A new hominoid species from the middle Miocene site of Paşalar, Turkey

A new species of fossil hominoid is described from the middle Miocene deposits at Pa?alar, Turkey. It is the less common of the two Pa?alar species discussed by Martin and Andrews (1993), making up approximately 10% of the individuals in the Pa?alar hominoid sample according to analyses of the minimum number of individuals. To the diagnostic features of I(1) described by Alpagut et al. (1990) and Martin and Andrews (1993) can now be added further diagnostic features of all the anterior teeth, as well as both upper premolars and P(3). These include discrete, nonmetric features and metric differences at all the noted tooth positions. Attempts to distinguish the upper and lower molars of the two species have so far been unsuccessful, with the possible exception of M(3). The morphology of the new species is similar in most respects to that of Kenyapithecus wickeri from Fort Ternan, especially concerning maxillary morphology. They share robust and moderately deep maxillary alveolar processes, a restricted maxillary sinus with an elevated and uncomplicated floor, lacking the compartmentalization evident to varying degrees in many other taxa, and a zygomatic process that originates and turns laterally fairly high above the alveolar margin. There are also a number of distinctive similarities in the dentition, particularly for I(1), C(1), P(4) and P(3). The I(1) morphology in particular, with greatly hypertrophied lingual marginal ridges bounding a uniformly thickened basal crown area, is distinctive among Miocene hominoids. All of these similarities serve to reinforce the differences noted by others between the derived morphology of K. wickeri and the more primitive morphology of Equatorius africanus from Maboko and Kipsaramon. The new species differs from K. wickeri in morphological details of most of the anterior and premolar teeth that are known for both species, despite the general morphological similarity, and in the size of I(1) versus I(2). One striking feature of the new species is a relatively large incisive fossa, although it cannot be determined if this is associated with an open palatine fenestra, as in many early Miocene hominoids, or a minimally overlapping palate and nasoalveolar clivus, as in some middle and late Miocene hominoids.     

Metric variation in the postcanine teeth from Paşalar, Turkey

The coefficient of variation (CV) has frequently been used to evaluate variation in a morphologically homogeneous fossil assemblage and to make inferences about its species composition. Comparisons of coefficients of variation in single- and mixed-species assemblages have typically involved mixed-species assemblages made up of an equal number of data points from two or more different species, but in reality, a fossil assemblage may be biased in favor of one or more of the component species. In this paper, we evaluate the effects of unequal species composition in mixed-species assemblages on CV distributions by carrying out an extensive series of resampling experiments. The experiments were designed to replicate a paleontological situation in which the species affiliation of particular specimens is not known. We use this technique to explore the pattern of metric variation in the postcanine dental assemblage from the middle Miocene site of Pa?alar in a comparative context. The distributions of CVs from mixed assemblages that are heavily biased toward one species may be characterized by a greater range of CVs, increased skewing, and a tail of low values, but only heavily biased assemblages comprising species that differ markedly in size could be reliably identified on this basis. Evidence from the simulated CV distributions supports nonmetric evidence in indicating that the Pa?alar assemblage represents a heavily biased assemblage of two species that are similar in size but not entirely overlapping in the size distributions of their postcanine teeth.     

Hominoid phalanges from the middle Miocene site of Paşalar, Turkey

Eleven proximal and ten intermediate partial or complete hominoid phalanges have been recovered from the middle Miocene site of Pa?alar in Turkey. Based on species representation at Pa?alar, it is likely that most or all of the phalanges belong to Griphopithecus alpani rather than Kenyapithecus kizili, but both species may be represented. All of the complete or nearly complete phalanges appear to be manual, so comparisons to extant and other fossil primate species were limited to manual phalanges. Comparisons were made to extant hominoid and cercopithecoid primate genera expressing a variety of positional repertoires and varying degrees of arboreality and terrestriality. The comparisons consisted of a series of bivariate indices derived from previous publications on Miocene catarrhine phalangeal morphology. The proximal phalanges have dorsally expanded proximal articular surfaces, which is characteristic of cercopithecoids and most other Miocene hominoids, and indicates that the predominant positional behaviors involved pronograde quadrupedalism. Among the extant primates, many of the proximal and intermediate phalangeal indices clearly distinguish more habitually terrestrial taxa from those that are predominantly arboreal, and especially from taxa that commonly engage in suspensory activities. For nearly every index, the values of the Pa?alar phalanges occupy an intermediate position-most similar to values for Pan and, to a lesser extent, Macaca-indicating a generalized morphology and probably the use of both arboreal and terrestrial substrates. At least some terrestrial activity is also compatible with reconstructions of the Pa?alar habitat. Most proximal and intermediate phalanges of other middle and late Miocene hominoids have similar index values to those of the Pa?alar specimens, revealing broadly similar manual phalangeal morphology among many Miocene hominoids.     

The deciduous dentition of Griphopithecus alpani from Paşalar, Turkey

Seventy-four hominoid primary teeth have been recovered from the middle Miocene site of Pa?alar, Turkey, constituting the largest sample of deciduous teeth for any species of fossil ape. Morphological features that characterize the permanent teeth of Griphopithecus alpani from the site have also been identified in some of these deciduous teeth, including a lingual pillar on the di(1)s. These features plus the overwhelming preponderance of G. alpani permanent teeth at the site suggest that all of the deciduous teeth belong to this species. Contrary to the situation in the permanent teeth, nothing in the morphology of the primary dentition suggests the representation of a second species. The age profile of the non-adult hominoids was reconstructed based on the degree and type of wear recorded on the dp4s, the most abundant deciduous tooth in the sample, assuming a similar eruption chronology to that of Pan troglodytes. This analysis indicates underrepresentation of very young individuals in the sample and high mortality for individuals belonging to the 3-5-years age cohort, a situation that could be due to the effects of stress related to weaning. The coefficient of variation and range-index values obtained for the majority of tooth types are equal to or greater than the comparable values in a sample of P. troglodytes, in some cases at much smaller sample sizes. One possible explanation for this is that there was greater sexual dimorphism in the G. alpani deciduous dentition than in Pan, which would mirror the condition of the permanent dentition.     

Identification of a single birth cohort in Kenyapithecus kizili and the nature of sympatry between K. kizili and Griphopithecus alpani at Paşalar

The sample of the less common hominoid species at Pa?alar, Kenyapithecus kizili, is characterized by a number of unusual attributes. All ten of the upper central incisors attributed to this species show a distinct, identical pattern of two linear enamel hypoplasias. The two hypoplasias occur on the same portion of the labial crown face, revealing that the two hypoplasia-causing events occurred at the same stage of development in all individuals. The morphology of the two hypoplasias and the amount of time between them, as determined by both their separation and counts of perikymata, are also the same on all teeth. In addition, all of the approximately 70 teeth assigned to K. kizili appear to come from young adults based on degrees of wear; there are no younger or older individuals (diagnostic morphology at most tooth positions would be evident even with heavy wear). Thus, all of the K. kizili individuals (minimum number of individuals is nine: seven males, two females) appear to have died at essentially the same age. It is concluded that the most plausible interpretation of all these features is that the incisor hypoplasias were caused by the same two events in all the K. kizili individuals and that these individuals therefore represent a single birth cohort. As such, and because they died at essentially the same age, they would also have died at the same time, which is consistent with the catastrophic nature of the Pa?alar deposits. The number of coincidences needed to explain all of the attributes of the K. kizili sample if these animals were born in, and died in, different years seems highly improbable. Moreover, the lack of a typical age-class structure for the K. kizili sample, or any age-class structure at all beyond the one age class of young adult, strongly suggests that the species was not resident in the area that contributed to the Pa?alar accumulation, and that K. kizili was not permanently sympatric with the other Pa?alar hominoid, Griphopithecus alpani. Rather, the nine K. kizili individuals must have been transients in, or recent immigrants to, the area at the time of the events that led to the formation of the site. Recent observations on social associations in male chimpanzees offer at least a possible interpretive framework to explain this unprecedented occurrence in the primate fossil record.     

Differential roles of proteasome and immunoproteasome regulators Pa28αβ, Pa28γ and Pa200 in the degradation of oxidized proteins

The response and functions of proteasome regulators Pa28αβ (or 11S), Pa28γ and Pa200 in oxidative-stress adaptation (also called hormesis) was studied in murine embryonic fibroblasts (MEFs), using a well-characterized model of cellular adaptation to low concentrations (1.0-10.0 μM) of hydrogen peroxide (H(2)O(2)), which alter gene expression profiles, increasing resistance to higher levels of oxidative-stress. Pa28αβ bound to 20S proteasomes immediately upon H(2)O(2)-treatment, whereas 26S proteasomes were disassembled at the same time. Over the next 24h, the levels of Pa28αβ, Pa28γ and Pa200 proteasome regulators increased during H(2)O(2)-adaptation, whereas the 19S regulator was unchanged. Purified Pa28αβ, and to a lesser extent Pa28γ, significantly increased the ability of purified 20S proteasome to selectively degrade oxidized proteins; Pa28αβ also increased the capacity of purified immunoproteasome to selectively degrade oxidized proteins but Pa28γ did not. Pa200 regulator actually decreased 20S proteasome and immunoproteasome's ability to degrade oxidized proteins but Pa200 and poly-ADP ribose polymerase may cooperate in enabling initiation of DNA repair. Our results indicate that cytoplasmic Pa28αβ and nuclear Pa28γ may both be important regulators of proteasome's ability to degrade oxidatively-damaged proteins, and induced-expression of both 20S proteasome and immunoproteasome, and their Pa28αβ and Pa28γ regulators are important for oxidative-stress adaptation.     

Enhanced Wnt/β-catenin signalling during tooth morphogenesis impedes cell differentiation and leads to alterations in the structure and mineralisation of the adult tooth

BACKGROUND INFORMATION: Previous studies have indicated that over-activation of the wingless interaction site (Wnt)/β-catenin signalling pathway has important implications for tooth development, at the level of cell differentiation and morphology, as well as for the production of supernumerary teeth. Here, we provide evidence for a crucial role of this signalling pathway during the stage of tooth morphogenesis. We have developed an in vitro model consisting of 14.5-day-old mouse embryo first molars, in which the Wnt pathway is overactivated by the glycogen synthase kinase-3 inhibitor 6-bromoindirubin-3'-oxime (BIO; 20 μM). RESULTS: We found that over-activation of the Wnt/β-catenin pathway delayed the differentiation and growth of the inner dental epithelium. In addition, in contrast to controls in which Nestin protein expression was restricted to differentiated odontoblasts, in BIO-treated molars, Nestin expression spread through sub-odontoblastic cellular layers. This alteration appears to be related to: (i) the over-expression of Bmp4 in the same region, (ii) the delay in odontoblast precursor cell differentiation and (iii) increased proliferation of mesenchymal cells. Furthermore, treatments longer than 6 days induced the malformation of typical dental structures and led to a total lack of cell differentiation. Finally, over-activation of the Wnt route during odontogenesis resulted in adult teeth which presented altered size, morphology and mineralisation. CONCLUSIONS: Our results indicate that Wnt/β-catenin over-activation during tooth morphogenesis is sufficient to cause dramatic alterations in the adult tooth, by delaying cellular differentiation and stimulating proliferation of the dental mesenchyme of developing teeth.     

Age determination in roe deer — a new approach to tooth wear evaluated on known age in dividuals

A novel, simple, and objective method is presented for ageing roe deerCapreolus capreolus (Linnaeus, 1758) evaluated on 471 lower jaws from roe deer of known age (351 with permanent premolars). It is based on tooth eruption patterns and presence/absence of wear characters in jaws from roe deer integrated in a scoring system. Permanent cheek teeth emerge in May–July in the year af ter birth, which enables precise age determination of individuals with deciduous premolars. For individuals with permanent cheek teeth, the method provides the correct age for all individ uals younger than 13 months and > 80% of all individuals between 13 and 24 months old. For older in dividuals the accuracy decreases, but decent accuracy is achieved to the age of 48 months. Males have higher wear rates than females corroborating recent documentation of sex-specific life history tactics in ungulates. The data originate from two separated Danish roe deer populations exposed to contrasting habitats, but no difference in wear rate is found between populations. Thus, previous concern about the validity of age determination methods based on tooth wear may have been overstated. The findings demonstrate that objective measures of tooth wear can provide the basis for age determination in ungulate species that are otherwise difficult to age.     

The associations of morningness–eveningness with anger and impulsivity in the general population

The aim of this study was to investigate the relationships among morningness–eveningness, impulsivity and anger in the general population. A total of 1000 community-dwelling subjects (500 males) aged 20–77 years (mean± SD age: 39.6 ± 11.6 years) completed the morningness–eveningness questionnaire (MEQ), Barratt impulsiveness scale (BIS), Spielberger State–Trait Anger Expression Inventory (STAXI) and Center for Epidemiologic Studies Depression Scale. Moderation and mediation analyses were performed to determine whether the relationship between two variables depended on the third variable, referred to as a moderator, and whether the third variable, known as a mediator, was associated with the other two variables establishing causation. The MEQ scores exhibited significant negative associations with BIS (p < 0.001) and STAXI (p < 0.001) scores, and high scores on the BIS were associated with high scores on the STAXI (p < 0.001). Impulsivity, as measured by the BIS, played a role as a moderator (p < 0.001) in the relationship between MEQ and STAXI, and anger, as measured by the STAXI, acted as moderator (p = 0.030) in the association between MEQ and BIS. However, after controlling for the interaction of the BIS and MEQ, the MEQ scores did not significantly predict STAXI scores (p = 0.070). Additionally, the effect size of the mediating effect of the BIS scores on the relationship between the MEQ and STAXI (percent mediation: 53.2%) was larger than that of the STAXI scores on the association between the MEQ and BIS (percent mediation: 31.8%). The present results demonstrate that morningness–eveningness was closely related with both impulsivity and anger in the general population. Furthermore, these findings suggest that impulsivity may exercise a great influence on the association between morningness–eveningness and anger in two ways: as a moderator by modulating this relationship based on the level of impulsivity and as a mediator by acting as an intermediary factor.     

Longer in the tooth,shorter in the record? The evolutionary correlates of hypsodonty in Neogene ruminants

The acquisition of hypsodont molars is often regarded as a key innovation in the history of ruminant ungulates. Hypsodont ruminants diversified rapidly during the later Neogene, circa 15-2 Myr ago, and came to dominate the ruminant fossil record in terms of species diversity. Here we show that hypsodont clades had higher speciation and diversification rates than other clades. Hypsodont species had, on average, shorter stratigraphic durations, smaller range size and lower occupancy than non-hypsodont species. Within hypsodont clades, some species were very common and acquired large geographical ranges, whereas others were quite rare and geographically limited. We argue that hypsodont clades diversified in an adaptive radiation-like fashion, with species often splitting cladogenetically while still in the expansive phase of their occupancy history.     

The many facets of the Tim-Tipin protein families’ roles in chromosome biology

Failures in DNA replication are a potent force for driving genome instability. The proteins which form the replisome, the DNA replication machinery, play a fundamental role in preventing replicative catastrophes. The Tim (TIMELESS/TIMEOUT) and Tipin proteins are two conserved replisome associated proteins which have functions in preventing replication fork collapse and replicative checkpoint signalling in response to factors which slow the progression of the replisome. Intriguingly, TIMELESS family members have been implicated in the regulation of the biological clock, giving a tantalising pointer to a possible link between DNA replication and circadian rhythm control. Here we report on our current understanding of the many facets of these protein families in maintaining genome stability and replication checkpoint control.     

Are there associations between grain-filling rate and photosynthesis in the flag leaves of field-grown rice?

Rate of grain filling in terms of dry mass accumulated per panicle per day was measured in field-grown rice in the dry season in the Philippines and compared to rates of light-saturated photosynthesis per unit leaf area (P(max)) measured at 350 micro l l(-1) CO(2) for 21 d after flowering. Five new plant type (tropical japonica) varieties (NPT) and one indica variety (IR72) were used and these gave some variation in rates and patterns of grain filling. A rapid grain-filling phase (RGFP) occurred approximately 10 d after flowering in most varieties. There was no consistent relationship in any variety between the rate of grain-filling and P(max) and chlorophyll content, both of which remained mostly unchanged throughout grain filling. Significant declines in the amount of total leaf protein and ribulose bisphosphate carboxylase-oxygenase (Rubisco) occurred, but these did not occur at the same time as the RGFP in all varieties. A decrease in the ratio of chlorophyll a/b preceded these changes and a transient rise in chlorophyll content was also observed in four varieties at this time. There was no significant change in leaf non-structural carbohydrate content during or following the RGFP. It is concluded that the decline in Rubisco and protein content in NPT was not reflected in photosynthetic activity. Hence in these field experiments Rubisco accumulated to a level in excess of photosynthetic requirements, serving as a store of nitrogen for grain filling.     

Assessing spatiotemporal associations in the occurrence of badger–human conflict in England

Examples from a variety of taxa demonstrate that under certain circumstances, the exclusion or translocation of ‘problem’ animals is ineffective in resolving human–wildlife conflicts and may even elicit new problems elsewhere. Damage caused by badger setts (burrows) is an important source of human–wildlife conflict in the UK and is commonly managed by excluding badgers from all or part of problem setts. We used records of licences issued for the management of such problems and a novel statistical approach to assess spatiotemporal associations between problem cases in England from 2002 to 2005. We predicted that management at urban badgers' setts, and particularly exclusion of badgers from urban main setts, would give rise to subsequent problems at focal setts and in neighbouring areas. Frequencies of problems occurring at individual setts were similar in urban and rural areas. In areas neighbouring setts subjected to management action, the background frequency of problems was higher in urban than in rural areas, reflecting the occurrence of problems at a higher proportion of urban setts. The frequency of new cases arising at or in the vicinity of managed setts within a critical time period after management action was not significantly different from the background frequency of problems for any combination of land use, sett type and management approach. This finding suggests that the measures currently employed for managing problem setts do not importantly increase the likelihood of problems reoccurring in the same location or emerging nearby.     

An Overview of the Ichnological and Ethological Studies in the Cave Bear Den in Urşilor Cave (Western Carpathians,Romania)

This paper presents a preliminary investigation of late Pleistocene cave bear traces from Ur?ilor Cave in the western Carpathians, Romania. The bears left thousands of traces on the walls, plateaus, and slopes of the cave interior. Some areas in the cave have been heavily trampled, leaving more than 140 hibernation beds as well as fur impressions. The footprints of cave bears are assigned to Ursichnus europaeus nov. ichnogen. and nov. ichnosp. and the cave bear-beds to Ursalveolus carpathicus nov. ichnogen. and nov. ichnosp. as behavioral traces. Tens of thousands of scratch marks on the slopes and top of a clay plateau are the result of bears moving from the hibernation area down to a stream and back. These traces reveal that the cave bears had short claws, similar to those of modern, primarily herbivorous black bears. Deep within the cave, three weathered, articulated cave bear skeletons still lie in their hibernation beds on the clay plateaus or a natural cave corner. One of these bears was a one-year-old male cub that did not survive its first hibernation; a second skeleton close was an adult female. The third, a young male, was found close to the end of the cave system. The bears of Ur?ilor Cave would have felt well protected against carnivores during their hibernation because their sleeping places were so deep within the cave. Their strategy may well have been to avoid any conflict with hyenas and lions during hibernation.