Ancient genomes reveal origin and rapid trans-Eurasian migration of 7th century Avar elites.

The Avars settled the Carpathian Basin in 567/68 CE, establishing an empire lasting over 200 years. Who they were and where they came from is highly debated. Contemporaries have disagreed about whether they were, as they claimed, the direct successors of the Mongolian Steppe Rouran empire that was destroyed by the Turks in ∼550 CE. Here, we analyze new genome-wide data from 66 pre-Avar and Avar-period Carpathian Basin individuals, including the 8 richest Avar-period burials and further elite sites from Avar's empire core region. Our results provide support for a rapid long-distance trans-Eurasian migration of Avar-period elites. These individuals carried Northeast Asian ancestry matching the profile of preceding Mongolian Steppe populations, particularly a genome available from the Rouran period. Some of the later elite individuals carried an additional non-local ancestry component broadly matching the steppe, which could point to a later migration or reflect greater genetic diversity within the initial migrant population.

Alkaline soda Lake Velika Rusanda (Serbia): the first insight into diatom diversity of this extreme saline lake.

Alkaline soda lakes are unique habitats found in specific geographic regions, usually with dry climate. The Carpathian Basin is one of those regions very important for habitat and biodiversity conservation in Europe, with natural soda lakes found in Austria, Hungary and Serbia. In comparison to other two countries from Central Europe, algal biodiversity studies of saline soda lakes in Serbia are scarce. Lake Velika Rusanda has the highest measured salinity of all saline lakes in the Carpathian Basin and there were no reports of its diatom species richness and diversity till now. We conducted 2-year investigation programme to study biodiversity and seasonal dynamics of diatoms in this lake. A total of 27 diatom taxa were found, almost all of them attached to reed and much less in benthos and plankton. Five new diatom species for Serbia were recorded, Craticula halopannonica, Navicymbula pusilla, Hantzschia weyprechtii, Nitzschia thermaloides and Navicula staffordiae. The last mentioned is new for Europe as well. Lake Velika Rusanda is inhabited mostly by alkaliphilous and halophilic diatoms. Since diatoms are used as bioindicators in soda lakes, our results will improve their further application in ecological status assessment of these fragile habitats in the Carpathian Basin.

Tracing the genetic origin of Europe's first farmers reveals insights into their social organization.

Farming was established in Central Europe by the Linearbandkeramik culture (LBK), a well-investigated archaeological horizon, which emerged in the Carpathian Basin, in today's Hungary. However, the genetic background of the LBK genesis is yet unclear. Here we present 9 Y chromosomal and 84 mitochondrial DNA profiles from Mesolithic, Neolithic Starcevo and LBK sites (seventh/sixth millennia BC) from the Carpathian Basin and southeastern Europe. We detect genetic continuity of both maternal and paternal elements during the initial spread of agriculture, and confirm the substantial genetic impact of early southeastern European and Carpathian Basin farming cultures on Central European populations of the sixth-fourth millennia BC. Comprehensive Y chromosomal and mitochondrial DNA population genetic analyses demonstrate a clear affinity of the early farmers to the modern Near East and Caucasus, tracing the expansion from that region through southeastern Europe and the Carpathian Basin into Central Europe. However, our results also reveal contrasting patterns for male and female genetic diversity in the European Neolithic, suggesting a system of patrilineal descent and patrilocal residential rules among the early farmers.

The first probable case with tuberculous meningitis from the Hun period of the Carpathian Basin - How diagnostics development can contribute to increase knowledge and understanding of the spatio-temporal distribution of tuberculosis in the past.

The aim of our paper is to demonstrate and discuss in detail the endocranial bony changes suggestive of tuberculous meningitis (TBM) that were recorded in an adult female's (SPF15) skeleton. The bone remains were uncovered from a solitary grave from the Hun period (5th-century-CE) archaeological site of Solt-Polya-fok (Bács-Kiskun county, Hungary). During the macromorphological examination of the very incomplete and poorly preserved skeleton of SPF15, the inner surface of the skull displayed abnormally pronounced digital impressions (APDIs) and granular impressions (GIs). Recently, it was confirmed that endocranial GIs can be considered as specific signs of TBM; and thus, they are sufficient enough on their own to make a definitive diagnosis of the disease in the palaeopathological practice. On the other hand, APDIs are not specific to TBM but can be tuberculous in origin; their concomitant presence with GIs in SPF15 makes their tuberculous origin very likely. Based on the above, it seems that the adult female from the 5th-century-CE archaeological site of Solt-Polya-fok suffered from TBM. SPF15 is the first reported probable case with tuberculosis (TB) from the Hun period of the present-day territory of Hungary, who gives us invaluable information about the spatio-temporal distribution of the disease in the past. Furthermore, it highlights the paramount importance of diagnostics development, especially the identification and refinement of diagnostic criteria, as without the application of APDIs and GIs, the diagnosis of TB could not have been established in SPF15.

Repeated stand structure inventory dataset in long abandoned deciduous forest reserves in Hungary.

Deeper understanding on natural forest dynamics requires long-term data series from forests that have not been affected by human interventions, which are often scarce especially in the Pannonian Bioregion. Unmanaged, but regularly inventoried forest reserves provide an opportunity to fill this gap. The dataset provides repeated inventory data for 233 permanent plots situated in the core areas of six forest reserves selected from primary forests (Kékes), long abandoned forests (Kecskés-galya, Szalafo, Várhegy) and abandoned ones (Hidegvíz-völgy, Nagy Istrázsa-hegy). The sampled old stands represent the four most widespread hilly forest types in Hungary: Carpathian submountainous beech forest; sessile oak-hornbeam forest; Turkey oak and sessile oak forest; downy oak forest. In each plot, stand level attributes included main mensuration variables (canopy closure, stand height, tree density, basal area, living and dead volume, lying deadwood and admixture of the main tree species). Tree level attributes (diameter at breast height, height measured and estimated, crown position in the canopy, health status, tree history of all trees or shrubs having diameter larger or equal to 5 cm) were also measured in two inventories (after 6-16 years) for a total of 6,986 individual trees sampled in all plots. Fagus sylvatica L., Quercus petraea agg., Q. cerris L., Q. pubescens Willd., Carpinus betulus L., Acer campestre L. and Cornus mas L. were the most abundant. The individual tree history classification refers to regeneration ingrowth, growing phase, mortality, decaying phase and disappearance events, that can be used for calculation of various stand dynamics attributes. The dataset offers valuable opportunities for quantifying changes in stand structures and tree population dynamic attributes after the abandonment of management. Inventory data can be integrated with environmental and climatic information to understand the drivers of forest stand dynamics under a changing climate.

Genomic analysis reveals complex population structure within the smooth newt, Lissotriton vulgaris, in Central Europe.

Species with wide-range distributions usually display high genetic variation. This variation can be partly explained by historical lineages that were temporally isolated from each other and are back into secondary reproductive contact, and partly by local adaptations. The smooth newt (Lissotriton vulgaris) is one of the most widely distributed amphibians species across Eurasia and forms a species complex with a partially overlapping distribution and morphology. In the present study, we explored the population genomic structure of smooth newt lineages in the Carpathian Basin (CB) relying on single-nucleotide polymorphisms. Our dataset included new and previously published data to study the secondary contact zone between lineages in the CB and also tested for the barrier effect of rivers to gene flow between these lineages. We confirmed the presence of the South L. v. vulgaris Lineage distributed in Transdanubia and we provided new distribution records of L. v. ampelensis inhabiting the eastern territories of the CB. High genetic diversity of smooth newts was observed, especially in the North Hungarian Mountains and at the interfluves of the main rivers in the South with four distinct lineages of L. v. vulgaris and one lineage of L. v. ampelensis showing a low level of admixture with the spatially closest L. v. vulgaris lineage. Moreover, admixture detected at the interfluve of the main rivers (i.e. Danube and Tisza) suggested a secondary contact zone in the area. Finally, we found that the river Danube has a very weak effect on population divergence, while the river Tisza is a geographical barrier limiting gene flow between smooth newt lineages. As the range boundaries of L. v. vulgaris and L. v. ampelensis in the CB coincide with the river Tisza, our study underpins the influence of rivers in lineage diversification.

Is there a massive glacial-Holocene flora continuity in Central Europe?

The prevailing paradigm about the Quaternary ecological and evolutionary history of Central European ecosystems is that they were repeatedly impoverished by regional extinctions of most species during the glacial periods, followed by massive recolonizations from southern and eastern refugia during interglacial periods. Recent literature partially contradicts this view and provides evidence to re-evaluate this Postglacial Recolonization Hypothesis and develop an alternative one. We examined the long-term history of the flora of the Carpathian (Pannonian) Basin by synthesising recent advances in ecological, phylogeographical, palaeoecological and palaeoclimatological research, and analysing the cold tolerance of the native flora of a test area (Hungary, the central part of the Carpathian Basin). We found that (1) many species have likely occurred there continuously since before the Last Glacial Maximum (LGM); (2) most of the present-day native flora (1404 species, about 80%) can occur in climates as cold as or colder than the LGM (mean annual temperature ≤+3.5°C); and (3) grasslands and forests can be species-rich under an LGM-like cold climate. These arguments support an alternative hypothesis, which we call the Flora Continuity Hypothesis. It states that long-term continuity of much of the flora in the Carpathian Basin is more plausible than regional extinctions during the LGM followed by massive postglacial recolonizations. The long-term continuity of the region's flora may have fundamental implications not only for understanding local biogeography and ecology (e.g. the temporal scale of processes), but also for conservation strategies focusing on protecting ancient species-rich ecosystems and local gene pools.

Conservation status of the blind mole rat populations in Hungary (Rodentia: Spalacinae: Nannospalax) revisited.

Regular reviews of long-term research and conservation programs are useful sources of information for future directions in science and for the assessment of current conservation status of taxa. In this paper, we compiled all available data from the last 10 years related to Hungarian blind mole rat populations and assessed this information according to the following main themes: systematics, distribution and threats, and conservation actions. Based on the most recent information, national and global risk assessments are provided for the three species of Nannospalax (N. montanosyrmiensis, N. hungaricus and N. syrmiensis), currently accepted as part of the Hungarian fauna.

The mixed phylogenetic origin of northern pike (Esox lucius Linnaeus 1758) populations in the Middle Danubian drainage.

BACKGROUND: Pikes, members of genus Esox, are widespread freshwater predators of the northern hemisphere, and important sport fish also. From the Carpathian basin only one species, the northern pike (E. lucius) is noted. At the same time the pike stocks living in this area show high level of phenotypic variance (e.g. various body pattern) and its growth varies highly both among and within populations. These features usually explained by the environmental diversity of the area. Whereas we think that genetic reasons -e.g. the appearance of other/new pike species in the area- may also be responsible for these observed features. Since as no detailed information have been published from the pike populations of this area, so far; we conducted phylogenetic and morphological assay on 88 pike specimens, collected from 49 Middle Danubian sampling sites. RESULTS: Our phylogenetic surveys showed that the northern pike appear in the study area solely, but all the three of its major lineages (Northern, Circumpolar, Southern) were indicated. Only six specimens represent the Northern lineage, collected from the western part of the study area. The Circumpolar and Southern lineages were common in the Carpathian basin, but the Southern lineage showed higher levels of haplotype diversity than the Circumpolar clade. Which indicates that only the Southern lineage is native in the area, while the other two groups could have appeared in the Middle Danubian system either spontaneously or by human introduction. Moreover, the different clades appeared in the same populations, suggesting the opportunity of inter-lineage hybridisation. From the studied morphologicalal features, the number of scales on the lateral line and the head length showed significant differences among the lineages. At the same time the body pattern of the studied individuals seems to be rather influenced by the ontogenic changes than phylogeny. CONCLUSIONS: The high phenotypic variability of Middle Danubian northern pike populations may be due that all of its three major clades appeared and came in secondary contact in the area. In the within watershed spread of the non-native lineages the human stocking/transfer may play a considerable role.

The genetic origin of Huns, Avars, and conquering Hungarians.

Huns, Avars, and conquering Hungarians were migration-period nomadic tribal confederations that arrived in three successive waves in the Carpathian Basin between the 5th and 9th centuries. Based on the historical data, each of these groups are thought to have arrived from Asia, although their exact origin and relation to other ancient and modern populations have been debated. Recently, hundreds of ancient genomes were analyzed from Central Asia, Mongolia, and China, from which we aimed to identify putative source populations for the above-mentioned groups. In this study, we have sequenced 9 Hun, 143 Avar, and 113 Hungarian conquest period samples and identified three core populations, representing immigrants from each period with no recent European ancestry. Our results reveal that this "immigrant core" of both Huns and Avars likely originated in present day Mongolia, and their origin can be traced back to Xiongnus (Asian Huns), as suggested by several historians. On the other hand, the "immigrant core" of the conquering Hungarians derived from an earlier admixture of Mansis, early Sarmatians, and descendants of late Xiongnus. We have also shown that a common "proto-Ugric" gene pool appeared in the Bronze Age from the admixture of Mezhovskaya and Nganasan people, supporting genetic and linguistic data. In addition, we detected shared Hun-related ancestry in numerous Avar and Hungarian conquest period genetic outliers, indicating a genetic link between these successive nomadic groups. Aside from the immigrant core groups, we identified that the majority of the individuals from each period were local residents harboring "native European" ancestry.

Historical fog climate dataset for Carpathian Basin from 1886 to 1919.

This paper presents the historical fog climate dataset from 1886 to 1919 for Hungary and its neighbouring countries in the Carpathian Basin. The dataset was obtained from the yearbooks of the Royal Hungarian Central Institute of Meteorology and Earth Magnetism (RHCIMEM) established in 1870 to investigate the climatic features of Hungary during the time of the Austro-Hungarian Monarchy. Monthly observations were recorded from 1871 and daily observations were recorded from 1886. The yearbooks contain daily meteorological records of temperature, relative humidity, rainfall, pressure, wind speed and direction, cloudiness and surface weather conditions along with monthly summaries for 24 meteorological stations. The daily weather observations were recorded three times a day, namely at 07:00, 14:00 and 21:00 local time. Station information (location, environment, instrumentation, observations etc.) can also be found in the yearbooks as metadata. For example, the definition of fog in the case of historical observations is the same as that of today, i.e., fog is detected if the maximum horizontal visibility is less than 1 km. In this way fog observations are easily comparable to today's observations without requiring further data correction and homogenisation. The longest 13 continuously recorded fog observation datasets have the length between 15 and 34 years. The stations are located in 5 countries of the Carpathian Basin at present. These datastests are suitable for conducting historical climatic investigations and can also serve as reference datasets. The historical dataset can be used to study the annual and seasonal changes in frequency and duration of fog events in the Carpathian Basin as a reference, thus facilitating research in the field of fog climatology and forecast.

Maternal Lineages from 10-11th Century Commoner Cemeteries of the Carpathian Basin.

Nomadic groups of conquering Hungarians played a predominant role in Hungarian prehistory, but genetic data are available only from the immigrant elite strata. Most of the 10-11th century remains in the Carpathian Basin belong to common people, whose origin and relation to the immigrant elite have been widely debated. Mitogenome sequences were obtained from 202 individuals with next generation sequencing combined with hybridization capture. Median joining networks were used for phylogenetic analysis. The commoner population was compared to 87 ancient Eurasian populations with sequence-based (Fst) and haplogroup-based population genetic methods. The haplogroup composition of the commoner population markedly differs from that of the elite, and, in contrast to the elite, commoners cluster with European populations. Alongside this, detectable sub-haplogroup sharing indicates admixture between the elite and the commoners. The majority of the 10-11th century commoners most likely represent local populations of the Carpathian Basin, which admixed with the eastern immigrant groups (which included conquering Hungarians).

Increased spatial resolution of sampling in the Carpathian basin helps to understand the phylogeny of central European stream-dwelling gudgeons.

BACKGROUND: Phylogenetic studies of widespread European fish species often do not completely cover their entire distribution area, and some areas are often excluded from analyses than others. For example, Carpathian stocks are often omitted from these surveys or are under-represented in the samples. However, this area served as an extra-Mediterranean refugia for many species; therefore, it is assumed that fish stocks here may show special phylogenetic features. For this reason, increased spatial resolution of sampling, namely revealing genetic information from unexamined Carpathian areas within the range of doubtful taxa, may help us better understand their phylogenetic features. To test this hypothesis, a phylogenetic investigation using a partial mtCR sequence data was conducted on 56 stream-dwelling freshwater fish (Gobio spp.) individuals collected from 11 rivers of the data-deficient Southeastern Carpathian area. Moreover, we revieved the available phylogenetic data of Middle-Danubian stream-dwelling gudgeon lineages to delineate their distribution in the area. RESULTS: Seven out of the nine detected haplotypes were newly described, suggesting the studied area hosts distinct and diverse Gobio stocks. Two valid species (G. obtusirostris, G. gobio), and a haplogroup with doubtful phylogenetic position" G. sp. 1" were detected in the area, showing a specific spatial distribution pattern. The distribution of the detected lineages in the Middle-Danubian area correspond with recent and paleo hydrogeographic features, at the same time mainly on their bordering areas show considerable overlap. CONCLUSIONS: Despite the relatively limited geographic range of the study, our results provide important information which contributes to a better understanding of the phylogenetic, taxonomic and distribution features of Central European gudgeons. The genetically confirmed distribution data of the indicated lineages corresponds well with the recent and near-recent hydrogeographic features of the area, and assumes several hybrid zones in the Carpathian Basin. Additionally, the results show that the middle and lower Danubian watershed cannot be excluded from the range of G. gobio. Moreover, the" G. sp. 1", is slightly differentiated but phylogenetically distinct entity, and is the only Gobio taxa thus far detected in the middle and lower Tisza-basin. However, further investigations are necessary to clarify the taxonomic position of this group.

Semiochemical baited traps of lepidopteran pests of economic importance can deliver reliable data also on wide range of non-target species: case study in the Hajdúság Region of East Pannonian Lowland (East Hungary).

Lepidopteran assemblages were studied at 16 sampling sites in the Hajdúság Region between 2013 and 2020. Although studies targeted development of synthetic phenylacetaldehyde-based and semi-synthetic isoamyl alcohol-based baits for pest monitoring, traps caught 179 species belonging to the Sphingidae, Thyatiridae, Geometridae, Erebidae and Noctuidaefamilies. Most species were pests or widely distributed generalists, but there were also many rare habitat specialists, for example, silvicol species, whose appearance was unexpected in the recently less forested region. The specificity of the two bait types tested differed notably both on family and subfamily levels. Semi-synthetic baits performed better and attracted a wide range of noctuids belonging mainly to the Xyleninae and Noctuinae subfamilies, while synthetic phenylacetaldehyde-based lures showed specificity to Plusiinae subfamilies with lower number of sampled species. Our data fill a gap of knowledge since the fauna studied formerly was nearly unknown and brings attention to the alternative use of volatile traps of agricultural pests in faunistical studies.

Population Genetic Structure of the Wild Boar (Sus scrofa) in the Carpathian Basin.

In the Carpathian Basin the wild boar (Sus scrofa) belongs among the most important game species both ecologically and economically, therefore knowing more about the basics of the genetics of the species is a key factor for accurate and sustainable management of its population. The aim of this study was to estimate the genetic diversity and to elucidate the genetic structure and location of wild boar populations in the Carpathian Basin. A total of 486 samples were collected and genotyped using 13 STR markers. The number of alleles varied between 4 and 14, at 9 of the 13 loci the observed heterozygosity was significantly different (p < 0.05) from the expected value, showing remarkable introgression in the population. The population was separated into two groups, with an Fst value of 0.03, suggesting the presence of two subpopulations. The first group included 147 individuals from the north-eastern part of Hungary, whereas the second group included 339 samples collected west and south of the first group. The two subpopulations' genetic indices are roughly similar. The lack of physical barriers between the two groups indicates that the genetic difference is most likely caused by the high reproduction rate and large home range of the wild boars, or by some genetic traces' having been preserved from both the last ice age and the period before the Hungarian water regulation.

Changing nature of Saharan dust deposition in the Carpathian Basin (Central Europe): 40 years of identified North African dust events (1979-2018).

Several billion tonnes of mineral dust is emitted, and transported through winds every year from arid-semiarid areas. North African dust hot spots located in the Sahara are responsible for 50-70% of the global mineral dust budget. Dust-loaded air-masses originated from these sources can be transported over long distances and can also affect remote areas, such as North and South Americas, Europe, and the Middle East. In this study, we analysed 218 identified Saharan dust events (SDEs) in the Carpathian Basin (Central Europe) during 1979 to 2018. Systematic identification of SDEs and analyses of dust emission, dust source area activity, dust transporting wind systems, and transport routes revealed that different synoptic meteorological patterns are responsible for SDEs, and these are occurring mostly in spring and summer. The characteristic synoptic meteorological background of episodes was also identified, and three major types of atmospheric pressure-system patterns were distinguished. In recent years, several intense wintertime dust deposition events have been recorded in Central Europe. All of the identified unusual episodes were characterised by severe washout of mineral dust material and were related to very similar synoptic meteorological situations. Enhanced southward propagation of a high-latitude upper-level atmospheric trough to north-western Africa and orographic blocking of Atlas Mountains played an essential role in the formation of severe dust storms, whereas the long-range transport was associated with the northward branch of the meandering jet. The occurrence and southerly penetration of high-latitude upper-level atmospheric trough to low-latitudes and the increased meridionality of the dominant flow patterns may be associated with enhanced warming of the Arctic, leading to more meandering jet streams. Particles size of sampled dust material of some intense deposition episodes were very coarse with a considerable volumetric proportion of > 20 µm particles.

Identification of microplastics in fish ponds and natural freshwater environments of the Carpathian basin, Europe.

In the past few years, there has been a significant development in freshwater microplastic research. Pollution has been detected in lakes and rivers of several continents, but the number of papers is still marginal compared to the ones investigating marine environments. In this study, we present the first detection of microplastics (MPs) in Central and Eastern European (CEE) surface waters and, globally, the first detection in fish ponds. Samples were taken from different types of fish ponds and natural water bodies along a novel concept down to a particle size of 100 µm, then, after sample preparation, MPs were characterized using an FTIR microscope. 92% of the water samples contained MPs ranging from 3.52 to 32.05 particles/m3. MPs were detected in 69% of the sediment samples ranging from 0.46 to 1.62 particles/kg. Dominant abundance of polypropylene (PP) and polyethylene was shown in water and PP and polystyrene in sediment samples. First results also indicate that fish ponds may act as a deposition area for MPs.

Contributions to the taxonomy, identification, and biogeography of the Western Palaearctic species of Campoletis Förster (Ichneumonidae: Campopleginae).

Two new species of Campoletis Förster (Ichneumonidae: Campopleginae) are described: Campoletis katalinarum sp. nov. from Georgia and Turkey, and Campoletis margaritae sp. nov. from Romania and Armenia. Affixes to the identification key of the most recent revision of the genus are given. Additionally, contributing to the knowledge on biogeographical distribution of several species of the genus, 27 new records of 14 Campoletis species are reported, representing first records for 12 countries in the Palaearctic region.

A new species of Aethra (Decapoda: Brachyura: Aethridae) from the early Serravallian (middle Miocene) of the Medobory Hills (Ukraine).

A new species of the genus Aethra (A. stalennyii n. sp.) from the Badenian (middle Miocene) reef frameworks of the Maksymivka quarry in the Medobory Hills, near Ternopil (Ukraine) is described herein. Aethra stalennyii n. sp. represents the first fossil occurrence of the genus known to date. Aethra stalennyii n. sp. inhabited a very similar habitat to those all the extant congeners live in. The presence of Aethra in the middle Miocene of the Central Paratethys adds further emphasis on close affinities between Tethyan reef-associated decapod assemblages and their modern counterparts in the Indo-West Pacific waters. Aethra stalennyii n. sp. represents the first report of Aethra in the fossil record. Possible origin of the genus and subsequent migration eastward is discussed.

Tuberculosis in Late Neolithic-Early Copper Age human skeletal remains from Hungary.

Alsónyék-Bátaszék in Southern Hungary is one of the largest late Neolithic settlements and cemeteries excavated in Central Europe. In total, 2359 burials from the Late Neolithic - Early Copper Age Lengyel culture were found between 2006 and 2009 [1]. Anthropological investigations previously carried out on individuals from this site revealed an interesting paleopathological case of tuberculosis in the form of Pott's disease dated to the early 5(th) millennium BC. In this study, selected specimens from this osteoarcheological series were subjected to paleomicrobiological analysis to establish the presence of MTBC bacteria. As all individuals showing clear osteological signs of TB infection belonged to a single grave group, 38 individuals from this grave group were analysed. The sample included the case of Pott's disease as well as individuals both with and without osseous TB manifestations. The detection of TB DNA in the individual with Pott's disease provided further evidence for the occurrence of TB in Neolithic populations of Europe. Moreover, our molecular analysis indicated that several other individuals of the same grave group were also infected with TB, opening the possibility for further analyses of this unique Neolithic skeletal series.