Diversity in the later Paleogene proboscidean radiation: a small barytheriid from the Oligocene of Dhofar Governorate,Sultanate of Oman

Despite significant recent improvements to our understanding of the early evolution of the Order Proboscidea (elephants and their extinct relatives), geographic sampling of the group’s Paleogene fossil record remains strongly biased, with the first ~30 million years of proboscidean evolution documented solely in near-coastal deposits of northern Africa. The considerable morphological disparity that is observable among the late Eocene and early Oligocene proboscideans of northern Africa suggests that other, as yet unsampled, parts of Afro-Arabia might have served as important centers for the early diversification of major proboscidean clades. Here we describe the oldest taxonomically diagnostic remains of a fossil proboscidean from the Arabian Peninsula, a partial mandible of Omanitherium dhofarensis (new genus and species), from near the base of the early Oligocene Shizar Member of the Ashawq Formation, in the Dhofar Governorate of the Sultanate of Oman. The molars and premolars of Omanitherium are morphologically intermediate between those of Arcanotherium and Barytherium from northern Africa, but its specialized lower incisors are unlike those of other known Paleogene proboscideans in being greatly enlarged, high-crowned, conical, and tusk-like. Omanitherium is consistently placed close to late Eocene Barytherium in our phylogenetic analyses, and we place the new genus in the Family Barytheriidae. Some features of Omanitherium, such as tusk-like lower second incisors, the possible loss of the lower central incisors, an enlarged anterior mental foramen, and inferred elongate mandibular symphysis and diminutive P₂, suggest a possible phylogenetic link with Deinotheriidae, an extinct family of proboscideans whose origins have long been mysterious.     

Contribution of anadromous fish to the diet of European catfish in a large river system

Many anadromous fish species, when migrating from the sea to spawn in fresh waters, can potentially be a valuable prey for larger predatory fish, thereby efficiently linking these two ecosystems. Here, we assess the contribution of anadromous fish to the diet of European catfish (Silurus glanis) in a large river system (Garonne, southwestern France) using stable isotope analysis and allis shad (Alosa alosa) as an example of anadromous fish. Allis shad caught in the Garonne had a very distinct marine δ¹³C value, over 8‰ higher after lipid extraction compared to the mean δ¹³C value of all other potential freshwater prey fish. The δ¹³C values of European catfish varied considerably between these two extremes and some individuals were clearly specializing on freshwater prey, whereas others specialized on anadromous fish. The mean contribution of anadromous fish to the entire European catfish population was estimated to be between 53% and 65%, depending on the fractionation factor used for δ¹³C.     

The hunter becomes the hunted: when cleptobiotic insects are captured by their target ants

Here we show that trying to rob prey (cleptobiosis) from a highly specialized predatory ant species is risky. To capture prey, Allomerus decemarticulatus workers build gallery-shaped traps on the stems of their associated myrmecophyte, Hirtella physophora. We wondered whether the frequent presence of immobilized prey on the trap attracted flying cleptoparasites. Nine social wasp species nest in the H. physophora foliage; of the six species studied, only Angiopolybia pallens rob prey from Allomerus colonies. For those H. physophora not sheltering wasps, we noted cleptobiosis by stingless bees (Trigona), social wasps (A. pallens and five Agelaia species), assassin bugs (Reduviidae), and flies. A relationship between the size of the robbers and their rate of capture by ambushing Allomerus workers was established for social wasps; small wasps were easily captured, while the largest never were. Reduviids, which are slow to extract their rostrum from prey, were always captured, while Trigona and flies often escaped. The balance sheet for the ants was positive vis-à-vis the reduviids and four out of the six social wasp species. For the latter, wasps began by cutting up parts of the prey’s abdomen and were captured (or abandoned the prey) before the entire abdomen was retrieved so that the total weight of the captured wasps exceeded that of the prey abdomens. For A. pallens, we show that the number of individuals captured during attempts at cleptobiosis increases with the size of the Allomerus’ prey.     

Salt glands in the Jurassic metriorhynchid <Emphasis Type="Italic">Geosaurus</Emphasis>: implications for the evolution of osmoregulation in Mesozoic marine crocodyliforms

The presence of salt-excreting glands in extinct marine sauropsids has been long suspected based on skull morphology. Previously, we described for the first time the natural casts of salt-excreting glands in the head of the Jurassic metriorhynchid crocodyliform Geosaurus araucanensis from the Tithonian of the Vaca Muerta Formation in the Neuquén Basin (Argentina). In the present study, salt-excreting glands are identified in three new individuals (adult, a sub-adult and a juvenile) referable to the same species. New material provides significant information on the salt glands form and function and permit integration of evolutionary scenarios proposed on a physiological basis in extant taxa with evidence from the fossil record. G. araucanensis represents an advanced stage of the basic physiological model to marine adaptations in reptiles. G. araucanensis salt glands were hypertrophied. On this basis, it can be hypothesized that these glands had a high excretory capability. This stage implies that G. araucanensis (like extant pelagic reptiles, e.g. cheloniids) could have maintained constant plasma osmolality even when seawater or osmoconforming prey were ingested. A gradual model of marine adaptation in crocodyliforms based on physiology (freshwater to coastal/estuarine to estuarine /marine to pelagic life) is congruent with the phylogeny of crocodyliforms based on skeletal morphology. The fossil record suggests that the stage of marine pelagic adaptation was achieved by the Early Middle Jurassic. Salt gland size in the juvenile suggests that juveniles were, like adults, pelagic.     

Armoured spiderman: morphological and behavioural adaptations of a specialised araneophagous predator (Araneae: Palpimanidae)

In a predator–prey system where both intervenients come from the same taxon, one can expect a strong selection on behavioural and morphological traits involved in prey capture. For example, in specialised snake-eating snakes, the predator is unaffetced by the venom of the prey. We predicted that similar adaptations should have evolved in spider-eating (araneophagous) spiders. We investigated potential and actual prey of two Palpimanus spiders (P. gibbulus, P. orientalis) to support the prediction that these are araneophagous predators. Specific behavioural adaptations were investigated using a high-speed camera during staged encounters with prey, while morphological adaptations were investigated using electron microscopy. Both Palpimanus species captured a wide assortment of spider species from various guilds but also a few insect species. Analysis of the potential prey suggested that Palpimanus is a retreat-invading predator that actively searches for spiders that hide in a retreat. Behavioural capture adaptations include a slow, stealthy approach to the prey followed by a very fast attack. Morphological capture adaptations include scopulae on forelegs used in grabbing prey body parts, stout forelegs to hold the prey firmly, and an extremely thick cuticle all over the body preventing injury from a counter bite of the prey. Palpimanus overwhelmed prey that was more than 200% larger than itself. In trials with another araneophagous spider, Cyrba algerina (Salticidae), Palpimanus captured C. algerina in more than 90% of cases independent of the size ratio between the spiders. Evidence indicates that both Palpimanus species possesses remarkable adaptations that increase its efficiency in capturing spider prey.     

Cytogenetic analysis of 1375 amniotic fluid specimens from pregnancies with gestational age less than 14 weeks

Our laboratory has received 1375 early amniotic fluid (EA) specimens during the past 5-year period for cytogenetics analysis. The gestational ages of the EA specimens were less than 14 weeks as estimated by ultrasound. The average volume of specimen received was 16 ml. Specimens were typically received in two collection tubes and cultured in Chang A and in supplemented MEM media using the in situ technique. Of the 1375 EA specimens received, 1356 were successfully cultured and yielded results. Abnormal results were found in 67 (4.9 per cent) of the cases. Nineteen specimens (1.4 per cent) failed to yield a result. The mean turn-around time (TAT) for all EA specimens was 8.28 days. In 1991, the average TAT for the EA specimens was 8.00 days compared with a TAT of 6.59 days for all specimens received over 14 weeks gestational age. The number of EA specimens received has increased from 1.5 per month in 1986 to 57 per month in 1991. In summary, our experience with EA specimens for cytogenetic analysis has demonstrated that the success rate is 98.6 per cent and that an increasing number of obstetricians are performing early amniocentesis as they seek to provide their patients with earlier results and an alternative to chorionic villus sampling.     

New remains of the enigmatic cetartiodactyl Bugtitherium grandincisivum Pilgrim, 1908, from the upper Oligocene of the Bugti Hills (Balochistan, Pakistan)

Newly discovered fossil material of the enigmatic cetartiodactyl Bugtitherium grandincisivum from the upper Oligocene of the Bugti Member of the Chitarwata Formation in the Bugti Hills (Balochistan, Pakistan) is reported. These new specimens consist of two fragmentary muzzles (one preserving the first incisors and belonging to a juvenile) and a fragmentary right mandible with m3. The morphologies of the anterior dentition and m3 provided by these new specimens confirm the validity of the genus Bugtitherium and advocate probable anthracotheriid affinity for the genus rather than entelodontid or suoid affinities, but do not definitively close the debate about Bugtitherium’s familial affinities within Cetartiodactyla. Although still poorly documented, this large-sized anthracotheriid-like cetartiodactyl is a possible key form for understanding the early evolution of hippos, and, in turn, the ancestry of whales, because of both its morphological similarities with hippos and primitive Paleogene whales and its Tethysian distribution.     

The European land leech: biology and DNA-based taxonomy of a rare species that is threatened by climate warming

The European land leech Xerobdella lecomtei was discovered in 1868 and is one of the rarest animals on Earth. During the 1960s, several individuals of these approx. 40 mm long, cold-adapted terrestrial annelids that inhabit the moist soils of birch forests around Graz, Austria, were investigated. Only one original research paper has been published on the biology of this species. Between 2001 and 2005, we re-investigated the morphology of preserved specimens and searched for living individuals in their natural habitat that appeared to be intact. We found only one juvenile individual (length approx. 10 mm), indicating that this local leech population became largely extinct over the past four decades. The feeding behaviour of our ‘lonesome George of the annelids’ was studied and is described here in detail. After its death, the Xerobdella individual was used for chemical extraction and molecular studies (deoxyribonucleic acid [DNA] barcoding, based on one gene, the mitochondrial cytochrome c oxidase subunit I). In addition, novel DNA barcodes for a land leech from Madagascar and a recently discovered species from Europe were obtained. Our phylogenetic tree shows that X. lecomtei is not a member of the tropical land leeches (family Haemadipsidae), as previously thought, but represents a separate line of descent (family Xerobdellidae). The decline of the local leech population around Graz correlates with a rise in average summer temperatures of +3°C between 1961 and 2004. This warming led to a drastic reduction in the moisture content of the soil where X. lecomtei lives. We suggest that human-induced climate change without apparent habitat destruction can lead to the extinction of populations of cold-adapted species that have a low colonization ability.     

Reconstructing the anatomy of the 42-million-year-old fossil †Mengea tertiaria (Insecta, Strepsiptera)

Fossilization in amber is unique in preserving specimens with microscopic fidelity; however, arthropod inclusions are rarely examined beyond the exoskeleton as this requires destructive sampling when traditional techniques are used. We report the first complete, digital 3D, non-destructive reconstruction of the anatomy of an insect fossil, a specimen of †Mengea tertiaria embedded in a 42-Ma Baltic amber. This was made possible using Synchrotron μ-CT. The species belongs to the stem group of the phylogenetically enigmatic and extremely specialized Strepsiptera. Most internal structures of the fossil are preserved, but small parts of the lumen had decayed due to incomplete infiltration of the resin. Data on internal organs provided additional information for resolving phylogenetic relationships. A sister group relationship between †Mengea and all extant lineages of the group was confirmed with characters previously not accessible. The newly gained information also yielded some insights in the biology of †Mengea and the early evolutionary history of Strepsiptera. The technique has a tremendous potential for a more accurate interpretation of diverse fossil arthropods preserved in ambers from 130 Ma to the present.     

Does leaf flushing in the dry season affect leaf traits and herbivory in a tropical dry forest?

We report a small hawk-like diurnal bird from the early Oligocene (30–31 million years ago) of Poland. Aviraptor longicrus, n. gen. et sp. is of a size comparable with the smallest extant Accipitridae. The new species is characterized by very long legs, which, together with the small size, suggest an avivorous (bird-eating) feeding behavior. Overall, the new species resembles extant sparrowhawks (Accipiter spp.) in the length proportions of the major limb bones, even though some features indicate that it convergently acquired an Accipiter-like morphology. Most specialized avivores amongst extant accipitrids belong to the taxon Accipiter and predominantly predate small forest passerines; the smallest Accipiter species also hunts hummingbirds. Occurrence of a possibly avivorous raptor in the early Oligocene of Europe is particularly notable because A. longicrus coexisted with the earliest Northern Hemispheric passerines and modern-type hummingbirds. We therefore hypothesize that the diversification of these birds towards the early Oligocene may have triggered the evolution of small-sized avivorous raptors, and the new fossil may exemplify one of the earliest examples of avian predator/prey coevolution.

     

Smaller Brains and Optic Lobes in Reproductive Workers of the Ant Harpegnathos

 Most animals show long-term modifications of their behavior which often reflect an adaptation to seasonal variations (e.g., hibernation) or result from changes in the animal's internal state (e.g., estrous cycle or sexual maturity). Such modifications may substantially affect the nervous system [1, 2]. A particularly striking behavioral change can occur in workers of the ant Harpegnathos. A few young workers in the colony may become reproductives and are thus confined to their dark nest chambers, whereas most workers spend their lives as foragers, employing acute vision when hunting prey. This behavioral difference coincides with a marked decrease in brain volume and with an even stronger reduction in the large visual brain centers. Instead of maintaining superfluous brain functions, these ants reduce brain matter which is expensive to support. Received: 8 September 1998 / Accepted: 16 March 1999     

Direct evidence of hybodont shark predation on Late Jurassic ammonites

Sharks are known to have been ammonoid predators, as indicated by analysis of bite marks or coprolite contents. However, body fossil associations attesting to this predator–prey relationship have never been described so far. Here, I report a unique finding from the Late Jurassic of western France: a complete specimen of the Kimmeridgian ammonite Orthaspidoceras bearing one tooth of the hybodont shark Planohybodus. Some possible tooth puncture marks are also observed. This is the first direct evidence of such a trophic link between these two major Mesozoic groups, allowing an accurate identification of both organisms. Although Planohybodus displays a tearing-type dentition generally assumed to have been especially adapted for large unshelled prey, our discovery clearly shows that this shark was also able to attack robust ammonites such as aspidoceratids. The direct evidence presented here provides new insights into the Mesozoic marine ecosystem food webs.     

The earliest modern mongoose (Carnivora, Herpestidae) from Africa (late Miocene of Chad)

We report on the earliest modern mongooses of Africa, from the late Miocene (ca. 7 Ma) of the hominid locality TM 266, Toros-Menalla, Chad. The material is based on fragmentary dentitions of three individuals. The main diagnostic feature of the Chadian species is the great development of the shear in the carnassials, which distinguishes the Chadian specimens from all extant herpestids except Herpestes and Galerella. In comparison with most extinct and extant Herpestes, the species from Toros-Menalla differs by a markedly smaller size and, depending on the species, relatively more elongated carnassials, more transversely elongated M1 and more reduced p4. On the basis of a great morphological similarity and the absence of significant differences, we assign our material to Galerella sanguinea; the Chadian finding therefore represents the earliest appearance of an extant species of Herpestidae. This record ties the first appearance of the genus to a minimum age of ca. 7 Ma, which is consistent with the estimated divergence date of 11.4 Ma known from the literature for the species of Galerella.     

Winged queens replaced by reproductives smaller than workers in <Emphasis Type="Italic">Mystrium</Emphasis> ants

In ants, winged queens that are specialized for independent colony foundation can be replaced by wingless reproductives better adapted for colony fission. We studied this shift in reproductive strategy by comparing two Mystrium species from Madagascar using morphometry, allometry and dissections. Mystrium rogeri has a single dealate queen in each colony with a larger thorax than workers and similar mandibles that allow these queens to hunt during non-claustral foundation. In contrast, Mystrium ‘red’ lacks winged queens and half of the female adults belong to a wingless ‘intermorph’ caste smaller and allometrically distinct from the workers. Intermorphs have functional ovaries and spermatheca while those of workers are degenerate. Intermorphs care for brood and a few mate and reproduce making them an all-purpose caste that takes charge of both work and reproduction. However, their mandibles are reduced and inappropriate for hunting centipedes, unlike the workers’ mandibles. This together with their small thorax disallow them to perform independent colony foundation, and colonies reproduce by fission. M. rogeri workers have mandibles polymorphic in size and shape, which allow for all tasks from brood care to hunting. In M. ‘red’, colonial investment in reproduction has shifted from producing expensive winged queens to more numerous helpers. M. ‘red’ intermorphs are the first case of reproductives smaller than workers in ants and illustrate their potential to diversify their caste system for better colonial economy.     

Attraction of two lacewing species to volatiles produced by host plants and aphid prey

It is well documented that host-related odors enable many species of parasitoids and predatory insects to locate their prey and prey habitats. This study reports the first characterization of prey and prey host odor reception in two species of lacewings, Chrysoperla carnea (Say) and Chrysopa oculata L. 2-Phenylethanol, one of the volatiles emitted from their preys host plants (alfalfa and corn) evoked a significant EAG response from antennae of C. carnea. Traps baited with this compound attracted high numbers of adult C. carnea, which were predominantly females. One of the sex pheromone components (1R,4aS,7S,7aR)-nepetalactol of an aphid species, Acyrthosiphon pisum (Harris) attracted only C. oculata adults. Single sensillum recordings showed that the olfactory neurons of C. carnea responded to both 2-phenylethanol and aphid sex pheromone components, but those of C. oculata only responded to the latter.     

Dental microwear patterns of extant and extinct Muridae (Rodentia, Mammalia): ecological implications

Extant species of Muridae occupy a wide array of habitats and have diverse dietary habits. Consequently, their dental microwear patterns represent a potential clue to better understand the paleoecology of their extinct relatives, which are abundant in many Old World Neogene localities. In this study, dental microwear is investigated for specimens of 17 extant species of murine and deomyine rodents in order to test the reliability of this method and infer dietary preferences on the fossil species Saïdomys afarensis. This extinct form comes from a mid-Pliocene site (AL 327) located at the Hadar Formation (Ethiopia) known to have delivered many hominid specimens of Australopithecus afarensis. A significant correlation between microwear patterns and diet is detected. Thus, grass, fruit, and insect eaters display, respectively, high amounts of fine scratches, wide scratches, and large pits. Moreover, some aspects of the paleoecology of S. afarensis, including feeding habits, could be assessed in regard to its dental microwear pattern. Indeed, it probably had feeding habits similar to that of living grass eaters. These results concur with the presence of open to woodland areas covered by an herbaceous vegetal layer, including monocotyledons, in the vicinity of this mid-Pliocene locality.     

Artiodactyls from the Pondaung Formation (Myanmar): new data and reevaluation of the South Asian Faunal Province during the Middle Eocene

Although Asia is thought to have played a critical role in the radiation of artiodactyls, the fossil record of stem selenodonts (“dichobunoids”) remains dramatically poor in tropical Asian regions. In this study, we report a new dichobunid genus and species Cadutherium kyaukmagyii and a new basal ruminant genus and species Irrawadymeryx pondaungi, from the late Middle Eocene Pondaung Formation, Central Myanmar. Although the scarcity of the present material prevents any attempts to investigate the phylogenetic relationships of Cadutherium with contemporaneous forms from other Holarctic landmasses, this new form shed new light on the diversity of these small rabbit-like ungulates during a key period of their evolutionary history. Reexamination of the small-bodied artiodactyls from Pondaung leads us to propose new identifications of certain published specimens and, in turn, to investigate the temporal and geographic distribution of taxa recognized in the Pondaung Formation. Although fragmentary, these potential new taxa reveal an unsuspected diversity of small forms among artiodactyls of Pondaung. This addition to the Eocene record of dichobunoids and early ruminants provides further insight in the diversity of dental patterns among small artiodactyls from the Pondaung Formation and attests to the antiquity of these groups in Southeast Asia.     

Prey-tracking behavior in the invasive terrestrial planarian Platydemus manokwari (Platyhelminthes, Tricladida)

Platydemus manokwari is a broadly distributed invasive terrestrial flatworm that preys heavily on land snails and has been credited with the demise of numerous threatened island faunas. We examined whether P. manokwari tracks the mucus trails of land snail prey, investigated its ability to determine trail direction, and evaluated prey preference among various land snail species. A plastic treatment plate with the mucus trail of a single species and a control plate without the trail were placed side by side at the exit of cages housing P. manokwari. All trials were then videotaped overnight. The flatworms moved along plates with mucus trails, but did not respond to plates without trails, blank control (distilled water), or with conspecific flatworm trails. When presented at the midpoint of a snail mucus trail, the flatworms followed the trail in a random direction. The flatworms showed a preference when choosing between two plates, each with a mucus trail of different land snail species. Our results suggest that P. manokwari follows snail mucus trails based on chemical cues to increase the chance of encountering prey; however, trail-tracking behavior showed no directionality.     

Regional,seasonal and interspecific variation in <Superscript>15</Superscript>N and <Superscript>13</Superscript>C in sympatric mouse lemurs

Madagascar provides some of the rare examples where two or more primate species of the same genus and with seemingly identical niche requirements occur in sympatry. If congeneric primate species co-occur in other parts of the world, they differ in size in a way that is consistent with Hutchinson’s rule for coexisting species, or they occupy different ecological niches. In some areas of Madagascar, mouse lemurs do not follow these “rules” and thus seem to violate one of the principles of community ecology. In order to understand the mechanisms that allow coexistence of sympatric congeneric species without obvious niche differentiation, we studied food composition of two identical sized omnivorous mouse lemur species, Microcebus griseorufus and M. murinus with the help of stable isotope analyses (δ ¹⁵N and δ ¹³C). The two species are closely related sister species. During the rich season, when food seems abundant, the two species do not differ in their nitrogen isotope composition, indicating that the two species occupy the same trophic level. But they differ in their δ ¹³C values, indicating that M. griseorufus feeds more on C₄ and CAM (Crassulacean-acid-metabolism) plants than M. murinus. During the lean season, M. murinus has lower δ ¹⁵N values, indicating that the two species feed at different trophic levels during times of food shortage. Hybrids between the two species showed intermediate food composition. The results reflect subtle differences in foraging or metabolic adaptations that are difficult to quantify by traditional observations but that represent possibilities to allow coexistence of species.     

Non-specific association between filamentous bacteria and fungus-growing ants

Fungus-growing ants and their fungal cultivar form a highly evolved mutualism that is negatively affected by the specialized parasitic fungus Escovopsis. Filamentous Pseudonocardia bacteria occurring on the cuticle of attine ants have been proposed to form a mutualistic interaction with these ants in which they are vertically transmitted (i.e. from parent to offspring colonies). Given a strictly vertical transmission of Pseudonocardia, the evolutionary theory predicts a reduced genetic variability of symbionts among ant lineages. The aim of this study was to verify whether actinomycetes, which occur on Acromyrmex octospinosus leaf-cutting ants, meet this expectation by comparing their genotypic variability with restriction fragment length polymorphisms. Multiple actinomycete strains could be isolated from both individual ant workers and colonies (one to seven strains per colony). The colony specificity of actinomycete communities was high: Only 15% of all strains were isolated from more than one colony, and just 5% were present in both populations investigated. Partial sequencing of 16S ribosomal deoxyribonucleic acid of two of the isolated strains assigned both of them to the genus Streptomyces. Actinomycetes could also be isolated from workers of the two non-attine ant species Myrmica rugulosa and Lasius flavus. Sixty-two percent of the strains derived from attine ants and 80% of the strains isolated from non-attine ants inhibited the growth of Escovopsis. Our data suggest that the association between attine ants and their actinomycete symbionts is less specific then previously thought. Soil-dwelling actinomycetes may have been dynamically recruited from the environment (horizontal transmission), probably reflecting an adaptation to a diverse community of microbial pathogens.