Ecomorphology of the European Hyaenodon

Hyaenodon war eine artenreiche Gattung im späten Eozän bis ins frühe Miozän der nördlichen Hemisphäre (Europa, Nordamerika und Asien). Unter Berücksichtigung der spärlichen Erhaltung von karnivoren Säugetieren, ist der Nachweis aus bestimmten oligozänen Fundstellen beachtlich. Das trifft besonders auf Nordamerika (z.B. White River Formation) zu. In Europa lieferten die Spaltenfüllungen von Quercy (Frankreich) einen reichhaltigen Fossilnachweis, darunter auch Hyaenodon. Das asiatische Material ist weit weniger umfangreich und die meisten Funde stammen aus der Mongolei. Der Mittelpunkt dieser Arbeit ist die Ökomorphologie des europäischen Hyaenodon und verfolgt das Ziel Ernährung, sensorische Fähigkeiten und Fortbewegung zu rekonstruieren. Taxonomisch verwertbare Informationen werden ebenfalls diskutiert. Hyaenodon wurde lange als Knochenbrecher gewertet. Die Zahnschmelzdicke, Ergebnisse aus der „stereomicrowear“ Analyse und die Ultrastruktur des Zahnschmelzes wurden als Zusatzinformation zu der Zahnmorphologie und dem Kaumuster verwendet, um die Ernährung dieses fossilen Räubers zu rekonstruieren. Hyaenodon besetzte eine breite Nische, die mit der des rezenten Löwen und der Tüpfelhyäne überlappt. Unterschiedliche Tendenzen in der Nahrungspräferenz konnten zwischen europäischen und nordamerikanischen Taxa festgestellt werden. Tast-, Gehörsinn und das Gehirn wurden je nach Möglichkeit untersucht. Erstmalig wird ein Gehörknöchelchen von Hyaenodon beschrieben. Das Foramen infraorbitale ist relativ klein. Hyaenodon besaß keine spezialisierten Tasthaare wie Katzen. Der Tastsinn, hier eingeschränkt beurteilt an den Tasthaaren, war ähnlich wie in Hunden entwickelt. Folgende Merkmale sind bei Betrachtung des bisherigen Wissens über das Gehirn zu erwähnen: Das Großhirn ist klein, aber gefurcht. Die olfaktorischen Loben sind groß. Der Geruchssinn war verglichen zu den anderen Sinnen sehr ausgeprägt. Der Gehörapparat wurde untersucht, darunter die Bulla auditiva, ein Gehörknöchelchen (der Incus) und das Innenohr (die Gehörschnecke und die Bogengänge). Die Bulla ist aus einem gekammerten Entotympanicum aufgebaut. Die Aufwickelung der Gehörschnecke ist von einem vergleichbaren Grad wie dem des Hundes. Die Bogengänge sind ausgedehnt. Der Incus zeigt die übliche sattelförmige Gelenksfläche. Sie ist in eine große superiore und eine kleine, runde, inferiore Fläche unterteilt. Verglichen mit modernen Raubtieren ist der Incus am ähnlichsten dem der Katze. Hyaenodon war nicht an geringe Frequenzen angepasst. Höchstwahrscheinlich verfügte es über ein weites Spektrum an Frequenzen (wie Raubtiere allgemein) und könnte an hohe Frequenzen angepasst gewesen sein (wie die Katze). Die Skelettelemente, auch wenn weniger zahlreich erhalten in Europa, waren die Grundlage für Rückschlüsse auf das Bewegungsmuster und die Jagdstrategie. Das Ligamentum nuchae unterstützte den proportional zum Körper übergroßen Kopf. Wenn dieses Ligament tatsächlich vorhanden war, dann setzte es nicht am Epistropheus an wie in Caninae. Stattdessen wird hier vorgeschlagen, dass es am Hinterhaupt wie in Huftieren ansetzte. Das Skelett von Hyaenodon zeigt terrestrische und kursoriale Merkmale. Generell war es von robustem Bau und mit starker Muskulatur ausgestattet. Der Fibio-Tarsal-Komplex weist auf Bewegung über unebene Flächen hin. Das Fortbewegungsmuster wird hier als kursorial beurteilt und in dem Sinne ausgelegt, dass Hyaenodon ein primär am Boden lebender Räuber mit Laufanpassungen war. Die Eckzähne waren mediolateral komprimiert und in diesem Aspekt ähnlich den Caniden und damit anders als die im Querschnitt runden Eckzähne der Feliden. Dieser Typ ist ideal für das Aufschlitzen der Beute. Daher fügte Hyaenodon dem Beutetier mehrere flache Wunden zu und tötete nicht mit einem einzigem Biss wie moderne Feliden (dafür spricht auch die Ausbildung der Schnurrhaare). Die primäre Jagdstrategie war der Angriff aus dem Hinterhalt, aber auch eine Verfolgungsjagd mit den meisten damaligen Beutetieren konnte erfolgreich ausgehen. Hyaenodon war ein gefährlicher Räuber in den Wäldern des Eozäns und der offeneren Landschaft des Oligozäns. Taxonomisch verwertbare Ergebnisse wurden ebenfalls erzielt: Der Zahnwechsel verläuft bei europäischen Formen anders als bei Nordamerikanischen: deutliche Unterschiede sind im Ablauf des Zahnwechsels im Unterkiefer (den p3, p4, m3 und den Eckzahn betreffend) und auch im Oberkiefer (den P1 betreffend) festzustellen. Die Milchbezahnung ist in Europa je nach Art ausgeprägt. Interessanterweise zeigen die oligozänen Arten Hyaenodon exiguus und Hyaenodon filholi unterschiedliche Morphologien der Oberkiefermilchzähne DP3 und DP4 (Entwicklung des anterioren Höckers und der Schneidekante) und der Unterkiefermilchzähne dp3 und dp4 (Entwicklung des anterioren Höckers und des Talonids). Im Vergleich mit juvenilem Material aus Nordamerika und Asien, erscheint die Morphologie von Hyaenodon exiguus als europäische Entwicklung. Die Annahme, dass Hyaenodon filholi ein asiatischer Einwanderer war, wird hier betont. Divergenz innerhalb der Gattung Hyaenodon wurde festgestellt. Manche Unterschiede zwischen den nordamerikanischen und den europäischen Formen, die schon in der Literatur beschrieben worden sind, erwiesen sich als ungültig, z.B. das Vorhandensein eines Scapholunare in den eruopäischen Arten (basierend auf falscher Zuordnung). Der unterschiedliche Zahnwechsel ist jedoch ein beständiger Unterschied, ebenso wie die plesiomorphe Morphologie des M1 in den nordamerikanischen Formen. Die Wurzelentwicklung (ein- oder zweiwurzelig) des p1 ist variabel, gibt aber keine deutlichen Hinweise: die älteste bekannte Art zeigt einen einwurzeligen p1 und die oligozänen Arten Europas besitzen einen zweiwurzeligen p1. Das Skelett des europäischen Hyaenodon weist Unterschiede zwischen eozänem und oligozänem Material auf. Die Evolution des nordamerikanischen Hyaenodon wird als eigene Linie anerkannt.Hyaenodon was a speciose genus in the late Eocene to early Miocene distributed over the Northern hemisphere (Europe, North America and Asia). Taking into account that carnivorous mammals are not as numerously preserved as others, certain Oligocene sites yield a considerable amount of material. This is true especially for North America (e.g. White River formation). In Europe, Quercy (France) is a complex of fissure fillings bearing a rich fossil record of many taxa including Hyaenodon. Concerning Asia the evidence is sparser and most findings come from Mongolia. The focus of this thesis is on the ecomorphology of the European Hyaenodon with the aim to reconstruct diet, sensory capabilities and the locomotion pattern. However, facts of taxonomic value are discussed as well. Hyaenodon has long been regarded as having been capable of processing bone. The enamel thickness, stereomicrowear results and the ultrastructure of the enamel were used as additional information to dental morphology and wear patterns to reconstruct the diet of this fossil predator. The ossiphagous tendencies can be confirmed. Hyaenodon occupies a broad dietary niche — overlapping with those of the recent lion and the spotted hyena. Different dietary tendencies were observed in the European and the North American taxa. The sense of touch and hearing, as well as the brain were examined in detail. For the first time an auditory ossicle of Hyaenodon is described. The infraorbital foramen is rather small. Hyaenodon did not possess as specialized whiskers as felids do. The sense of touch — concerning the whiskers — was equally developed as in canids. Reviewing the hitherto knowledge of the brain — the following features were apparent: the neocortex is small, but folded; the olfactory lobes are largely developed. The sense of smell was compared to other senses important in Hyaenodon. The hearing apparatus was examined including the bulla, an auditory ossicle (the incus) and the inner ear (cochlea and semicircular canals). The bulla is made up by a chambered entotympanic. The coiling of the cochlea is of the same degree as in dogs. The semicircular canals are extensive. The incus is showing the usual saddle-shaped facet, but with a large upper articulation surface and a small, circular lower articulation facet. Compared to recent carnivores it is most similar to cats. Hyaenodon was not adapted to low frequencies and most probably had a wide frequency range (as predators commonly do) with a possible adaptation to high frequencies (as in cats). The skeletal remains, although by far less frequently preserved in Europe than in North America, were the basis for the inferences of locomotion pattern and predation style. The nuchal ligament was assumed to support the proportionally oversized head (compared to recent Carnivora). If this ligament really was present, it did not insert on the epistropheus as it does in canine dogs. It is proposed here, that it inserted on the posterior aspect of the skull as in ungulates. The skeleton of Hyaenodon exhibits terrestrial and cursorial features. Its general structure was robust with a strong musculature. The fibio-tarsal complex indicates progression over uneven surface. Hyaenodon´s locomotion pattern is judged here as cursorial and interpreted in the sense that this predator was dwelling primarily on the ground with an adaptation to running. The canines of Hyaenodon are mediolaterally compressed and like those in canids and not like the rounded canines in felids. This type of canines is ideal for slashing. Thus, Hyaenodon inflicted its prey with shallow wounds and not with a killing bite like recent felids do (additional evidence comes from the non-specialized whiskers). The predation style of Hyaenodon was ambushing, although it did not succumb in a chase with most contemporaneous prey animals. It was a powerful predator in the forests of the Eocene as well as in the more open landscape of the Oligocene. Some taxonomic information can be provided as well: The tooth eruption sequence is differing between the European and the North American forms: there are significant differences in the sequence of the lower jaw (concerning the p3, the p4, the m3 and the canine) and a difference in the sequence of the upper jaw (concerning the eruption of the P1). The milk dentition in European Hyaenodon species shows different morphologies. For the first time differences are described concerning the morphology of the upper milk teeth DP3 and DP4 (development of the anterior cusp and the cutting blade) and the lower milk teeth dp3 and dp4 (development of the anterior cuspid and the talonid) in the Oligocene species Hyaenodon exiguus and Hyaenodon filholi. Compared with other known juvenile material from North America and Asia, the morphology of Hyaenodon exiguus appears to be a product of European evolution. The assumption that Hyaenodon filholi was an Asian immigrant is emphasized here. Divergence within the genus was found. Some differences between the North American and the European Hyaenodon described in literature turned out to be not valid, e.g. the presence of a scapholunatum in the European taxa (based on misidentifications). However, the different tooth eruption pattern is a consistent difference, as well as the more plesiomorphic morphology of the M1 in North American forms. The root development (single- or double-rooted) of the p1 is varying, but does not give distinct clues: the oldest species show a single-rooted p1 and the Oligocene species in Europe possess a double-rooted p1. The skeleton of the European Hyaenodon shows differences between Eocene and Oligocene material. The evolution of the North American Hyaenodon is acknowledged as an own lineage

Palaeodietary traits of large mammals from the middle Miocene of Gracanica (Bugojno Basin, Bosnia-Herzegovina)

Recent excavations at the Gracanica coal mine (Bugojno Basin, Bosnia-Herzegovina) have unearthed numerous skeletal parts of fossil vertebrates, including a noteworthy collection of mammalian remains. Previous palaeoecological investigations of the Dinarides Lake System were established using stratigraphical, palaeofloral, and malacological data. However, large mammal remains have so far not been used to reconstruct the terrestrial palaeoenvironment of this important fossil ecosystem. Here, the palaeodietary preferences of large mammals were investigated, using a multiproxy approach by employing dental microwear and dental mesowear analysis, in order to provide new perspectives on the terrestrial palaeoecology of the Dinarides Lake System. The dental microwear of all available adult mammalian teeth was analysed. Dental mesowear analysis was employed for ungulate and proboscidean taxa, using mesowear scores and mesowear angles, respectively. The analysis reveals the presence of browsing, "dirty browsing", and mixed-feeding herbivorous taxa, with seasonal fruit, or even grass intake. Additionally, the analysis of the carnivores suggests the presence of hyaena- and cheetah-like hypercarnivores, as well as generalists. The palaeodietary traits of the fossil mammals suggest a closed canopy-like environment, which is supported by the fossil plant assemblage. Palaeopalynological data confirm the omnipresence of fleshy fruit-bearing plants, herbaceous taxa, as well as grasses, which justifies the seasonal fruit browsing, the common "dirty browsing", and the occasional grazing behaviour visualized for some of the fossil mammals from Gracanica.Peer reviewe

Pollen season is reflected on symptom load for grass and birch pollen-induced allergic rhinitis in different geographic areas—An EAACI Task Force Report

Background: The effectiveness of allergen immunotherapy (AIT) in seasonal allergic rhinitis (AR) depends on the definition of pollen exposure intensity or time period. We recently evaluated pollen and symptom data from Germany to examine the new definitions of the European Academy of Allergy and Clinical Immunology (EAACI) on pollen season and peak pollen period start and end. Now, we aim to confirm the feasibility of these definitions to properly mirror symptom loads for grass and birch pollen-induced allergic rhinitis in other European geographical areas such as Austria, Finland and France, and therefore their suitability for AIT and clinical practice support. Methods: Data from twenty-three pollen monitoring stations from three countries in Europe and for 3 years (2014-2016) were used to investigate the correlation between birch and grass pollen concentrations during the birch and grass pollen season defined via the EAACI criteria, and total nasal symptom and medication scores as reported with the aid of the patient's hay-fever diary (PHD). In addition, we conducted a statistical analysis, together with a graphical investigation, to reveal correlations and dependencies between the studied parameters. Results: The analysis demonstrated that the definitions of pollen season as well as peak pollen period start and end as proposed by the EAACI are correlated to pollen-induced symptom loads reported by PHD users during birch and grass pollen season. A statistically significant correlation (slightly higher for birch) has been found between the Total Nasal Symptom and Medication Score (TNSMS) and the pollen concentration levels. Moreover, the maximum symptom levels occurred mostly within the peak pollen periods (PPP) following the EAACI criteria. Conclusions: Based on our analyses, we confirm the validity of the EAACI definitions on pollen season for both birch and grass and for a variety of geographical locations for the four European countrie

Real-world data using mHealth apps in rhinitis, rhinosinusitis and their multimorbidities

Digital health is an umbrella term which encompasses eHealth and benefits from areas such as advanced computer sciences. eHealth includes mHealth apps, which offer the potential to redesign aspects of healthcare delivery. The capacity of apps to collect large amounts of longitudinal, real-time, real-world data enables the progression of biomedical knowledge. Apps for rhinitis and rhinosinusitis were searched for in the Google Play and Apple App stores, via an automatic market research tool recently developed using JavaScript. Over 1500 apps for allergic rhinitis and rhinosinusitis were identified, some dealing with multimorbidity. However, only six apps for rhinitis (AirRater, AllergyMonitor, AllerSearch, Husteblume, MASK-air and Pollen App) and one for rhinosinusitis (Galenus Health) have so far published results in the scientific literature. These apps were reviewed for their validation, discovery of novel allergy phenotypes, optimisation of identifying the pollen season, novel approaches in diagnosis and management (pharmacotherapy and allergen immunotherapy) as well as adherence to treatment. Published evidence demonstrates the potential of mobile health apps to advance in the characterisation, diagnosis and management of rhinitis and rhinosinusitis patients.Peer reviewe

How to do a clinical trial? Recommendations from the aerobiological point of view

Background: Allergy immunotherapy is still the only treatment of pollen allergy, providing a long-term effect. Clinical trials with pollen allergic patients are in need of validated, high quality pollen data and forecasts in order to grant comparability and to adhere to scientific standards. The aerobiological part of clinical trials remained hitherto not well defined, leaving the definition and use of pollen and forecast data more or less open. Methods: Pollen data of eight Austrian pollen-monitoring stations were selected and used as an example to present a new method of pollen data replacement, in case of station failure. Gower's similarity provides an objective calculation based on a defined time frame and a specific aeroallergen (for example birch, grass, mugwort and ragweed). Results: The ideal planning of the aerobiological part of a clinical trial with a pollen extract is described in detail with specific recommendations concerning site selection, pollen and forecast data, definition of the pollen season, and risk management. A checklist for every clinical trial with an aerobiological part was developed. Conclusion: Virtual biogeographic regions are beneficial due to their objective establishment, and can be integrated into clinical trials. Pollen data is not the same as forecast data. Both datasets have to be critically evaluated by trained aerobiologists before they are used in clinical trials. Therefore, only institutions with aerobiological knowledge, at best ISO-certified, should be involved in clinical trials and handle the aerobiological tasks. Keywords: Pollen data, Aerobiology, Allergen immunotherapy, Clinical trials, Checklis

Journal of Medical Internet Research / Evaluation of Pollen Apps Forecasts: The Need for Quality Control in an eHealth Service

Background: Pollen forecasts are highly valuable for allergen avoidance and thus raising the quality of life of persons concerned by pollen allergies. They are considered as valuable free services for the public. Careful scientific evaluation of pollen forecasts in terms of accurateness and reliability has not been available till date. Objective: The aim of this study was to analyze 9 mobile apps, which deliver pollen information and pollen forecasts, with a focus on their accurateness regarding the prediction of the pollen load in the grass pollen season 2016 to assess their usefulness for pollen allergy sufferers. Methods: The following number of apps was evaluated for each location: 3 apps for Vienna (Austria), 4 apps for Berlin (Germany), and 1 app each for Basel (Switzerland) and London (United Kingdom). All mobile apps were freely available. Todays grass pollen forecast was compared throughout the defined grass pollen season at each respective location with measured grass pollen concentrations. Hit rates were calculated for the exact performance and for a tolerance in a range of 2 and 4 pollen per cubic meter. Results: In general, for most apps, hit rates score around 50% (6 apps). It was found that 1 app showed better results, whereas 3 apps performed less well. Hit rates increased when calculated with tolerances for most apps. In contrast, the forecast for the “readiness to flower” for grasses was performed at a sufficiently accurate level, although only two apps provided such a forecast. The last of those forecasts coincided with the first moderate grass pollen load on the predicted day or 3 days after and performed even from about a month before well within the range of 3 days. Advertisement was present in 3 of the 9 analyzed apps, whereas an imprint mentioning institutions with experience in pollen forecasting was present in only three other apps. Conclusions: The quality of pollen forecasts is in need of improvement, and quality control for pollen forecasts is recommended to avoid potential harm to pollen allergy sufferers due to inadequate forecasts. The inclusion of information on reliability of provided forecasts and a similar handling regarding probabilistic weather forecasts should be considered.(VLID)486420

World Allergy Organization Journal / The evaluation of pollen concentrations with statistical and computational methods on rooftop and on ground level in Vienna How to include daily crowd-sourced symptom data

Background It is recommended to position pollen monitoring stations on rooftop level to assure a large catchment area and to gain data that are representative for a regional scale. Herein, an investigation of the representativeness of pollen concentrations was performed for 20 pollen types in the pollen seasons 20152016 in Vienna for rooftop and ground level and was compared with weather data and for the first time with symptom data. Methods The complete data set was analyzed with various statistical methods including Spearmen correlation, ANOVA, KolmogorovSmirnov test and logistic regression calculation: Odds ratio and Yule's Q values. Computational intelligence methods, namely Self Organizing Maps (SOMs) were employed that are capable of describing similarities and interdependencies in an effective way taking into account the U-matrix as well. The Random Forest algorithm was selected for modeling symptom data. Results The investigation of the representativeness of pollen concentrations on rooftop and ground level concerns the progress of the season, the peak occurrences and absolute quantities. Most taxa examined showed similar patterns (e.g. Betula), while others showed differences in pollen concentrations exposure on different heights (e.g. the Poaceae family). Maximum temperature, mean temperature and humidity showed the highest influence among the weather parameters and daily pollen concentrations for the majority of taxa in both traps. Conclusion The rooftop trap was identified as the more adequate one when compared with the local symptom data. Results show that symptom data correlate more with pollen concentrations measured on rooftop than with those measured on ground level.(VLID)490428

The connection of pollen concentrations and crowd-sourced symptom data: new insights from daily and seasonal symptom load index data from 2013 to 2017 in Vienna

Abstract Background Online pollen diaries and mobile applications nowadays allow easy and fast documentation of pollen allergy symptoms. Such crowd-sourced symptom data provides insights into the development and the onset of a pollen allergy. Hitherto studies of the symptom load index (SLI) showed a discrepancy between the SLI and the total pollen amount of a season, but did not analyze the daily data. Methods The Patient’s Hayfever Diary (PHD) was used as data pool for symptom data. Symptom data of Vienna (Austria) was chosen as a large and local sample size within the study period of 2013 until 2017. The city was divided into three different areas based on equal population densities and different environmental factors. Correlation factors, regression lines, locally weighted smoothing (LOESS) curves and line plots were calculated to examine the data. Results Daily SLI and pollen concentration data correlates well and the progress of the SLI within a pollen season is mirrored by the pollen concentrations. The LOESS curves do not deviate much from the regression line and support the linearity of the symptom-pollen correlation on a daily basis. Seasonal SLI data does not follow the same pattern as the respective seasonal pollen indices. Results did not vary in the three areas within Vienna or when compared with the Eastern region of Austria showing no significant spatial variation of the SLI. Discussion Results indicate a linear relationship of the SLI and pollen concentrations/seasonal polllen index (SPIn) on a daily basis for both in general and throughout the season, but not on a seasonal basis. These findings clarify the frequent misinterpretation of the SLI as index that is tightly connected to pollen concentrations, but reflects as well the seasonal variation of the burden of pollen allergy sufferers. Conclusion More than just the seasonal pollen index has to be considered when the SLI of a selected pollen season has to be explained. Cross-reactivity to other pollen types, allergen content and air pollution could play a considerable role. The similar behavior of the SLI in Vienna and a whole region indicate the feasibility of a possible symptom forecast in future and justifies the use of a single pollen monitoring station within a city of the size of Vienna

Defining Pollen Seasons : Background and Recommendations

Purpose of Review The definition of a pollen season determines the start and the end of the time period with a certain amount of pollen in the ambient air. Different pollen season definitions were used for a long time including the use of different terms for data and methods used to define a pollen season. Recently suggested pollen season definitions for clinical trials were tested and applied for the first time to more aeroallergens. Recent Findings This is a review on pollen season definitions and the latest recommendations. Recently, proposed terminology in aerobiology is promoted here in order to support reproducibility and repeatability in research. Two pollen season definitions, one based on percentages and one based on pollen concentrations, were tested. Summary Percentage definitions can be recommended for standard aerobiological routines and for retrospective applications, whereas pollen concentrations definitions can be recommended for prospective applications such as clinical trials.(VLID)361609