Pseudopus pannonicus (Squamata), the largest known anguid lizard-Redescription of the type material and new specimens from the Neogene and Quaternary of Hungary and Poland.

Herein, we revise the material of the extinct taxon Pseudopus pannonicus from Central Europe, the largest known anguid lizard and iconic member of herpetofaunas from the Upper Cenozoic of Europe. The geographical position of Polgárdi 2, the type locality of P. pannonicus, as well as several other closely located important localities make Central Europe a valuable area of high interest for studies regarding European Cenozoic palaeoherpetological assemblages. We clarified the nature of the type material of Pseudopus pannonicus, showing that it consisted not only of the five specimens originally figured. Instead, the syntype series also includes a considerable number of specimens from Polgárdi 2 that were only described or figured 12 years after the original description of P. pannonicus. Detailed osteological descriptions are provided for specimens from the type series and Polish specimens, with the aid of high-resolution imaging (SEM and µCT scanning), and intraspecific variability is discussed. The articular surface with the lappet of the parietal overlapping the frontal is discussed as a character potentially relevant for the diagnosis of P. pannonicus. We updated the identification of several specimens of P. pannonicus and discussed the biogeographic implications of such revisions. In Poland, P. pannonicus is an abundant component of Neogene and early Quaternary herpetofaunas, known with certainty from the Middle Miocene of Przeworno (the oldest known occurrence of the species globally), the Early-Late Pliocene of Weze I, the Late Pliocene of Rebielice Królewskie I, the Late Pliocene of Weze II, and the Early Pleistocene of Kadzielnia (one of the youngest occurrences documented globally). An indeterminate anguine with Anguis affinities is newly reported from Rebielice Królewskie II. The taxonomic status of other large anguids from the Neogene of Europe is discussed and we conclude that most are junior synonyms of P. pannonicus. We also show that another purported synonym, that is, Ophisaurus intermedius from the Early Pleistocene of Romania, is instead a nomen nudum. We conducted phylogenetic analyses (18 taxa, 65 characters) to understand the relationship of P. pannonicus relative to other anguid representatives and anguid-related group (i.e., glyptosaurids). A single most parsimonious tree (length: 134 steps) was recovered. The clade Pseudopus is stable, comprising the two distinct sister subclades [Pseudopus laurillardi + Pseudopus ahnikoviensis] and [Pseudopus pannonicus + Pseudopus apodus]. These phylogenetic results are in accordance with previously published works.

Combined uranium-series and electron spin resonance dating from the Pliocene fossil sites of Aves and Milo's palaeocaves, Bolt's Farm, Cradle of Humankind, South Africa.

Bolt's Farm is the name given to a series of non-hominin bearing fossil sites that have often been suggested to be some of the oldest Pliocene sites in the Cradle of Humankind, South Africa. This article reports the results of the first combined Uranium-Series and Electron Spin Resonance (US-ESR) dating of bovid teeth at Milo's Cave and Aves Cave at Bolt's Farm. Both tooth enamel fragments and tooth enamel powder ages were presented for comparison. US-ESR, EU and LU models are calculated. Overall, the powder ages are consistent with previous uranium-lead and palaeomagnetic age estimates for the Aves Cave deposit, which suggest an age between ~3.15 and 2.61 Ma and provide the first ages for Milo's Cave dates to between ~3.1 and 2.7 Ma. The final ages were not overly dependent on the models used (US-ESR, LU or EU), which all overlap within error. These ages are all consistent with the biochronological age estimate (<3.4->2.6 Ma) based on the occurrence of Stage I Metridiochoerus andrewsi. Preliminary palaeomagnetic analysis from Milo's Cave indicates a reversal takes place at the site with predominantly intermediate directions, suggesting the deposit may date to the period between ~3.03 and 3.11 Ma within error of the ESR ages. This further suggests that there are no definitive examples of palaeocave deposits at Bolt's Farm older than 3.2 Ma. This research indicates that US-ESR dating has the potential to date fossil sites in the Cradle of Humankind to over 3 Ma. However, bulk sample analysis for US-ESR dating is recommended for sites over 3 Ma.

Old World Fruit Bats (Pteropodidae, Chiroptera) from the Pleistocene of Vietnam.

Isolated teeth of a lesser short-nosed fruit bat Cynopterus brachyotis (Müller, 1838), a fulvous fruit bat Rousettus leschenaultii (Desmarest, 1820), and a dawn bat Eonycteris spelaea (Dobson, 1871) are described from the Middle Pleistocene Tham Hai cave locality in northern Vietnam (Lang Son Province). These are the first fossil findings of the Old World fruit bats in Vietnam. The Middle Pleistocene association of Pteropodidae from the Tham Hai locality may largely reflect the composition of species that roosted in local caves.

First automatic size measurements for the separation of dwarf birch and tree birch pollen in MIS 6 to MIS 1 records from Northern Germany.

During past glacial periods, the land cover of Northern Eurasia and North America repeatedly shifted between open steppe tundra and boreal/temperate forest. Tracking these changes and estimating the coverage of open versus forested vegetation in past glacial and interglacial landscapes is notoriously difficult because the characteristic dwarf birches of the tundra and the tree birches of the boreal and temperate forests produce similar pollen grains that are difficult to distinguish in the pollen record. One objective approach to separating dwarf birch pollen from tree birch pollen is to use grain size statistics. However, the required grain size measurements are time-consuming and, therefore, rarely produced. Here, we present an approach to automatic size measurement based on image recognition with convolutional neural networks and machine learning. It includes three main steps. First, the TOFSI algorithm is applied to detect and classify pollen, including birch pollen, in lake sediment samples. Second, a Resnet-18 neural network is applied to select the birch pollen suitable for measurement. Third, semantic segmentation is applied to detect the outline and the area and mean width of each detected birch pollen grain. Test applications with two pollen records from Northern Germany, one covering the Lateglacial-Early Holocene transition and the other covering the Mid to Late Pleistocene transition, show that the new technical approach is well suited to measure the area and mean width of birch pollen rapidly (>1000 per hour) and with high accuracy. Our new network-based tool facilitates more regular size measurements of birch pollen. Expanded analysis of modern birch pollen will help to better understand size variations in birch pollen between birch species and in response to environmental factors as well as differential sample preparation. Analysis of fossil samples will allow better quantification of dwarf birch versus tree birch in past environments.

Fauna and Landscapes of the Southern Minusinsk Basin (Southern Siberia) at the End of the Late Pleistocene.

Paleolandscapes of the first half of MIS 2 or the Last Glacial Maximum (LGM) were reconstructed based on the study of the first mammoth fauna locality found in the Abakan River valley and a comparison with other localities of the South Minusinsk Basin. Sediments of the Uytag geological section under study included silt interlayers underlain and overlain by sandstone rock slack fragments and were interpreted as weakly eroded diluvium. Studies of the Uytag locality made it possible to clarify the time and range of distribution in southern Siberia for several mammals (Ovis ammon, Equus ferus, Marmota baibacina, Sicista subtilis, Lagurus lagurus, and Microtus gregalis) and the Pleistocene bird Aquila chrysaetos. The age of the locality was confirmed by radiocarbon dating. The species composition of the Uytag fauna was similar to that of other localities of the same age and region. All known localities and single finds of faunal remains indicate that open steppe landscapes were widely developed in the South Minusinsk Basin during the LGM.

First record of well-preserved canid coprolites from Eurasia: New insights into the unique ecological niche of Yuanmou Basin.

This study reconstructs the Early Pleistocene paleoenvironment of the Yuanmou Basin through coproecology of the third member of the Yuanmou Formation. We examined 38 exceptionally well-preserved coprolites from a new fossil locality, and attributed the putative defecating agent to the hypercarnivorous diet canid, Sinocuon yuanmouensis through geochemical and quantitative analyses. A new ichnogenus and ichnospecies, Cuocopros yuanmouensis igen. et. isp. nov., was established based on distinctive characteristics. Multi-disciplinary analysis, including sediment palynology and lithostratigraphy, helped primarily reconstruct a significant climatic event during the early Pleistocene, coinciding with the emergence of Yuanmou Man during the fourth member of the Yuanmou Formation's deposition. The findings provide insights into coexistence between canids, hyaenas, hominoids, and other fauna, revealing a rich paleoecosystem and food chain in the region's history. This study contributes to understanding the complex ecological dynamics during this period in the Yuanmou Basin.

The role of Pleistocene dispersal in shaping species richness of sky island wintergreens from the Himalaya-Hengduan Mountains.

In addition to topography and climate, biogeographic dispersal has been considered to influence plant diversity in the Himalaya-Hengduan Mountains (HHM), yet, the mode and tempo of sky island dispersal and its influence on species richness has been little explored. Through phylogenetic analysis of Gaultheria ser. Trichophyllae, a sky island alpine clade within the HHM, we test the hypothesis that dispersal has affected current local species richness. We inferred the dynamics of biogeographic dispersal with correlation tests on direction, distance, occurrence time, and regional species richness. We found that G. ser. Trichophyllae originated at the end of the Miocene and mostly dispersed toward higher longitudes (eastward). In particular, shorter intra-regional eastward dispersals and longer inter-regional westward dispersals were most frequently observed. We detected a prevalence of eastward intra-region dispersals in both glacial periods and interglacials. These dispersals may have been facilitated by the reorganization of paleo-drainages and monsoon intensification through time. We suggest that the timing of dispersal corresponding to glacial periods and the prevalence of intra-region dispersal, rather than dispersal frequency, most influenced the pattern of species richness of G. ser. Trichophyllae. This study facilitates a more comprehensive understanding of biodiversity in the sky islands within the HHM.

Virtual reconstruction and geometric morphometric analysis of the Kocabas hominin fossil from Turkey: Implications for taxonomy and evolutionary significance.

The Kocabas specimen comes from a travertine quarry near the homonymous village in the Denizli basin (Turkey). The specimen comprises three main fragments: portions of the right and left parietal and left and right parts of the frontal bone. The fossil was assumed to belong to the Homo erectus s.l. hypodigm by some authors, whereas others see similarities with Middle Pleistocene fossils (Broken Hill 1/Kabwe, Bodo, or Ceprano). Here, we present the first attempt to make a complete reconstruction of the missing medial portion of the frontal bone and a comprehensive geometric morphometric analysis of this bone. We restored the calotte by aligning and mirroring the three preserved fragments. Afterward, we restored the missing portion by applying the thin-plate spline interpolation algorithm of target fossils onto the reconstructed Kocabas specimen. For the geometric morphometric analyses, we collected 80 landmarks on the frontal bone (11 osteometric points, 14 bilateral curve semilandmarks, and 41 surface semilandmarks). The comparative sample includes 21 fossils from different chronological periods and geographical areas and 30 adult modern humans from different populations. Shape analyses highlighted the presence in Kocabas of features usually related to Middle Pleistocene Homo, such as a developed supraorbital torus associated with a relatively short frontal squama and reduced post-toral sulcus. Cluster analysis and linear discriminant analysis classification procedure suggest Kocabas being part of the same taxonomic unit of Eurasian and African Middle Pleistocene Homo. In light of our results, we consider that attributing the Kocabas hominin to H. erectus s.l. may be unwarranted. Results of our analyses are compatible with different evolutionary scenarios, but a more precise chronological framework is needed for a thorough discussion of the evolutionary significance of this specimen. Future work should clarify its geological age, given uncertainties regarding its stratigraphic provenance.

Camelus knoblochi genome reveals the complex evolutionary history of Old World camels.

Extant Old World camels (genus Camelus) contributed to the economic and cultural exchanges between the East and West for thousands of years.1,2 Although many remains have been unearthed,3,4,5 we know neither whether the prevalent hybridization observed between extant Camelus species2,6,7 also occurred between extinct lineages and the ancestors of extant Camelus species nor why some populations became extinct while others survived. To investigate these questions, we generated paleogenomic and stable isotope data from an extinct two-humped camel species, Camelus knoblochi. We find that in the mitochondrial phylogeny, all C. knoblochi form a paraphyletic group that nests within the diversity of modern, wild two-humped camels (Camelus ferus). In contrast, they are clearly distinguished from both wild and domesticated (Camelus bactrianus) two-humped camels on the nuclear level. Moreover, the divergence pattern of the three camel species approximates a trifurcation, because the most common topology is only slightly more frequent than the two other possible topologies. This mito-nuclear phylogenetic discordance likely arose due to interspecific gene flow between all three species, suggesting that interspecific hybridization is not exclusive to modern camels but a recurrent phenomenon throughout the evolutionary history of the genus Camelus. These results suggest that the genomic complexity of Old World camels' evolutionary history is underestimated when considering data from only modern species. Finally, we find that C. knoblochi populations began declining prior to the last glacial maximum and, by integrating palaeoecological evidence and stable isotope data, suggest that this was likely due to failure to adapt to a changing environment.

Phylogeographic Pattern of the Assassin Bug Sycanus bifidus Inferred from Mitochondrial Genomes and Nuclear Genes.

The assassin bug Sycanus bifidus has a wide distribution across southern China. This study explored its distribution and evolution by analyzing mitochondrial and nuclear ribosomal RNA genes, revealing how Pleistocene climate and geological changes shaped its phylogeography. We identified two main clades, A and B, that diverged in the Middle Pleistocene. Hainan Island's populations form a unique group within Clade A, suggesting that the Qiongzhou Strait served as a dispersal corridor during glaciation. Rising sea levels likely separated the Hainan population afterward. Ecological niche modeling showed that both populations have been viable since the last interglacial period, with demographic analyses indicating possible expansions during the Middle and Late Pleistocene, driven by favorable climates. This study highlights the significant effects of Pleistocene sea-level and climatic changes on the distribution and evolution of S. bifidus in China.

An extinct north American porcupine with a South American tail.

New World porcupines (Erethizontinae) originated in South America and dispersed into North America as part of the Great American Biotic Interchange (GABI) 3-4 million years ago.1 Extant prehensile-tailed porcupines (Coendou) today live in tropical forests of Central and South America.2,3 In contrast, North American porcupines (Erethizon dorsatum) are thought to be ecologically adapted to higher-latitude temperate forests, with a larger body, shorter tail, and diet that includes bark.4,5,6,7 Limited fossils8,9,10,11,12,13 have hindered our understanding of the timing of this ecological differentiation relative to intercontinental dispersal during the GABI and expansion into temperate habitats.14,15,16,17,18 Here, we describe functionally important features of the skeleton of the extinct Erethizon poyeri, the oldest nearly complete porcupine skeleton documented from North America, found in the early Pleistocene of Florida. It differs from extant E. dorsatum in having a long, prehensile tail, grasping foot, and lacking dental specializations for bark gnawing, similar to tropical Coendou. Results from phylogenetic analysis suggest that the more arboreal characteristics found in E. poyeri are ancestral for erethizontines. Only after it expanded into temperate, Nearctic habitats did Erethizon acquire the characteristic features that it is known for today. When combined with molecular estimates of divergence times, results suggest that Erethizon was ecologically similar to a larger species of Coendou when it crossed the Isthmus of Panama by the early Pleistocene. It is likely that the range of this more tropically adapted form was limited to a continuous forested biome that extended from South America through the Gulf Coast.

How the Sima de los Huesos was won.

Although the first discovery of a human fossil in the Sima de los Huesos took place in 1976, systematic excavations did not begin there until 1984. Since then, this site has been continuously excavated in month-long camps. The site is dated by different radiometric techniques to between 430,000 and 300,000 years ago. Until the 2023 campaign, just over 7000 human fossils have been recovered, constituting the largest collection of fossils prior to Homo sapiens ever discovered. The fossils correspond to a minimum of 29 individuals of both sexes and different ages at death, from preadolescents to a specimen of advanced age. Comparative anatomy and ancient DNA studies both suggest that this is a population closely related to Homo neanderthalensis. The great variety and extraordinary quality of the fossils recovered have allowed us to carry out a series of investigations that have greatly increased our knowledge about the evolution of Homo in the Middle Pleistocene. Among the most important discoveries, it has been possible to establish body size and proportions, the confirmation that the origin of the accumulation of human fossils was of an anthropic nature, that those past humans took care of disabled individuals and who were capable of having an oral language almost as complex and efficient as that of our own species.

Plecotus macrobullaris sarmaticus subsp. nov. (Vespertilionidae, Chiroptera) from the Early Pleistocene of Crimea.

The extinct mountain long-eared bat Plecotus macrobullaris sarmaticus subsp. nov. is described on the base of jaw remains from the Lower Pleistocene of the Taurida cave in central Crimea. This is the first record of the genus Plecotus in the Lower Pleistocene of Russia and the first fossil find of P. macrobullaris Kuzyakin, 1965, documenting the early stage of the evolutionary history of the species.

Vrba was right: Historical climate fragmentation, and not current climate, explains mammal biogeography.

Climate plays a crucial role in shaping species distribution and evolution over time. Dr Vrba's Resource-Use hypothesis posited that zones at the extremes of temperature and precipitation conditions should host a greater number of climate specialist species than other zones because of higher historical fragmentation. Here, we tested this hypothesis by examining climate-induced fragmentation over the past 5 million years. Our findings revealed that, as stated by Vrba, the number of climate specialist species increases with historical regional climate fragmentation, whereas climate generalist species richness decreases. This relationship is approximately 40% stronger than the correlation between current climate and species richness for climate specialist species and 77% stronger for generalist species. These evidences suggest that the effect of climate historical fragmentation is more significant than that of current climate conditions in explaining mammal biogeography. These results provide empirical support for the role of historical climate fragmentation and physiography in shaping the distribution and evolution of life on Earth.

The phylogenetic relationship and demographic history of rhesus macaques (Macaca mulatta) in subtropical and temperate regions, China.

Pleistocene climatic oscillations exerted significant influences on the genetic structure and demography of rhesus macaque (Macaca mulatta) in eastern China. However, the evolutionary history of rhesus macaques in subtropical and temperate China remained unclear and/or controversial. Herein, we analyzed the autosomes, mitochondrial genomes, and Y-chromosomes from 84 individuals of Chinese rhesus macaque. The results revealed that (1) all individuals were clustered into pan-west and pan-east genetic groups, which exhibited Shaanxi Province as the northernmost region of western dispersal route of rhesus macaques in China; (2) in subtropical and temperate China, rhesus macaques were divided into four lineages (TH, DB, HS, and QL), and their divergence times corresponded to the Penultimate Glaciation (300-130 kya) and Last Glaciation (70-10 kya), respectively; (3) the individuals from Mt. Taihangshan (TH) are closely related to individuals from Mt. Dabashan (DB) in the autosomal tree, rather than individuals from Mt. Huangshan (HS) as indicated by the mitogenome tree, which supports the hypothesis that the ancestral rhesus macaques radiated into Mt. Taihangshan from Mt. Huangshan via Mt. Dabashan; and (4) the demographic scenario of the four lineages showed the ancestral rhesus macaques bottleneck and expansion corresponding to the suitable habitat reduction and expansion, which confirmed they had experienced northward recolonization and southward retreat events from Mt. Huangshan area via Northern China Plain to Northernmost China along with Pleistocene glacial cycles. This study provides a new insight into understanding how Pleistocene glaciation has influenced faunal diversity in subtropical and temperate China, especially for those exhibiting differential patterns of sex dispersal.

The scimitar-cat Homotherium from the submerged continental shelf of the Gulf Coast of Texas.

The machairodontine felid Homotherium achieved a global geographic distribution throughout much of the Pleistocene. Accordingly, that large carnivore is important for understanding patterns of community composition. We report on a new record of Homotherium based on a fragmentary premaxilla-maxilla discovered on McFaddin Beach, Texas, along the Gulf of Mexico. Skeletal remains of extinct, Pleistocene vertebrates accumulate on McFaddin Beach. Those fossils appear to originate from submerged deposits on the continental shelf in the Gulf of Mexico, an area that was subaerially exposed in the Late Pleistocene during glacial intervals. Marine erosion and transport altered the externally visible morphology of the current specimen, obscuring and/or damaging taxonomically informative details of the preserved dentition. However, high-resolution X-ray computed tomography revealed diagnostic portions of the unerupted crown of an upper canine within its alveolus. The serrated edges of the canine combined with the position of the incisors demonstrate that the specimen from McFaddin Beach represents a species of Homotherium. That specimen is the latest in a larger sample of Homotherium in Texas that spans most of the Pliocene-Pleistocene. This is the first occurrence of Homotherium from the continental shelf of the Gulf Coast. That landscape may have formed a broad subtropical Gulf Coast corridor that facilitated the dispersal of Neotropical taxa along the coast between Texas and Florida. The associated fauna from McFaddin Beach contains Neotropical mammals common to southern Texas and Florida and indicates that Homotherium was a member of the fauna inhabiting the Gulf Coast corridor during the Late Pleistocene.

New Oldowan locality Sare-Abururu (ca. 1.7 Ma) provides evidence of diverse hominin behaviors on the Homa Peninsula, Kenya.

The Homa Peninsula, in southwestern Kenya, continues to yield insights into Oldowan hominin landscape behaviors. The Late Pliocene locality of Nyayanga (∼3-2.6 Ma) preserves some of the oldest Oldowan tools. At the Early Pleistocene locality of Kanjera South (∼2 Ma) toolmakers procured a diversity of raw materials from over 10 km away and strategically reduced them in a grassland-dominated ecosystem. Here, we report findings from Sare-Abururu, a younger (∼1.7 Ma) Oldowan locality approximately 12 km southeast of Kanjera South and 18 km east of Nyayanga. Sare-Abururu has yielded 1754 artifacts in relatively undisturbed low-energy silts and sands. Stable isotopic analysis of pedogenic carbonates suggests that hominin activities were carried out in a grassland-dominated setting with similar vegetation structure as documented at Kanjera South. The composition of a nearby paleo-conglomerate indicates that high-quality stone raw materials were locally abundant. Toolmakers at Sare-Abururu produced angular fragments from quartz pebbles, representing a considerable contrast to the strategies used to reduce high quality raw materials at Kanjera South. Although lithic reduction at Sare-Abururu was technologically simple, toolmakers proficiently produced cutting edges, made few mistakes and exhibited a mastery of platform management, demonstrating that expedient technical strategies do not necessarily indicate a lack of skill or suitable raw materials. Lithic procurement and reduction patterns on the Homa Peninsula appear to reflect variation in local resource contexts rather than large-scale evolutionary changes in mobility, energy budget, or toolmaker cognition.

Hominin population bottleneck coincided with migration from Africa during the Early Pleistocene ice age transition.

Two recently published analyses make cases for severe bottlenecking of human populations occurring in the late Early Pleistocene, one case at about 0.9 Mya based on a genomic analysis of modern human populations and the low number of hominin sites of this age in Africa and the other at about 1.1 Mya based on an age inventory of sites of hominin presence in Eurasia. Both models point to climate change as the bottleneck trigger, albeit manifested at very different times, and have implications for human migrations as a mechanism to elude extinction at bottlenecking. Here, we assess the climatic and chronologic components of these models and suggest that the several hundred-thousand-year difference is largely an artifact of biases in the chronostratigraphic record of Eurasian hominin sites. We suggest that the best available data are consistent with the Galerian hypothesis expanded from Europe to Eurasia as a major migration pulse of fauna including hominins in the late Early Pleistocene as a consequence of the opening of land routes from Africa facilitated by a large sea level drop associated with the first major ice age of the Pleistocene and concurrent with widespread aridity across Africa that occurred during marine isotope stage 22 at ~0.9 Mya. This timing agrees with the independently dated bottleneck from genomic analysis of modern human populations and allows speculations about the relative roles of climate forcing on the survival of hominins.

Quantifying hominin morphological diversity at the end of the middle Pleistocene: Implications for the origin of Homo sapiens.

OBJECTIVES: The Middle Pleistocene (MP) saw the emergence of new species of hominins: Homo sapiens in Africa, H. neanderthalensis, and possibly Denisovans in Eurasia, whose most recent common ancestor is thought to have lived in Africa around 600 ka ago. However, hominin remains from this period present a wide range of morphological variation making it difficult to securely determine their taxonomic attribution and their phylogenetic position within the Homo genus. This study proposes to reconsider the phenetic relationships between MP hominin fossils in order to clarify evolutionary trends and contacts between the populations they represent. MATERIALS AND METHODS: We used a Geometric Morphometrics approach to quantify the morphological variation of the calvarium of controversial MP specimens from Africa and Eurasia by using a comparative sample that can be divided into 5 groups: H. ergaster, H. erectus, H. neanderthalensis, and H. sapiens, as well as individuals from current modern human populations. We performed a Generalized Procrustes Analysis, a Principal Component Analysis, and Multinomial Principal Component Logistic Regressions to determine the phenetic affinities of the controversial Middle Pleistocene specimens with the other groups. RESULTS: MP African and Eurasian specimens represent several populations, some of which show strong affinities with H. neanderthalensis in Europe or H. sapiens in Africa, others presenting multiple affinities. DISCUSSION: These MP populations might have contributed to the emergence of these two species in different proportions. This study proposes a new framework for the human evolutionary history during the MP.