The heteromorph ammonite genus Ammonitoceras Dumas, 1876 (Ancyloceratidae) in the lower Aptian (Lower Cretaceous) of the Les Ferres Aptian Basin (southeastern France)

Ammonitoceras Dumas, 1876 is a genus of heteromorph ammonites characterized by the presence of a peculiar ontogenetic stage in its inner whorls: the Ammonitoceras stage. But in spite of its wide paleogeographic and biostratigraphic extension throughout the Aptian (Lower Cretaceous), this genus remains poorly known. In the present work we study specimens of Ammonitoceras from the lower Aptian Deshayesites multicostatus (Deshayesites deshayesi Zone) to Dufrenoyia furcata (Dufrenoyia furcata Zone) subzones of the Les Ferres Aptian Basin (southeastern France). The results are as follows: (1) representatives of Ammonitoceras from this area are regarded dimorphic with criocone macroconchs and ancylocone microconchs, (2) their ontogenetic sequence is described, (3) their intraspecific variability is significant and concerns the adult size and the duration of the ontogenetic stages, especially the Ammonitoceras stage, (4) two species are recognized: the earlier Ammonitoceras ucetiae Dumas, 1876, characterized by a brief Ammonitoceras stage on average, and the latter Ammonitoceras lahuseni (Sinzow, 1906), characterized by a longer Ammonitoceras stage on average. The sample of the Deshayesites grandis Subzone (Deshayesites deshayesi Zone) is composed of specimens too fragmentary to be identified at species level.     

Integrating Ontogenetic Shift,Growth and Mortality to Determine a Species' Ecological Role from Isotopic Signatures

Understanding species linkages and energy transfer is a basic goal underlying any attempt at ecosystem analysis. Although the first food-web studies were based on gut contents of captured specimens, the assessment of stable isotopes, mainly δ¹³C and δ¹⁵N, has become a standard methodology for wide-range analyses in the last 30 years. Stable isotopes provide information on the trophic level of species, food-web length, and origin of organic matter ingested by consumers. In this study, we analyzed the ontogenetic variability of δ¹³C and δ¹⁵N obtained from samples of three Neotropical fish species: silver sardine (Lycengraulis grossidens, n=46), white lambari (Cyanocharax alburnus, n= 26), and the red-tail lambari (Astyanax fasciatus, n=23) in Pinguela Lagoon, southern Brazil. We developed a new metric, called the Weighted Isotopic Signature (φ ¹⁵N or φ ¹³C, ‰), that incorporates ontogenetic variability, body growth, and natural mortality into a single number.     

Life history of Rhamphorhynchus inferred from bone histology and the diversity of pterosaurian growth strategies

Background

Rhamphorhynchus from the Solnhofen Limestones is the most prevalent long tailed pterosaur with a debated life history. Whereas morphological studies suggested a slow crocodile-like growth strategy and superprecocial volant hatchlings, the only histological study hitherto conducted on Rhamphorhynchus concluded a relatively high growth rate for the genus. These controversial conclusions can be tested by a bone histological survey of an ontogenetic series of Rhamphorhynchus.

Methodology/Principal Findings

Our results suggest that Bennett''s second size category does not reflect real ontogenetic stage. Significant body size differences of histologically as well as morphologically adult specimens suggest developmental plasticity. Contrasting the ‘superprecocial hatchling’ hypothesis, the dominance of fibrolamellar bone in early juveniles implies that hatchlings sustained high growth rate, however only up to the attainment of 30–50% and 7–20% of adult wingspan and body mass, respectively. The early fast growth phase was followed by a prolonged, slow-growth phase indicated by parallel-fibred bone deposition and lines of arrested growth in the cortex, a transition which has also been observed in Pterodaustro. An external fundamental system is absent in all investigated specimens, but due to the restricted sample size, neither determinate nor indeterminate growth could be confirmed in Rhamphorhynchus.

Conclusions/Significance

The initial rapid growth phase early in Rhamphorhynchus ontogeny supports the non-volant nature of its hatchlings, and refutes the widely accepted ‘superprecocial hatchling’ hypothesis. We suggest the onset of powered flight, and not of reproduction as the cause of the transition from the fast growth phase to a prolonged slower growth phase. Rapidly growing early juveniles may have been attended by their parents, or could have been independent precocial, but non-volant arboreal creatures until attaining a certain somatic maturity to get airborne. This study adds to the understanding on the diversity of pterosaurian growth strategies.     

Morphological Variations within the Ontogeny of Deinonychus antirrhopus (Theropoda,Dromaeosauridae)

This research resulted from the determination that MCZ 8791 is a specimen of Deinonychus antirrhopus between one and two years of age and that the morphological variations within particular growth stages of this taxon have yet to be described. The primary goal of the research is to identify ontogenetic variations in this taxon. Histological analyses determined that the Deinonychus specimens AMNH 3015 and MOR 1178 were adults. Comparisons are made between MCZ 8791 and these adult specimens. The holotype, YPM 5205, and the other associated specimens of this taxon within the YPM collection are similar in size and morphology to AMNH 3015. Further comparisons were made with the three partial specimens OMNH 50268, MCZ 4371, and MOR 1182. Although these specimens represent only a partial ontogenetic series, a number of morphological variations can be described. One secondary goal of this research is to compare the known pattern of variable, informative, ontogenetic characters in MCZ 8791 to a similar pattern of morphological characters in the sub-adult dromaeosaurid specimen Bambiraptor feinbergorum, AMNH FR: 30556. If the characters that have been determined to represent variable juvenile morphology in the ontogeny of Deinonychus are exhibited in Bambiraptor, this study will begin the process of determining whether a similar, conservative, ontogenetic pattern exists throughout the rest of Dromaeosauridae. If defensible, it may reduce the number of sympatric taxa within this clade. The other secondary goal relates to the forelimb function. The approximate body size, forelimb length, wrist development, and the presence of a more prominent olecranon on the ulna of MCZ 8791 support the hypothesis that juveniles of this taxon possessed some form of flight capability.     

Ontogenetic trajectory and allometry of Diplonychus rusticus (Fabricius), an Oriental aquatic bug (Hemiptera: Belostomatidae) from the Western Ghats of India

Despite being one of the dominant groups in freshwater ecosystems, morphological and ontogenetic studies on aquatic Hemiptera have received little attention in the Oriental region. We present the ontogenetic trajectory and allometry of the widespread Oriental belostomatid species, Diplonychus rusticus (Fabricius) for the first time. We have measured nine different morphological variables throughout the growth of the bug using both field captured and laboratory reared specimens. Our results suggest that the developmental instars can be distinguished by the size variables, as seen in the Principal Component Analysis. On the basis of a CHAID (Chi-squared Automatic Interaction Detection) based regression tree, we also show that the characters – total length without head and maximum width – prove to be adequate for effective instar identification. The multivariate allometric growth pattern shows that different body parts exhibit different types of allometry. This is apparent in the allometry exhibited by forelegs and mid and hind legs, which show allometry of opposite polarities. This may be due to the different functions attributed to these body parts. Our results show that the growth pattern in D. rusticus is comparable with the New World genus Belostoma, suggesting a conserved growth pattern in the family Belostomatidae.     

Testing the ontogenetic base for the transient model of inflorescence development

Backgrounds and Aims

Current research in plant science has concentrated on revealing ontogenetic processes of key attributes in plant evolution. One recently discussed model is the ‘transient model’ successful in explaining some types of inflorescence architectures based on two main principles: the decline of the so called ‘vegetativeness’ (veg) factor and the transient nature of apical meristems in developing inflorescences. This study examines whether both principles find a concrete ontogenetic correlate in inflorescence development.

Methods

To test the ontogenetic base of veg decline and the transient character of apical meristems the ontogeny of meristematic size in developing inflorescences was investigated under scanning electron microscopy. Early and late inflorescence meristems were measured and compared during inflorescence development in 13 eudicot species from 11 families.

Key Results

The initial size of the inflorescence meristem in closed inflorescences correlates with the number of nodes in the mature inflorescence. Conjunct compound inflorescences (panicles) show a constant decrease of meristematic size from early to late inflorescence meristems, while disjunct compound inflorescences present an enlargement by merging from early inflorescence meristems to late inflorescence meristems, implying a qualitative change of the apical meristems during ontogeny.

Conclusions

Partial confirmation was found for the transient model for inflorescence architecture in the ontogeny: the initial size of the apical meristem in closed inflorescences is consistent with the postulated veg decline mechanism regulating the size of the inflorescence. However, the observed biphasic kinetics of the development of the apical meristem in compound racemes offers the primary explanation for their disjunct morphology, contrary to the putative exclusive transient mechanism in lateral axes as expected by the model.     

Trophic niche ontogeny and palaeoecology of early Toarcian Stenopterygius (Reptilia: Ichthyosauria)

Reconstructing ecological niche shifts during ontogeny in extinct animals with no living analogues is difficult without exceptional fossil collections. Here we demonstrate how a previously identified ontogenetic shift in the size and shape of the dentition in the early Toarcian ichthyosaur Stenopterygius quadriscissus accurately predicts a particular dietary shift. The smallest S. quadriscissus fed on small, burst‐swimming fishes, with a steady shift towards faster moving fish and cephalopods with increasing body size. Larger adult specimens appear to have been completely reliant on cephalopods, with fish completely absent from gut contents shortly after onset of sexual maturity. This is consistent with a previously proposed ontogenetic niche shift based on tooth shape and body size, corroborating the idea that dental ontogeny may be a useful predictor of dietary shifts in marine reptiles. Applying the theoretical framework used here to other extinct species will improve the resolution of palaeoecological reconstructions, where appropriate sample sizes exist.     

The Domestication Syndrome in Phoenix dactylifera Seeds: Toward the Identification of Wild Date Palm Populations

Investigating crop origins is a priority to understand the evolution of plants under domestication, develop strategies for conservation and valorization of agrobiodiversity and acquire fundamental knowledge for cultivar improvement. The date palm (Phoenix dactylifera L.) belongs to the genus Phoenix, which comprises 14 species morphologically very close, sometimes hardly distinguishable. It has been cultivated for millennia in the Middle East and in North Africa and constitutes the keystone of oasis agriculture. Yet, its origins remain poorly understood as no wild populations are identified. Uncultivated populations have been described but they might represent feral, i.e. formerly cultivated, abandoned forms rather than truly wild populations. In this context, this study based on morphometrics applied to 1625 Phoenix seeds aims to (1) differentiate Phoenix species and (2) depict the domestication syndrome observed in cultivated date palm seeds using other Phoenix species as a “wild” reference. This will help discriminate truly wild from feral forms, thus providing new insights into the evolutionary history of this species. Seed size was evaluated using four parameters: length, width, thickness and dorsal view surface. Seed shape was quantified using outline analyses based on the Elliptic Fourier Transform method. The size and shape of seeds allowed an accurate differentiation of Phoenix species. The cultivated date palm shows distinctive size and shape features, compared to other Phoenix species: seeds are longer and elongated. This morphological shift may be interpreted as a domestication syndrome, resulting from the long-term history of cultivation, selection and human-mediated dispersion. Based on seed attributes, some uncultivated date palms from Oman may be identified as wild. This opens new prospects regarding the possible existence and characterization of relict wild populations and consequently for the understanding of the date palm origins. Finally, we here describe a pipeline for the identification of the domestication syndrome in seeds that could be used in other crops.     

The Proximate Causes of Sexual Size Dimorphism in Phrynocephalus przewalskii

Sexual size dimorphism (SSD) is a common phenomenon and is a central topic in evolutionary biology. Recently, the importance of pursuing an ontogenetic perspective of SSD has been emphasized, to elucidate the proximate physiological mechanisms leading to its evolution. However, such research has seldom focused on the critical periods when males and females diverge. Using mark-recapture data, we investigated the development of SSD, sex-specific survivorship, and growth rates in Phrynocephalus przewalskii (Agamidae). We demonstrated that both male and female lizards are reproductively mature at age 10–11 months (including 5 months hibernation). Male-biased SSD in snout-vent length (SVL) was only found in adults and was fully expressed at age 11 months (June of the first full season of activity), just after sexual maturation. However, male-biased SSD in tail length (TL), hind-limb length (LL), and head width (HW) were fully expressed at age 9–10 months, just before sexual maturation. Analysis of age-specific linear growth rates identified sexually dimorphic growth during the fifth growth month (age 10–11 months) as the proximate cause of SSD in SVL. The males experienced higher mortality than females in the first 2 years and only survived better than females after SSD was well developed. This suggests that the critical period of divergence in the sizes of male and female P. przewalskii occurs between 10 and 11 months of age (May to June during the first full season of activity), and that the sexual difference in growth during this period is the proximate cause. However, the sexual difference in survivorship cannot explain the male-biased SSD in SVL. Our results indicate that performance-related characteristics, such as TL, HW, and LL diverged earlier than SVL. The physiological mechanisms underlying the different growth patterns of males and females may reflect different energy allocations associated with their different reproductive statuses.     

Genotyping validates the efficacy of photographic identification in a capture‐mark‐recapture study based on the head scale patterns of the prairie lizard (Sceloporus consobrinus)

Population studies often incorporate capture‐mark‐recapture (CMR) techniques to gather information on long‐term biological and demographic characteristics. A fundamental requirement for CMR studies is that an individual must be uniquely and permanently marked to ensure reliable reidentification throughout its lifespan. Photographic identification involving automated photographic identification software has become a popular and efficient noninvasive method for identifying individuals based on natural markings. However, few studies have (a) robustly assessed the performance of automated programs by using a double‐marking system or (b) determined their efficacy for long‐term studies by incorporating multi‐year data. Here, we evaluated the performance of the program Interactive Individual Identification System (I³S) by cross‐validating photographic identifications based on the head scale pattern of the prairie lizard (Sceloporus consobrinus) with individual microsatellite genotyping (N = 863). Further, we assessed the efficacy of the program to identify individuals over time by comparing error rates between within‐year and between‐year recaptures. Recaptured lizards were correctly identified by I³S in 94.1% of cases. We estimated a false rejection rate (FRR) of 5.9% and a false acceptance rate (FAR) of 0%. By using I³S, we correctly identified 97.8% of within‐year recaptures (FRR = 2.2%; FAR = 0%) and 91.1% of between‐year recaptures (FRR = 8.9%; FAR = 0%). Misidentifications were primarily due to poor photograph quality (N = 4). However, two misidentifications were caused by indistinct scale configuration due to scale damage (N = 1) and ontogenetic changes in head scalation between capture events (N = 1). We conclude that automated photographic identification based on head scale patterns is a reliable and accurate method for identifying individuals over time. Because many lizard or reptilian species possess variable head squamation, this method has potential for successful application in many species.     

Organelle Size Scaling of the Budding Yeast Vacuole Is Tuned by Membrane Trafficking Rates

Organelles serve as biochemical reactors in the cell, and often display characteristic scaling trends with cell size, suggesting mechanisms that coordinate their sizes. In this study, we measure the vacuole-cell size scaling trends in budding yeast using optical microscopy and a novel, to our knowledge, image analysis algorithm. Vacuole volume and surface area both show characteristic scaling trends with respect to cell size that are consistent among different strains. Rapamycin treatment was found to increase vacuole-cell size scaling trends for both volume and surface area. Unexpectedly, these increases did not depend on macroautophagy, as similar increases in vacuole size were observed in the autophagy deficient mutants atg1Δ and atg5Δ. Rather, rapamycin appears to act on vacuole size by inhibiting retrograde membrane trafficking, as the atg18Δ mutant, which is defective in retrograde trafficking, shows similar vacuole size scaling to rapamycin-treated cells and is itself insensitive to rapamycin treatment. Disruption of anterograde membrane trafficking in the apl5Δ mutant leads to complementary changes in vacuole size scaling. These quantitative results lead to a simple model for vacuole size scaling based on proportionality between cell growth rates and vacuole growth rates.     

New ankylosaurian material from the Upper Cretaceous of Transylvania

Ankylosaurian remains from the Transylvanian Basin, Romania, are extremely rare. More than 100 years after the discovery of the first and only better-known assemblage, namely the type material of Struthiosaurus transylvanicus, new ankylosaurian material has been discovered in the Maastrichtian of the Ha?eg Basin, as well as at another locality (Vurp?r), in the Transylvanian Basin, that is described here. The material consists of one tooth in a small jaw fragment (from the Ha?eg Basin) and at least two accummulations of associated, as well as several isolated, postcranial elements (from Vurp?r). No diagnostic elements are preserved that would overlap with the type of S. transylvanicus, so we cannot assign any of the new specimens to this species. The tooth shows marked differences compared to those of other anklyosaurs including S. austriacus and Hungarosaurus in having only six, more or less equally sized, apically pointed cusps separated by deep grooves. The postcranial material from Vurp?r represents at least three different individuals. The humerus is the most diagnostic element among the postcranial remains being most similar both in size and morphology to humeri referred to as Struthiosaurus from different European localities, thus here we refer the humerus and probably associated elements preserved in one assemblage to as cf. Struthiosaurus sp.; the remaining specimens from Vurp?r are retained as Nodosauridae indet. Histological studies have confirmed the adult nature of all sampled bones in the Vurp?r ankylosaur material suggesting that these fully grown animals were of similar size to Struthiosaurus, a small-bodied nodosaurid the ontogenetic status of which, however, has never been investigated histologically. The obviously diminished body size of the Transylvanian ankylosaurs compared to other members of the clade could be explained by insular dwarfism using the same histology-based argument as presented for Magyarosaurus.     

An ontogenetic perspective on the study of sexual dimorphism, phylogenetic variability, and allometry of the skull of European ground squirrel, Spermophilus citellus (Linnaeus, 1766)

Most studies of morphological variability in or among species are performed on adult specimens. However, it has been proven that knowledge of the patterns of size and shape changes and their covariation during ontogeny is of great value for the understanding of the processes that produce morphological variation. In this study, we investigated the patterns of sexual dimorphism, phylogenetic variability, and ontogenetic allometry in the Spermophilus citellus with geometric morphometrics applied to cross-sectional ontogenetic data of 189 skulls from three populations (originating from Burgenland, Banat, and Dojran) belonging to two phylogenetic lineages (the Northern and Southern). Our results indicate that sexual dimorphism in the ventral cranium of S. citellus is expressed only in skull size and becomes apparent just before or after the first hibernation because of accelerated growth in juvenile males. Sexes had the same pattern of ontogenetic allometry. Populations from Banat and Dojran, belonging to different phylogroups, were the most different in size but had the most similar adult skull shape. Phylogenetic relations among populations, therefore, did not reflect skull morphology, which is probably under a significant influence of ecological factors. Populations had parallel allometric trajectories, indicating that alterations in development probably occur prenatally. The species’ allometric relations during cranial growth showed characteristic nonlinear trajectories in the two northern populations, with accelerated shape changes in juveniles and continued but almost isometric growth in adults. The adult cranial shape was reached before sexual maturity of both sexes and adult size after sexual maturity. The majority of shape changes during growth are probably correlated with the shift from a liquid to a solid diet and to a lesser degree due to allometric scaling, which explained only 20 % of total shape variation. As expected, viscerocranial components grew with positive and neurocranial with negative allometry.     

Prey Capture Ecology of the Cubozoan Carukia barnesi

Adult Carukia barnesi medusae feed predominantly on larval fish; however, their mode of prey capture seems more complex than previously described. Our findings revealed that during light conditions, this species extends its tentacles and ‘twitches’ them frequently. This highlights the lure-like nematocyst clusters in the water column, which actively attract larval fish that are consequently stung and consumed. This fishing behavior was not observed during dark conditions, presumably to reduce energy expenditure when they are not luring visually oriented prey. We found that larger medusae have longer tentacles; however, the spacing between the nematocyst clusters is not dependent on size, suggesting that the spacing of the nematocyst clusters is important for prey capture. Additionally, larger specimens twitch their tentacles more frequently than small specimens, which correlate with their recent ontogenetic prey shift from plankton to larval fish. These results indicate that adult medusae of C. barnesi are not opportunistically grazing in the water column, but instead utilize sophisticated prey capture techniques to specifically target larval fish.     

Oligomeric Size of the M2 Muscarinic Receptor in Live Cells as Determined by Quantitative Fluorescence Resonance Energy Transfer

Fluorescence resonance energy transfer (FRET), measured by fluorescence intensity-based microscopy and fluorescence lifetime imaging, has been used to estimate the size of oligomers formed by the M₂ muscarinic cholinergic receptor. The approach is based on the relationship between the apparent FRET efficiency within an oligomer of specified size (n) and the pairwise FRET efficiency between a single donor and a single acceptor (E). The M₂ receptor was fused at the N terminus to enhanced green or yellow fluorescent protein and expressed in Chinese hamster ovary cells. Emission spectra were analyzed by spectral deconvolution, and apparent efficiencies were estimated by donor-dequenching and acceptor-sensitized emission at different ratios of enhanced yellow fluorescent protein-M₂ receptor to enhanced green fluorescent protein-M₂ receptor. The data were interpreted in terms of a model that considers all combinations of donor and acceptor within a specified oligomer to obtain fitted values of E as follows: n = 2, 0.495 ± 0.019; n = 4, 0.202 ± 0.010; n = 6, 0.128 ± 0.006; n = 8, 0.093 ± 0.005. The pairwise FRET efficiency determined independently by fluorescence lifetime imaging was 0.20–0.24, identifying the M₂ receptor as a tetramer. The strategy described here yields an explicit estimate of oligomeric size on the basis of fluorescence properties alone. Its broader application could resolve the general question of whether G protein-coupled receptors exist as dimers or larger oligomers. The size of an oligomer has functional implications, and such information can be expected to contribute to an understanding of the signaling process.     

Morphology and postnatal ontogeny of the cranial endocast and paranasal sinuses of capybara (Hydrochoerus hydrochaeris), the largest living rodent

Recent studies have analyzed and described the endocranial cavities of caviomorph rodents. However, no study has documented the changes in the morphology and relative size of such cavities during ontogeny. Expecting to contribute to the discussion of the endocranial spaces of extinct caviomorphs, we aimed to characterize the cranial endocast morphology and paranasal sinuses of the largest living rodent, Hydrochoerus hydrochaeris, by focusing on its ontogenetic growth patterns. We analyzed 12 specimens of different ontogenetic stages and provided a comparison with other cavioids. Our study demonstrates that the adult cranial endocast of H. hydrochaeris is characterized by olfactory bulbs with an irregular shape, showing an elongated olfactory tract without a clear circular fissure, a marked temporal region that makes the endocast with rhombus outline, and gyrencephaly. Some of these traits change as the brain grows. The cranial pneumatization is present in the frontal and lacrimal bones. We identified two recesses (frontal and lacrimal) and one sinus (frontal). These pneumatic cavities increase their volume as the cranium grows, covering the cranial region of the cranial endocast. The encephalization quotient was calculated for each specimen, demonstrating that it decreases as the individual grows, being much higher in younger specimens than in adults. Our results show that the ontogenetic stage can be a confounding factor when it comes to the general patterns of encephalization of extinct rodents, reinforcing the need for paleobiologists to take the age of the specimens into account in future studies on this subject to avoid age-related biases.     

Life-History Traits of the Miocene Hipparion concudense (Spain) Inferred from Bone Histological Structure

Histological analyses of fossil bones have provided clues on the growth patterns and life history traits of several extinct vertebrates that would be unavailable for classical morphological studies. We analyzed the bone histology of Hipparion to infer features of its life history traits and growth pattern. Microscope analysis of thin sections of a large sample of humeri, femora, tibiae and metapodials of Hipparion concudense from the upper Miocene site of Los Valles de Fuentidueña (Segovia, Spain) has shown that the number of growth marks is similar among the different limb bones, suggesting that equivalent skeletochronological inferences for this Hipparion population might be achieved by means of any of the elements studied. Considering their abundance, we conducted a skeletechronological study based on the large sample of third metapodials from Los Valles de Fuentidueña together with another large sample from the Upper Miocene locality of Concud (Teruel, Spain). The data obtained enabled us to distinguish four age groups in both samples and to determine that Hipparion concudense tended to reach skeletal maturity during its third year of life. Integration of bone microstructure and skeletochronological data allowed us to identify ontogenetic changes in bone structure and growth rate and to distinguish three histologic ontogenetic stages corresponding to immature, subadult and adult individuals. Data on secondary osteon density revealed an increase in bone remodeling throughout the ontogenetic stages and a lesser degree thereof in the Concud population, which indicates different biomechanical stresses in the two populations, likely due to environmental differences. Several individuals showed atypical growth patterns in the Concud sample, which may also reflect environmental differences between the two localities. Finally, classification of the specimens’ age within groups enabled us to characterize the age structure of both samples, which is typical of attritional assemblages.