus is in the Aktau Formation.     

Sponge borehole size as a relative measure of bioerosion and paleoproductivity

Bioerosion intensity has been proposed as a measure of paleoproductivity in fossil reefs, but it is difficult to measure directly because fossil corals are often incomplete and because it is difticult to infer the length of time a given coral was exposed to bioeroding organisms. Both nutrient availability and taphonomic factors can affect bioerosion intensity as measured in dead corals. Here, we examine these two effects separately using data from previous studies on bioerosion in modern and fossil corals. Size of individual sponge borings accurately reflects total bioerosion in modern massive and branching corals on the Great Barrier Reef. Total bioerosion in both massive and branching corals decreases outward across the continental shelf, paralleling trends in nutrient availability. Size of individual Cliothosa hancocki borings decreases across the shelf in branching Acropora but not in massive Porites. Fossil sponge borings Entobia convoluta and Uniglobites glomerata in massive corals from Oligocene and Miocene reefs in Puerto Rico are smallest in Oligocene shelf-edge reefs, intermediate in Oligocene patch reefs, and largest in Miocene patch reefs. Both facies-related influence, represented by Oligocene shelf-edge reefs vs. Oligocene patch reefs, and nutrient-related influence, represented by Oligocene vs. Miocene patch reefs, were reflected in the size of sponge boreholes. Size of sponge borings also varies among species of host corals, apparently in relation to skeletal architecture. Borehole size is inversely correlated with skeletal density as measured by the relative proportion of skeleton and pore space in transverse thin section. There is a weak positive correlation between borehole size and corallite diameter. These findings contradict reported positive correlations between total bioerosion and bulk density in modern corals. Borehole size appears accurately to reflect intensity of total internal bioerosion in fossil corals. Facies-controlled taphonomic overprints and influence of skeletal differences between coral species limit the use of sponge borehole size to a rough indicator of paleoproductivity in fossil coral reef environments.     

A Review of the Eurasian fossil species of the bee Apis

Fossil Apis species from the Oligocene, Miocene and Plio-Pleistocene of Eurasia are described and their relationships with Recent species are discussed. Several new populations of fossil bees are reported from the Oligocene and Miocene of France and Spain, including Apis aquisextusensis sp. nov. The present state of knowledge of fossil bee systematics is poor because of the general lack of preserved characters. Some of the problems, and items requiring further investigation, are identified.     

Stratigraphy and microfacies of the oligocene sequence at Gabal Bu Husah, Marada Oasis, South Sirte Basin, Libya

Summary Stratigraphic and microfacies investigations carried out on the Oligocene sequence exposed at Gabal Bu Husah, northwest of Marada Oasis, south of the Sirte Basin, Libya, showed that the Oligocene sequence conformably overlies the Late Eocene Thamat Formation and unconformably underlies the Early to Middle Miocene Marada Formation (Qarat Jahannam Member). The lithostratigraphic studies of the Oligocene sequence revealed the presence of two rock units, from base to top: 1- Umm Ad Dahiy Formation (Early Oligocene, Rupelian) and 2- Bu Hashish Formation (Late Oligocene, Chattian). The Oligocene sequence yields a rich foraminiferal assemblage with fifty-one benthonic and large formaminiferal species. The foraminiferal analysis allowed to subdivide the sequence into three local foraminiferal assemblage zones, arranged stratigraphically from base to top: 1)Elphidium minutum zone comprising the Umm Ad Dahiy Formation. 2)Miogypsinoides complanatus/Nonion granosus assemblage zone covering the lower two thirds of the Bu Hashish Formation. 3) Zone with abundantNummulites spp., including the upper part of the Bu Hashish Formation. The paleoenvironmental significance of the recorded species is described and discussed. The microfacies studies led to the recognition of eleven microfacies types. These microfacies indicate that the lower part of the Umm Ad Dahiy Formation was deposited in a shallow warm marine environment, but the conditions changed to fluviomarine in the uppermost part. The Bu Hashish Formation was deposited in a shallow-marine, inner-shelf environment (as indicated by the operculinid limestone) but a probable hiatus in its middle part is indicated by the presence of a nummulitic coquina and gypsum beds formed in a lagoonal environment. After returning to a shallow marine environment at the end of Oligocene the marine sedimentation ended.     

Major Radiations in the Evolution of Caviid Rodents: Reconciling Fossils,Ghost Lineages,and Relaxed Molecular Clocks

Background

Caviidae is a diverse group of caviomorph rodents that is broadly distributed in South America and is divided into three highly divergent extant lineages: Caviinae (cavies), Dolichotinae (maras), and Hydrochoerinae (capybaras). The fossil record of Caviidae is only abundant and diverse since the late Miocene. Caviids belongs to Cavioidea sensu stricto (Cavioidea s.s.) that also includes a diverse assemblage of extinct taxa recorded from the late Oligocene to the middle Miocene of South America (“eocardiids”).

Results

A phylogenetic analysis combining morphological and molecular data is presented here, evaluating the time of diversification of selected nodes based on the calibration of phylogenetic trees with fossil taxa and the use of relaxed molecular clocks. This analysis reveals three major phases of diversification in the evolutionary history of Cavioidea s.s. The first two phases involve two successive radiations of extinct lineages that occurred during the late Oligocene and the early Miocene. The third phase consists of the diversification of Caviidae. The initial split of caviids is dated as middle Miocene by the fossil record. This date falls within the 95% higher probability distribution estimated by the relaxed Bayesian molecular clock, although the mean age estimate ages are 3.5 to 7 Myr older. The initial split of caviids is followed by an obscure period of poor fossil record (refered here as the Mayoan gap) and then by the appearance of highly differentiated modern lineages of caviids, which evidentially occurred at the late Miocene as indicated by both the fossil record and molecular clock estimates.

Conclusions

The integrated approach used here allowed us identifying the agreements and discrepancies of the fossil record and molecular clock estimates on the timing of the major events in cavioid evolution, revealing evolutionary patterns that would not have been possible to gather using only molecular or paleontological data alone.     

A new species of Macaranga from the middle Miocene of Fujian,China and its significance

A well-preserved Macaranga leaf fossil from the middle Miocene Fotan Group of Zhangpu County, Fujian, South-eastern China is examined and described as a new species, Macaranga zhangpuensis Z.X. Wang et B.N. Sun sp. nov. The present fossil species represents the highest latitudinal distribution of a reliable Macaranga fossil in the world, and we present the first Macaranga fossil described with detailed cuticular characteristics from China. Based on the global palaeogeographic distribution of Macaranga, we infer that the genus probably originated during the Oligocene in Africa and spread from Africa to India and then to South Fujian, China, further into the Mariana Islands and finally into the Philippine Islands and Malaysia, leading to its present distribution. In addition, the new leaf material described herein is the first Macaranga fossil record with three leaf tips globally. This finding demonstrates that Macaranga with three leaf tips already existed during the Miocene in Fujian, South China, and it provides new information for understanding climatic changes between the Miocene and the present -day.     

Early Oligocene fruits and leaves of Burretiodendron (Malvaceae s.l.) from South China

The genus Burretiodendron Rehder is currently endemic to an area near the China–Vietnam border and the limestone mountains of Thailand and Myanmar. The fossil records of this genus were previously found only from the Miocene of Yunnan, Southwest China, and the Oligocene of Guangxi, South China. Here, we describe fossil fruits and associated leaves of Burretiodendron, which were discovered in the lower Oligocene Shangcun Formation of the Maoming Basin, Guangdong, South China. Morphological comparison with extant and fossil Burretiodendron taxa indicates that fruit fossils belong to the species Burretiodendron parvifructum J. Lebreton Anberrée & Z. K. Zhou. This is one of the earliest fossil records of the genus, providing additional evidence for the early biogeographic history of this genus and supporting the inference that the genus originated in South China. According to the habitat conditions of modern species, we speculate that there were limestone mountains around the Maoming Basin in the early Oligocene.     

Des chambres de pupation de Dermestidae (Insecta : Coleoptera) sur un os de mammifère tertiaire (phosphorites du Quercy) : implications taphonomiques et paléoenvironnementales

The larvae of some species of Dermestidae (Coleoptera) are able to bore the bones of a carrion for pupation. Several pupal chambers have been observed on an unciform belonging to a fossil rhinocerotid (Mesaceratherium sp.) from the late Oligocene/earliest Miocene of the phosphorites of Quercy (SW France). Such an unusual observation demonstrates the occurrence of dermestids in the entomofauna of the phosphorites of Quercy, and suggests at least the existence of paleoclimatic episodes with dry seasons, during a period usually considered as a relatively humid one. This is the earliest occurrence of such ichnofossils on a fossil mammal bone.     

Italian Cenozoic crocodilians: taxa, timing and palaeobiogeographic implications

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

New paleoenvironmental insights on the Miocene condensed phosphatic layer of Salento (southern Italy) unlocked by the coral-mollusc fossil archive

From the Late Oligocene to the Late Miocene, the central Mediterranean area was characterized by the extensive deposition of phosphate-rich sediments. They are usually represented by 10 to 20-cm-thick hardgrounds made of phosphatic and glauconitic sediments containing a rich macrofossil association. This study represents the first thorough investigation of the biotic assemblage of Mediterranean phosphorites aimed at collecting new information on the environmental factors controlling their deposition. The Serravallian/Tortonian phosphatic deposits of the Salento Peninsula (“Aturia level”) have been selected for the abundance of fossil remains and special attention is given to the coral–mollusc association. Two different facies have been recognized: a basal coral rudstone that includes most of the macrofossils, superimposed by a detrital rudstone made of thin layers mainly composed of phosphatic fragments. These two facies are separated by a phosphatic crust several millimeters in thickness. The coral assemblage contains at least 17 azooxanthellate taxa belonging to four families, while the molluscs are represented by a rich gastropod fauna (26 species), associated with bivalves (18 species) and cephalopods (two species). Four distinct depositional phases have been recognized, with the coral rudstone representing the key-facies to reconstruct the onset of the “Aturia level” and the original environment of its fossil content. The composition of the coral–mollusc association has been reliably compared with present-day analog taxa, suggesting the occurrence of a heterogeneous seafloor formed by rocky substrates and accumulations of soft sediment, at around 100–350-m water depth, and under the influence of moderate-to-strong bottom currents rich in nutrients and resuspended organic matter.     

FILLING A GAP: THE FIRST OCCURRENCES OF EPIPHAXUM (CNIDARIA: HELIOPORACEA: LITHOTELESTIDAE) IN THE EOCENE, OLIGOCENE AND MIOCENE

Abstract:  A new species of the genus Epiphaxum (family Lithotelestidae) is described and illustrated in detail, and compared to other species. Epiphaxum arbuscula sp. nov. has been collected from Upper Eocene (Priabonian), Upper Oligocene (Chattian) and Lower Miocene (Upper Burdigalian) deposits of the Aquitaine Basin, south-west France. Epiphaxum is a poorly documented genus but its fossil record extends back to the Late Cretaceous; it was previously known only from the Paleocene (Danian). Epiphaxum arbuscula differs from all others species of the genus in the form of its colony. In contrast to the creeping colonies of previously known species, it has branched colonies. It is very common at one Upper Oligocene outcrop from which an assemblage with submarine cave remains has been described. A close relationship between the three extant species (two from the Caribbean Sea and one from the Indo-West Pacific region) and the Paleogene species is also noted. These constitute a group that has not undergone any important morphological changes for the last 65 million years.     

A comprehensive phylogeny of extinct and extant Rhizomyinae (Rodentia): evidence for multiple intercontinental dispersals

The subfamily Rhizomyinae is known from the Late Oligocene up to the present. Today this group comprises six species, which live in southern Asia and eastern Africa. Despite the current moderate diversity of the rhizomyines, they had a greater diversification and wider distribution in the past: from Asia, their land of origin, to Africa, which they entered during the Early Miocene. So far 33 fossil species can be referred to this group. A cladistic analysis involving fossil and living species has been carried out. Prokanisamys spp. turned out to be the most basal taxa of the ingroup. This analysis calls into question the monophyly of several genera, and allows the proposal of a phylogenetic definition of the tribes Tachyoryctini and Rhizomyini. It also provides information about the origin of the African rhizomyines and allows inferring multiple dispersal phenomena from Asia to Africa in Early and Late Miocene times.     

An assemblage of early Oligocene lizards (Squamata) from the locality of Boutersem (Belgium), with comments on the Eocene–Oligocene transition

A diverse, new lizard assemblage from the early Oligocene of Belgium is described. The Boutersem railway local fauna is the most species‐rich lizard assemblage yet reported from the European early Oligocene. Four lizard taxa are present: Lacertidae, Anguidae, Scincoidea and Platynota. One new species is described, Folisaurus boutersemensis sp. nov . This fauna provides new insight into the profound turnover that took place during the Eocene/Oligocene boundary in Europe. The new fauna confirms a marked decrease in diversity across the Eocene/Oligocene boundary. Two groups encountered in the European late Eocene became extinct (Iguanidae*, Glyptosaurinae). Estimates of species‐level extinctions range up to 80%. These estimates include members of virtually all the families present in the late Eocene. The relative importance of climate change and biotic interactions in controlling this pattern is discussed, and negative interactions between lizards and new carnivorous mammals are favoured. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 148–170.     

塔里木盆地吐克尔明剖面第三系苏维依组的介形类和轮藻化石

报道产自新疆天山南麓轮台县吐克尔明地区第三系苏维依组的５属１０种介形类化石和２属５种轮藻化石,并讲座其地质时代和上、下第三系的界线。