Functional and phylogenetic implications of molar flare variation in Miocene hominoids

Comparative analyses of molar shape figure prominently in Miocene hominoid evolutionary studies, and incomplete understanding of functional and phylogenetic influences on molar shape variation can have direct consequences for the interpretation of fossil taxa. Molar flare is a shape trait whose polarity, phylogenetic distribution, and functional significance have been sources of contention. To clarify the determinants of molar flare variation in the hominoid radiation, a combination of statistical methods was employed to investigate the effects of diet, phylogeny, and geologic age upon several measures of molar shape, to identify interactions among these factors, and to estimate their relative influence. Classic indices of molar crown shape and cusp relief are highly significantly associated with diet and show no clear phylogenetic or temporal patterning. Correlations with diet are insignificant when phylogenetic effects are controlled, a result which is interpreted as an artifact of the distribution of folivory in the Miocene hominoid radiation. Possession of pronounced molar flare was found to be the primitive condition for Miocene hominoids, but molar flare reduction cannot be considered a crown hominoid synapomorphy. Molar flare is strongly correlated with geologic age but differs significantly among dietary categories when the effects of time are controlled. Among contemporaneous taxa, hard-object feeders consistently show the highest levels of flare. Molar flare reduction is hypothesized to arise from realignment of cusp positions to maximize molar shearing and increase working occlusal surface area, while variation in flare among contemporaneous taxa may be due, at least in part, to enamel thickness variation. The pronounced molar flare of Otavipithecus is interpreted as a primitive retention, although alternative dietary and phylogenetic interpretations cannot be excluded. A dramatic reversal of molar flare reduction in Mio-Pliocene hominins is interpreted as a synapomorphy of the crown hominin clade, thus supporting the hominin status of the Lukeino hominine. The last common ancestor of the Pan-Homo clade is predicted to have possessed relatively non-flaring molars, and implications of this hypothesis for early hominin recognition are discussed.     

The Diet of the Brown Trout Salmo trutta (L.) during the Reproductive Period: Size‐Related and Sexual Effects

We investigated the autumnal diet of the brown trout Salmo trutta, in a Prepyrenean stream (NW Iberian Peninsula) focusing on intraspecific dietary differences related to size and sex. The diet of trout included 18 types of prey, with Plecoptera and Ephemeroptera nymphs and Diptera larvae as the most consumed taxa. Large trout ate larger prey, than did small trout, and also increased the consumption of terrestrial‐surface prey with respect to aquatic‐benthic prey. As terrestrial‐surface preys were larger than aquatic‐benthic prey, the size‐related differences in the diet of trout were related to gape‐limitations. Although male and female trout did not differ in size, we found that males foraged on a more diverse type of prey than females, probably owing to male territoriality during the reproductive period. This study provides new evidence of dietary plasticity in the brown trout and confirms the importance of local dietary studies to better understand factors which drive trophic ecology of predators. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Relevance of aquatic environments for hominins: a case study from Trinil (Java, Indonesia)

Knowledge about dietary niche is key to understanding hominin evolution, since diet influences body proportions, brain size, cognition, and habitat preference. In this study we provide ecological context for the current debate on modernity (or not) of aquatic resource exploitation by hominins. We use the Homo erectus site of Trinil as a case study to investigate how research questions on possible dietary relevance of aquatic environments can be addressed. Faunal and geochemical analysis of aquatic fossils from Trinil Hauptknochenschicht (HK) fauna demonstrate that Trinil at ∼1.5 Ma contained near-coastal rivers, lakes, swamp forests, lagoons, and marshes with minor marine influence, laterally grading into grasslands. Trinil HK environments yielded at least eleven edible mollusc species and four edible fish species that could be procured with no or minimal technology. We demonstrate that, from an ecological point of view, the default assumption should be that omnivorous hominins in coastal habitats with catchable aquatic fauna could have consumed aquatic resources. The hypothesis of aquatic exploitation can be tested with taphonomic analysis of aquatic fossils associated with hominin fossils. We show that midden-like characteristics of large bivalve shell assemblages containing Pseudodon and Elongaria from Trinil HK indicate deliberate collection by a selective agent, possibly hominin.     

Riparian contributions to the diet of terapontid grunters (Pisces: Terapontidae) in wet–dry tropical rivers

The diets of terapontid assemblages in 22 catchments across Australia's wet–dry tropics were investigated in relation to the direct use of terrestrial‐riparian inputs, as well as the role of ontogeny and morphology in mediating consumption of allocthonous material. The diet of several species was restricted almost entirely to instream trophic resources throughout their life history. In contrast, ontogenetic diet shifts towards increasing consumption of terrestrial prey types were a prominent feature of the dietary ecology of some terapontids, with collective allocthonous dietary items making a significant contribution (up to 42%) to diet in larger size classes of several species. For those species consuming terrestrial‐riparian material in their diet, terrestrial invertebrates were the most common prey item; however, terrestrial vegetation, principally riparian fruits, and terrestrial vertebrates were also important dietary inclusions in the larger size classes of particular species. A large mouth gape was the morphological feature most strongly associated with consumption of terrestrial food resources within the Terapontidae. Results indicate that the direct consumption of terrestrially derived food sources in northern Australian aquatic systems may be more important than previously asserted, and that additional research is required to better clarify the role of terrestrial subsidies to these ecosystems.     

Biomechanical evolution of solid bones in large animals: a microanatomical investigation

Graviportal taxa show an allometric increase of the cross‐sectional area of supportive bones and are assumed to display microanatomical changes associated with an increase in bone mass. This evokes osteosclerosis (i.e. an increase in bone compactness observed in some aquatic amniotes). The present study investigates the changes in bones' microanatomical organization associated with graviportality and how comparable they are with aquatically acquired osteosclerosis aiming to better understand the adaptation of bone to the different associated functional requirements. Bones of graviportal taxa show microanatomical changes that are not solely attributable to allometry. They display a thicker cortex and a proportionally smaller medullary cavity, with a wider transition zone between these domains. This inner cancellous structure may enable to better enhance energy absorption and marrow support. Moreover, the cross‐sectional geometric parameters indicate increased resistance to stresses engendered by bending and torsion, as well as compression. Adaptation to a graviportal posture should be taken into consideration when analyzing possibly amphibious taxa with a terrestrial‐like morphology. This is particularly important for palaeoecological inferences about large extinct tetrapods that might have been amphibious and, more generally, for the study of early stages of adaptation to an aquatic life in amniotes.     

Isotopic dietary reconstructions of Pliocene herbivores at Laetoli: Implications for early hominin paleoecology

Major morphological and behavioral innovations in early human evolution have traditionally been viewed as responses to conditions associated with increasing aridity and the development of extensive grassland-savanna biomes in Africa during the Plio-Pleistocene. Interpretations of paleoenvironments at the Pliocene locality of Laetoli in northern Tanzania have figured prominently in these discussions, primarily because early hominins recovered from Laetoli are generally inferred to be associated with grassland, savanna or open woodland habitats. As these reconstructions effectively extend the range of habitat preferences inferred for Pliocene hominins, and contrast with interpretations of predominantly woodland and forested ecosystems at other early hominin sites, it is worth reevaluating the paleoecology at Laetoli utilizing a new approach. Isotopic analyses were conducted on the teeth of twenty-one extinct mammalian herbivore species from the Laetolil Beds (∼ 4.3–3.5 Ma) and Upper Ndolanya Beds (∼ 2.7–2.6 Ma) to determine their diet, as well as to investigate aspects of plant physiognomy and climate. Enamel samples were obtained from multiple localities at different stratigraphic levels in order to develop a high-resolution spatio-temporal framework for identifying and characterizing dietary and ecological change and variability within the succession. In general, dietary signals at Laetoli suggest heterogeneous ecosystems with both C₃ and C₄ dietary plants available that could support grassland, woodland, and forested communities. All large-bodied herbivores analyzed yielded dietary signatures indicating mixed grazing/browsing strategies or exclusive reliance on C₃ browse, more consistent with wooded than grassland-savanna biomes. There are no clear isotopic patterns documenting shifting ecology within the Laetolil Beds or between the Laetolil and overlying Upper Ndolanya Beds, although limited data from the U. Ndolanya Beds constrains interpretations. Comparison of the results from Laetoli with isotopic enamel profiles of other African fossil and modern communities reveals significant differences in dietary patterns. Relative to extant taxa in related lineages, carbon isotopic ranges of a number of Laetoli fossil herbivores are anomalous, indicating significantly more generalized intermediate C₃/C₄ feeding behaviors, perhaps indicative of dietary niches and habitat types with no close modern analogs. Enamel oxygen isotope ranges of fossil taxa from Laetoli are consistently more ¹⁸O depleted than modern E. African herbivores, possibly indicating more humid conditions during that interval in the past. These data have important implications for reconstructing dietary trajectories of mammalian herbivore lineages, as well as the evolution of ecosystems in East Africa. Isotopic analyses of similar or related taxa at other hominin fossil sites yield signatures generally consistent with Laetoli, suggesting that mammalian communities in East Africa were sampling ecosystems with similar proportions of browse and grass. Collectively, the isotopic dietary signatures indicate heterogeneous habitats with significant wooded or forested components in the Laetoli area during deposition of the Laetolil and Upper Ndolanya Beds. Early hominin foraging activity in this interval may have included access to forest or woodland biomes within this ecosystem, complicating traditional interpretations linking early human evolutionary innovations with a shift to savanna habitats.     

Spatial and seasonal variability in the trophic role of aquatic insects: An assessment of functional feeding group applicability

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The diet and digestive energetics of an Australian short-necked turtle, Emydura macquarii

We described the diet of Emydura macquarii, an omnivorous turtle from south-eastern Australia, compared its digestive performance on diets of fish or plants at two temperatures, and related how both diet and temperature affect its food selection in nature. Filamentous algae constituted 61% of the stomach content of E. macquarii. The turtles rarely fed on motile prey, but selected carrion from the lagoon bottom and terrestrial insects (Diptera, Hymenoptera and Coleoptera) trapped on the surface of the water. Digestive efficiency of E. macquarii was affected little by body temperature, in contrast to consumption rates and rates of passage which were strongly influenced by both temperature and diet. In combination, these responses resulted in a slower rate of digestion at 20 degrees C than at 30 degrees C. Digestive efficiency of E. macquarii on a herbivorous diet at 30 degrees C (49%) was about half that of turtles on a carnivorous diet (91%), but they had longer transit times (118 h on the plant diet versus 70 h). Lower consumption rates and longer mean retention times in turtles fed plants compared those fed fish relate to slower digestive processing of the plant. Rapid processing and higher consumption rates of fish by E. macquarii resulted in higher energy gains compared to turtles consuming plants (almost 100 times more energy at 30 degrees C). The laboratory results suggest that fish carrion and aquatic and terrestrial invertebrates are probably essential dietary items of E. macquarii in the wild, because its metabolic requirements cannot be met from aquatic macrophytes alone.     

Microanatomical diversity of the humerus and lifestyle in lissamphibians

A study of body size and the compactness profile parameters of the humerus of 37 species of lissamphibians demonstrates a relationship between lifestyle (aquatic, amphibious or terrestrial) and bone microstructure. Multiple linear regressions and variance partitioning with Phylogenetic eigenVector Regressions reveal an ecological and a phylogenetic signal in some body size and compactness profile parameters. Linear discriminant analyses segregate the various lifestyles (aquatic vs. amphibious or terrestrial) with a success rate of up to 89.2%. The models built from data on the humerus discriminate aquatic taxa relatively well from the other taxa. However, like previous models built from data on the radius of amniotes or on the femur of lissamphibians, the new models do not discriminate amphibious taxa from terrestrial taxa on the basis of body size or compactness profile data. To make our inference method accessible, spreadsheets (see supplementary material on the website), which allow anyone to infer a lissamphibian lifestyle solely from body size and bone compactness parameters, were produced. No such easy implementation of habitat inference models is found in earlier papers on this topic.     

Polyunsaturated fatty acids in fish tissues more closely resemble algal than terrestrial diet sources

The River Continuum Concept implies that consumers in headwater streams have greater dietary access to terrestrial basal resources, but recent studies have highlighted the dietary importance of high-quality algae. Algae provide consumers with physiologically important omega-3 (n-3) polyunsaturated fatty acids (PUFA), particularly eicosapentaenoic acid (EPA). However, terrestrial plants and most benthic stream algae lack the long-chain (LC) n-3 PUFA docosahexaenoic acid (DHA, 22:6n-3), which is essential for neural development in fish and other vertebrates. We sampled subalpine streams to investigate how the PUFA composition of neural (brain and eyes), muscle, and liver tissues of freshwater fish is related to their potential diets (macroinvertebrates, epilithon, fresh and conditioned terrestrial leaves). The PUFA composition of consumers was more similar to epilithon than to terrestrial leaves. Storage lipids of eyes most closely resembled dietary PUFA (aquatic invertebrates and algae). However, DHA and arachidonic acid (ARA, 20:4n-6) were not directly available in the diet but abundant in organs. This implies that algal PUFA were selectively retained or were produced internally via enzymatic PUFA conversion by aquatic consumers. This field study demonstrates the nutritional importance of algal PUFA for neural organs in aquatic consumers of headwater regions.

     

Differences in dispersal- and colonization-related traits between taxa from the freshwater and the terrestrial realm

The aquatic and terrestrial realms differ in many physical properties that not only require specific physiological adaptations but also cause differences in dispersal options. We thus expect that life-history traits related to dispersal and colonization are under selection pressure because freshwater habitats are more isolated and thus more difficult to reach. We compared traits from European databases of three taxonomic groups along the passive–active dispersal gradient: plants (Plantes), snails (Mollusca: Gastropoda: Prosobranchia et Pulmonata) and hoverflies (Diptera: Syrphidae), all of which have both terrestrial and freshwater species (plants and snails) or early life stages (hoverflies). Aquatic taxa seem to be more successful long-distance dispersers than are terrestrial taxa. Our analysis also revealed lower numbers of seeds or eggs produced in the aquatic habitats. However, aquatic taxa often allocate resources to offspring guarding (vegetative propagules in plants, egg capsules in snails) and breeding-site selection (syrphids). Colonization of the aquatic realm is reinforced by increases in life span (plants), clonal spread (plants), shorter generation times (snails), selfing ability (marginal effect in pulmonate snails) or paedogenesis (two incidences in hoverflies, needs further studies). Probably, the variety of strategies reflects the different evolutionary backgrounds that elicit different combinations of trade-offs, but all traits also might increase invasibility of species.     

Chimpanzee fauna isotopes provide new interpretations of fossil ape and hominin ecologies

Carbon and oxygen stable isotopes within modern and fossil tooth enamel record the aspects of an animal''s diet and habitat use. This investigation reports the first isotopic analyses of enamel from a large chimpanzee community and associated fauna, thus providing a means of comparing fossil ape and early hominin palaeoecologies with those of a modern ape. Within Kibale National Park forest, oxygen isotopes differentiate primate niches, allowing for the first isotopic reconstructions of degree of frugivory versus folivory as well as use of arboreal versus terrestrial resources. In a comparison of modern and fossil community isotopic profiles, results indicate that Sivapithecus, a Miocene ape from Pakistan, fed in the forest canopy, as do chimpanzees, but inhabited a forest with less continuous canopy or fed more on leaves. Ardipithecus, an early hominin from Ethiopia, fed both arboreally and terrestrially in a more open habitat than inhabited by chimpanzees.     

Diet and habit explain head-shape convergences in natricine snakes

The concept of ecomorphs, whereby species with similar ecologies have similar phenotypes regardless of their phylogenetic relatedness, is often central to discussions regarding the relationship between ecology and phenotype. However, some aspects of the concept have been questioned, and sometimes species have been grouped as ecomorphs based on phenotypic similarity without demonstrating ecological similarity. Within snakes, similar head shapes have convergently evolved in species living in comparable environments and/or with similar diets. Therefore, ecomorphs could exist in some snake lineages, but this assertion has rarely been tested for a wide-ranging group within a single framework. Natricine snakes (Natricinae) are ecomorphologically diverse and currently distributed in Asia, Africa, Europe and north-central America. They are primarily semiaquatic or ground-dwelling terrestrial snakes, but some are aquatic, burrowing or aquatic and burrowing in habit and may be generalist or specialist in diet. Thus, natricines present an interesting system to test whether snakes from different major habit categories represent ecomorphs. We quantify morphological similarity and disparity in head shape among 191 of the ca. 250 currently recognized natricine species and apply phylogenetic comparative methods to test for convergence. Natricine head shape is largely correlated with habit, but in some burrowers is better explained by dietary specialism. Convergence in head shape is especially strong for aquatic burrowing, semiaquatic and terrestrial ecomorphs and less strong for aquatic and burrowing ecomorphs. The ecomorph concept is useful for understanding natricine diversity and evolution, though would benefit from further refinement, especially for aquatic and burrowing taxa.     

Phylogenetic tree of vascular plants reveals the origins of aquatic angiosperms

Although aquatic plants are discussed as a unified biological group, they are phylogenetically well dispersed across the angiosperms. In this study, we annotated the aquatic taxa on the tree of vascular plants, and extracted the topology of these aquatic lineages to construct the tree of aquatic angiosperms. We also reconstructed the ancestral areas of aquatic families. We found that aquatic angiosperms could be divided into two different categories: the four aquatic orders and the aquatic taxa in terrestrial orders. Aquatic lineages evolved early in the radiation of angiosperms, both in the orders Nymphaeales and Ceratophyllales and among basal monocots (Acorales and Alismatales). These aquatic orders do not have any extant terrestrial relatives. They originated from aquatic habitats during the Early Cretaceous. Asia would have been one of the centers for early diversification of aquatic angiosperms. The aquatic families within terrestrial orders may originate from other areas besides Asia, such as America or Australia. The lineages leading to extant angiosperms diversified early in underexploited freshwater habitats. The four extant aquatic orders were relicts of an early radiation of angiosperm in aquatic environments. Their extinct ancestors might be aquatic early angiosperms.     

Dietary Inference from Upper and Lower Molar Morphology in Platyrrhine Primates

The correlation between diet and dental topography is of importance to paleontologists seeking to diagnose ecological adaptations in extinct taxa. Although the subject is well represented in the literature, few studies directly compare methods or evaluate dietary signals conveyed by both upper and lower molars. Here, we address this gap in our knowledge by comparing the efficacy of three measures of functional morphology for classifying an ecologically diverse sample of thirteen medium- to large-bodied platyrrhines by diet category (e.g., folivore, frugivore, hard object feeder). We used Shearing Quotient (SQ), an index derived from linear measurements of molar cutting edges and two indices of crown surface topography, Occlusal Relief (OR) and Relief Index (RFI). Using SQ, OR, and RFI, individuals were then classified by dietary category using Discriminate Function Analysis. Both upper and lower molar variables produce high classification rates in assigning individuals to diet categories, but lower molars are consistently more successful. SQs yield the highest classification rates. RFI and OR generally perform above chance. Upper molar RFI has a success rate below the level of chance. Adding molar length enhances the discriminatory power for all variables. We conclude that upper molar SQs are useful for dietary reconstruction, especially when combined with body size information. Additionally, we find that among our sample of platyrrhines, SQ remains the strongest predictor of diet, while RFI is less useful at signaling dietary differences in absence of body size information. The study demonstrates new ways for inferring the diets of extinct platyrrhine primates when both upper and lower molars are available, or, for taxa known only from upper molars. The techniques are useful in reconstructing diet in stem representatives of anthropoid clade, who share key aspects of molar morphology with extant platyrrhines.     

Fluxes of terrestrial and aquatic carbon by emergent mosquitoes: a test of controls and implications for cross-ecosystem linkages

Adult aquatic insects are a common resource for many terrestrial predators, often considered to subsidize terrestrial food webs. However, larval aquatic insects themselves consume both aquatic primary producers and allochthonous terrestrial detritus, suggesting that adults could provide aquatic subsidy and/or recycled terrestrial energy to terrestrial consumers. Understanding the source of carbon (aquatic vs. terrestrial) driving aquatic insect emergence is important for predicting magnitude of emergence and effects on recipient food web dynamics; yet direct experimental tests of factors determining source are lacking. Here, we use Culex mosquitoes in experimental pools as an exemplar to test how variation in general factors common to aquatic systems (terrestrial plant inputs and light) may alter the source and amount of energy exported to terrestrial ecosystems in adult aquatic insects that rely on terrestrial resources as larvae. We found strong sequential effects of terrestrial plant inputs and light on aquatic insect oviposition, diet, and emergence of Culex mosquitoes. Ovipositing mosquitoes laid ~3 times more egg masses in high terrestrial input pools under low light conditions. This behavior increased adult emergence from pools under low light conditions; however, high input pools (which had the highest mosquito densities) showed low emergence rates due to density-dependent mortality. Mosquito diets consisted mainly of terrestrial resources (~70–90 %). As a result, the amount of aquatic carbon exported from pools by mosquitoes during the experiment was ~18 times higher from low versus high light pools, while exports of terrestrial carbon peaked from pools receiving intermediate levels of inputs (3–6 times higher) and low light (~6 times higher). Our results suggest that understanding the interplay among terrestrial plant inputs, light availability and biotic responses of aquatic insects may be key in predicting source and magnitude of emergence, and thus the strength and effects of aquatic–terrestrial linkages in freshwater systems.     

Relating body size to the role of aquatic subsidies for the riparian spider <Emphasis Type="Italic">Nephila clavata</Emphasis>

We examined the relationship between body size of the riparian spider Nephila clavata and the contribution of allochthonous (aquatic insects) and autochthonous (terrestrial insects) sources to its diet using stable isotope analysis. During the study period from July to September, the body size of the females increased remarkably (about 60-fold) but that of males remained small. The biomass of both aquatic and terrestrial insects trapped on the spider webs increased with spider size, with the biomass of the former ranging between 30 and 70% of that of the terrestrial insects. The average relative contribution of aquatic insects to the diet of the spiders, calculated from δ¹³C values, was 40–50% in spiders in the early juvenile and juvenile stages, 35% in adult males and 4% in adult females. There was a significant negative relationship between the relative contribution of aquatic insects and body size of the female spiders. We conclude that aquatic insects might be an important seasonal dietary subsidy for small spiders and that these allochthonous subsidies may facilitate the growth of riparian spiders, which may in turn enable the spiders to feed on larger prey.     

Potential alteration of cross‐ecosystem resource subsidies by an invasive aquatic macroinvertebrate: implications for the terrestrial food web

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Importance of terrestrial subsidies for native brook trout in Appalachian intermittent streams

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Combining isotopic and ecomorphological data to refine bovid paleodietary reconstruction: a case study from the Makapansgat Limeworks hominin locality

The relationship between environmental change and hominin evolution remains obscure. For the most part, this stems from the difficulty of reconstructing ancient hominin habitats. Bovids are among the most frequently utilized paleoenvironmental indicators, but little is known about the habitat preferences of extinct taxa. It is generally assumed that fossil bovids both ate the same things and occupied the same habitats as their closest extant relatives. We test the first part of this assumption by reconstructing the diets of seven bovids from Makapansgat Limeworks, South Africa. Since diet and habitat are linked, these reconstructions have implications for our understanding of fossil bovid habitat tolerances. Ecomorphological and stable carbon isotope analyses are employed, allowing us to take advantage of the strengths and overcome the weaknesses of both. In most cases, fossil bovids did have similar diets to their extant relatives, and probably occupied similar habitats. Gazella vanhoepeni and Aepyceros sp., however, were almost exclusive browsers, and not mixed feeders like their living counterparts.