Search

Successful invaders often become established in new ranges by outcompeting native species. The "evolution of increased competitive ability" hypothesis predicts that invasive species are subjected to less predation and parasitization than sympatric native species, and thus can allocate resources from defence and immunity to growth and fecundity, thereby achieving higher fitness. In this study, we examined whether American invasive Polistes dominula paper wasps have reduced immunocompetence. To explore this scenario, we tested their susceptibility towards parasites and pathogens at both the individual (immune defence) and colony levels, i.e. hygienic behaviour (removal of diseased individuals by nestmates). First, we examined the response to the specific coevolved parasite Xenos vesparum (lost after invasion) in terms of individual host susceptibility and hygienic behaviour. Second, we explored the response against general pathogens by quantifying the bacterial clearance in individual wasps after a challenge with Escherichia coli and hygienic behaviour after a challenge with the fungus Beauveria bassiana. Our results show that American invasive P. dominula have a higher response against X. vesparum at the colony level, but at the individual level their susceptibility is not significantly different from conspecifics of the native range. On the other hand, invasive P. dominula display lower response after a challenge with general pathogens at both the individual and colony levels. While supporting the hypothesis of a reduction of immunocompetence towards general pathogens in invasive species, these findings also suggest that the response against coevolved parasites might follow different evolutionary pathways which are not always easily predictable.

Social wasps from temperate zones have clear annual colony cycles, and the young queens hibernate during winter. In the subtropics, the only previously reported evidence for the existence of "hibernation" is the facultative winter aggregations of females during harsh climate conditions. As in temperate-zone species analyzed so far, we show in this study that in the paper wasp, Polistes versicolor, a subtropical species, body size increases as an unfavorable season approaches. Our morphological studies indicate that larger females come from winter aggregations--that is, they are new queens. Multivariate analyses indicate that size is the only variable analyzed that shows a relationship to the differences. Given the absence of a harsh climate, we suggest that the occurrence of winter aggregations in tropical P. versicolor functions to allow some females to wait for better environmental conditions to start a new nest, rather than all being obliged to start a new nest as soon as they emerge.

In social insects, biogenic amines are thought to play regulatory roles in the transition between reproductive states in females. To determine the effect of dopamine on the reproductive development of workers in primitively eusocial societies, isolated workers of the paper wasp Polistes chinensis were supplied with oral dopamine. Ovarian development was accelerated in dopamine-fed workers as compared to control workers of the same age fed only sucrose solution. Oral dopamine increased brain levels of dopamine and its metabolite (N-acetyldopamine). Brain levels of tyramine or octopamine were also increased by dopamine application in one of two colonies; levels of the tyramine metabolite N-acetyltyramine were unchanged. These results indicate that dopamine plays a gonadotropic role in isolated workers in the primitively eusocial wasp, similar to the gonadotropic role previously reported for juvenile hormone. This is the first study to report effects of dopamine on ovarian development in workers of the paper wasp.

Phosphatic (possibly secondarily phosphatised) remains of antipatharian coralla, previously unknown in the fossil record, occur abundantly in the early Ordovician Fenxiang Formation in the Hubei Province, southern China. Probably two species (and genera) are represented, which differ in spinosity of branches. The more spinose one, Sinopathes reptans, has its lateral spines bearing regular, longitudinally arranged costellae. The early Floian geological age of this finding, about 470 Ma, supports predictions on the timing of anthozoan phylogeny derived from the molecular phylogenetic evidence. Black corals (Antipatharia) are basal to the scleractinians in the Hexacorallia clade, being more derived than sea anemones and the Zoantharia. Based on calibration of the molecular clock with Mesozoic data, the first split of lineages within the scleractinian hexacorals was proposed to take place approximately 425 million years ago. This implies that the origin of Antipatharia should precede this date. They have not been known in the fossil record because of unmineralised skeleton composed primarily of laminar chitin complexed with a protein. Unlike all recent species, the encrusting basal part of the colony dominated in the Ordovician ones and only occasionally erect branches developed, rather chaotically ramified. This presumably plesiomorphic trait seems consistent with ancient geological age and suggests that some problematic fossils from the Late Cambrian may be their, even less-derived, relatives.

Global pollinator declines have been attributed to habitat destruction, pesticide use, and climate change or some combination of these factors, and managed honey bees, Apis mellifera, are part of worldwide pollinator declines. Here we exposed honey bee colonies during three brood generations to sub-lethal doses of a widely used pesticide, imidacloprid, and then subsequently challenged newly emerged bees with the gut parasite, Nosema spp. The pesticide dosages used were below levels demonstrated to cause effects on longevity or foraging in adult honey bees. Nosema infections increased significantly in the bees from pesticide-treated hives when compared to bees from control hives demonstrating an indirect effect of pesticides on pathogen growth in honey bees. We clearly demonstrate an increase in pathogen growth within individual bees reared in colonies exposed to one of the most widely used pesticides worldwide, imidacloprid, at below levels considered harmful to bees. The finding that individual bees with undetectable levels of the target pesticide, after being reared in a sub-lethal pesticide environment within the colony, had higher Nosema is significant. Interactions between pesticides and pathogens could be a major contributor to increased mortality of honey bee colonies, including colony collapse disorder, and other pollinator declines worldwide.

Cichlids are an excellent model to study explosive speciation and adaptive radiation. Their evolutionary success has been attributed to their ability to undergo rapid morphological changes related to diet, and their particular breeding biology. Relatively minor changes in morphology allow for exploitation of novel food resources. The importance of phenotypic plasticity and genetically based differences for diversification was long recognized, but their relationship and relative magnitude remained unclear. We compared morphology of individuals of four wild populations of the Lake Tanganyika cichlid Tropheus moorii with their pond-raised F1 offspring. The magnitude of morphological change via phenotypic plasticity between wild and pond-bred F1 fish exceeds pairwise population differences by a factor of 2.4 (mean Mahalanobis distances). The genetic and environmental effects responsible for among population differentiation in the wild could still be recognized in the pond-bred F1 fish. All four pond populations showed the same trends in morphological change, mainly in mouth orientation, size and orientation of fins, and thickness of the caudal peduncle. As between population differentiation was lower in the wild than differentiation between pond-raised versus wild fish, we suggest the narrow ecological niche and intense interspecific competition in rock habitats is responsible for consistent shape similarity, even among long-term isolated populations. Electronic supplementary material The online version of this article (doi:10.1007/s00114-010-0751-2) contains supplementary material, which is available to authorized users.

Whereas some bird species are heavily affected by blood parasites in the wild, others reportedly are not. Seabirds, in particular, are often free from blood parasites, even in the presence of potential vectors. By means of polymerase chain reaction, we amplified a DNA fragment from the cytochrome b gene to detect parasites of the genera Plasmodium, Leucocytozoon, and Haemoproteus in 14 seabird species, ranging from Antarctica to the tropical Indian Ocean. We did not detect parasites in 11 of these species, including one Antarctic, four subantarctic, two temperate, and four tropical species. On the other hand, two subantarctic species, thin-billed prions Pachyptila belcheri and dolphin gulls Larus scoresbii, were found infected. One of 28 thin-billed prions had a Plasmodium infection whose DNA sequence was identical to lineage P22 of Plasmodium relictum, and one of 20 dolphin gulls was infected with a Haemoproteus lineage which appears phylogenetically clustered with parasites species isolated from passeriform birds such as Haemoproteus lanii, Haemoproteus magnus, Haemoproteus fringillae, Haemoproteus sylvae, Haemoproteus payevskyi, and Haemoproteus belopolskyi. In addition, we found a high parasite prevalence in a single tropical species, the Christmas Island frigatebird Fregata andrewsi, where 56% of sampled adults were infected with Haemoproteus. The latter formed a monophyletic group that includes a Haemoproteus line from Eastern Asian black-tailed gulls Larus crassirostris. Our results are in agreement with those showing that (a) seabirds are poor in hemosporidians and (b) latitude could be a determining factor to predict the presence of hemosporidians in birds. However, further studies should explore the relative importance of extrinsic and intrinsic factors on parasite prevalence, in particular using phylogenetically controlled comparative analyses, systematic sampling and screening of vectors, and within-species comparisons.

Giant honeybees (Apis dorsata) nest in the open and have therefore evolved a variety of defence strategies. Against predatory wasps, they produce highly coordinated Mexican wavelike cascades termed ‘shimmering’, whereby hundreds of bees flip their abdomens upwards. Although it is well known that shimmering commences at distinct spots on the nest surface, it is still unclear how shimmering is generated. In this study, colonies were exposed to living tethered wasps that were moved in front of the experimental nest. Temporal and spatial patterns of shimmering were investigated in and after the presence of the wasp. The numbers and locations of bees that participated in the shimmering were assessed, and those bees that triggered the waves were identified. The findings reveal that the position of identified trigger cohorts did not reflect the experimental path of the tethered wasp. Instead, the trigger centres were primarily arranged in the close periphery of the mouth zone of the nest, around those parts where the main locomotory activity occurs. This favours the ‘special-agents’ hypothesis that suggest that groups of specialized bees initiate the shimmering. Electronic supplementary material The online version of this article (doi:10.1007/s00114-009-0605-y) contains supplementary material, which is available to authorized users.

The evolution of winged stick insects (phasmids) from secondarily wingless ancestors was proposed in recent studies. We explored the cuticle of flying phasmids for wind sensors that could be involved in their flight control, comparable to those known for locusts. Surprisingly, wind-sensitive hairs (wsH) occur on the palps of mouthparts and on the antennae of the winged phasmid Sipyloidea sipylus which can fly in tethered position only when air currents blow over the mouthparts. The present study describes the morphology and major functional properties of these “new” wsH with soft and bulging hair bases which are different from the beaker-like hair bases of the wsH on the cerci of phasmids and the wsH described in other insects. The most sensitive wsH of antennae and palps respond with phasic-tonic afferents to air currents exceeding 0.2 ms−1. The fields of wsH on one side of the animal respond mainly to ventral, lateral, and frontal wind on the ipsilateral side of the head. Afferent inputs from the wsH converge but also diverge to a group of specific interneurons at their branches in the suboesophageal ganglion and can send their integrated input from wsH fields of the palps and antennae to the thoracic central nervous system. Response types of individual wsH-interneurons are either phasic or phasic-tonic to air puffs or constant air currents and also, the receptive fields of individual interneurons differ. We conclude that the “new” wsH system and its interneurons mainly serve to maintain flight activity in airborne phasmids and also, the “new” wsH must have emerged together with the integrating interneurons during the evolution from wingless to the recent winged forms of phasmids. peerReviewed

In all higher nonhuman primates, species survival depends upon safe carrying of infants clinging to body hair of adults. In this work, measurements of mechanical properties of ape hair (gibbon, orangutan, and gorilla) are presented, focusing on constraints for safe infant carrying. Results of hair tensile properties are shown to be species-dependent. Analysis of the mechanics of the mounting position, typical of heavier infant carrying among African apes, shows that both clinging and friction are necessary to carry heavy infants. As a consequence, a required relationship between infant weight, hair-hair friction coefficient, and body angle exists. The hair-hair friction coefficient is measured using natural ape skin samples, and dependence on load and humidity is analyzed. Numerical evaluation of the equilibrium constraint is in agreement with the knuckle-walking quadruped position of African apes. Bipedality is clearly incompatible with the usual clinging and mounting pattern of infant carrying, requiring a revision of models of hominization in relation to the divergence between apes and hominins. These results suggest that safe carrying of heavy infants justify the emergence of biped form of locomotion. Ways to test this possibility are foreseen here.

Associative learning is key to how bees recognize and return to rewarding floral resources. It thus plays a major role in pollinator floral constancy and plant gene flow. Honeybees are the primary model for pollinator associative learning, but bumblebees play an important ecological role in a wider range of habitats, and their associative learning abilities are less well understood. We assayed learning with the proboscis extension reflex (PER), using a novel method for restraining bees (capsules) designed to improve bumblebee learning. We present the first results demonstrating that bumblebees exhibit the memory spacing effect. They improve their associative learning of odor and nectar reward by exhibiting increased memory acquisition, a component of long-term memory formation, when the time interval between rewarding trials is increased. Bombus impatiens forager memory acquisition (average discrimination index values) improved by 129% and 65% at inter-trial intervals (ITI) of 5 and 3 min, respectively, as compared to an ITI of 1 min. Memory acquisition rate also increased with increasing ITI. Encapsulation significantly increases olfactory memory acquisition. Ten times more foragers exhibited at least one PER response during training in capsules as compared to traditional PER harnesses. Thus, a novel conditioning assay, encapsulation, enabled us to improve bumblebee-learning acquisition and demonstrate that spaced learning results in better memory consolidation. Such spaced learning likely plays a role in forming long-term memories of rewarding floral resources. © 2009 Springer-Verlag.

Environmental stressors during early life may have persistent consequences for phenotypic development and fitness. In group-living species, an important stressor during juvenile development is the presence and familiarity status of conspecific individuals. To alleviate intraspecific conflicts during juvenile development, many animals evolved the ability to discriminate familiar and unfamiliar individuals based on prior association and use this ability to preferentially associate with familiar individuals. Assuming that familiar neighbours require less attention than unfamiliar ones, as predicted by limited attention theory, assorting with familiar individuals should increase the efficiency in other tasks. We assessed the influence of social familiarity on within-group association behaviour, development and foraging of juvenile life stages of the group-living, plant-inhabiting predatory mite Phytoseiulus persimilis. The observed groups consisted either of mixed-age familiar and unfamiliar juvenile mites or of age-synchronized familiar or unfamiliar juvenile mites or of pairs of familiar or unfamiliar larvae. Overall, familiar mites preferentially grouped together and foraged more efficiently, i.e. needed less prey at similar developmental speed and body size at maturity, than unfamiliar mites. Preferential association of familiar mites was also apparent in the inter-exuviae distances. Social familiarity was established by imprinting in the larval stage, was not cancelled or overridden by later conspecific contacts and persisted into adulthood. Life stage had an effect on grouping with larvae being closer together than nymphal stages. Ultimately, optimized foraging during the developmental phase may relax within-group competition, enhance current and future food supply needed for optimal development and optimize patch exploitation and leaving under limited food.

Chemically mediated communication is common in spiders but has been poorly studied in burrowing tarantulas. This study aimed to determine whether chemical cues influence the behaviour of females of Brachypelma vagans, a Mexican species of tarantula, during encounters with previously inhabited burrows or with extracts from the silk of conspecific females. In laboratory choice tests, female tarantulas entered a burrow that had previously been inhabited by a conspecific female significantly more frequently than a burrow that had never been inhabited. The identity of the previous inhabitant also affected the number of spiders that chose to enter a burrow. Spiders were quicker to choose and enter a burrow previously inhabited by themselves than a burrow previously inhabited by a conspecific or a burrow that had not been previously inhabited. Hexane, methanol and dichloromethane extracts of conspecific silk elicited different responses from female tarantulas when extracts were placed on filter paper disks at one end of an experimental arena with a control filter paper disk, on to which the corresponding solvent alone had been pipetted, placed on the other end of the arena. Spiders showed the strongest responses to hexane extracts of silk, with a significant preference to move towards the hexane extract and a significantly greater period of time spent in proximity to the hexane extract compared to the control disk. Overall and in contrast to expectations, tarantulas were most strongly attracted to the cues left by other conspecific females. As encounters between B. vagans females usually lead to aggression and mortality of one of the participants, we conclude that chemical cues are not signals that are deliberately released by burrow-inhabiting females but may inadvertently escape and cannot be easily suppressed.

Strong and frequent rain is essentialfor the existence of tropical rain for-ests. Heavy showers, however, pose athreat especially for arboreal insectswhich can be washed down fromtrees or even be drowned in theirshelters. Queens of the plant-antgenus Cladomyrma, which must bitenest chambers into their host plantstems for colony foundation, fre-quently disappear during heavy rainand are sometimes drowned whenseeking refuge in their unfinished,still unclosed chambers due to runoffwater which collects in the hollow in-ternodes via the entrance holes. Ma-ture colonies of these ant species facethe same problem of intruding rain-water endangering ant brood and tro-phobiotic mealybugs, but they re-spond with an unusual behaviour onlyonce previously reported for ants: theworkers ingest the liquid and regurgi-tate it to the outside until no standingwater remains. This behaviour can bereadily provoked by injecting smallamounts of liquid into inhabited inter-nodes. The ability of Cladomyrma toremove water from its nest site ap-pears to be an adaptation to their obli-gate colonization of myrmecophytes(ant-plants).Cladomyrma is a formicine genuswhose species live exclusively in avariety of trees and vines from Bor-neo and the Malay Peninsula. To ourpresent knowledge Cladomyrma inha-bits 18 host plant species from 9 gen-era and 8families ([1–4], unpublishedresults). These ants protect their hostplants against herbivores ([4, 5], un-published results) and prune (at leastin part) encroaching vegetation. Theants gnaw open the host plant inter-nodes and hollow them out by remov-ing the pith. With both increasingplant and colony size other internodesare colonized successively. Eventuallythe nest chambers run through stem,branches and every twig of the host.These hollows contain brood and tro-phobiotic pseudococcids which arecultivated by the ants. As a rule onenest entrance per internode is found.In regard to nest structure the hostNeonauclea Merr. (Rubiaceae) differsfrom other host plants of Cladomyr-ma. Here only internodes which areswollen ( = myrmecodomatia) are hol-lowed out and colonized by the ants.The number of entrance holes per do-matium varies greatly, on averageabout 4–6 are found, depending onboth domatium and colony size.In all host plants the entrances are si-tuated more or less at the lateral sidesof the internodes. Thus the risk ofgetting runoff-water into the nestchambers is reduced (exept for thevertical stem). However, rainwatermay still be able to collect in the nest.We dissected longitudinally several in-habited internodes of the trees Sara-ca, Ryparosa, Neonauclea, Cryptero-nia, and the woody climber Spatholo-bus. By watering the outer walls ofthe internodes we could check frominside that small amounts of water dointrude into the nest. Most of the run-off water, however, passes the en-trances due to very small callus ridgesaround the holes. Rain water mayalso penetrate into the nest hollowsthrough openings of damaged twigs,the latter being torn open by predatorsof ants and brood.To test how Cladomyrma ants react tothe intrusion of water we injected be-tween 0.2 and 6 ml artificially co-loured liquid into nest entrances witha syringe. Table 1 summarizes the re-sults. In total we performed 16 testswith nine colonies of three Cladomyr-ma species, C. near andrei (Emery; 1colony, 3 tests), C. maschwitzi Agosti(4 colonies, 6 tests), C. petalae Agosti(4 colonies, 7 tests); and in nine hostplants of five various other species:Saraca thaipingensis Cantley ex Prain(n = 2), Ryparosa fasciculata King(n = 1), Spatholobus bracteolatusPrain (n = 1), Crypteronia griffithiiClarke (n = 4), and Neonauclea gigan-tea (Valeton) Merr. (n = 1). On eachplant we fixed the experimental twigsin a horizontal position to prevent theinjected water from draining away in-side the nest channels. Prior to the in-jection of coloured liquid we fixed asmall filter paper (4 ×4 cm) beneatheach experimental entrance hole(n = 16). Within minutes after flood-ing the internodes (1–15 min, median4, mean 5.38 ± 4.76) the number ofworkers increased noticeably (at leastthree-fold the activity level prior to in-jection) around the affected areas.Shortly afterwards the first workerswith distended gasters, containing co-loured liquid (tested by squashingworkers on filter paper), appeared onthe plant surface (3–65 min, median5, mean 12.08 ± 17.52). There alwayspassed a considerable period of timebefore the first appearance of thewater-ejection behaviour (3–41 min,median 27, mean 24.92 ± 13.18).The results are highly variable andmay be due to (a) different amountused of injected liquid, (b) the num-ber of workers involved, i.e., colonysize, (c) incomparability of risk levels,i.e., different number of larvae orpseudococcids inside the affected in-ternodes, and (d) species-specific dif-ferences in behavioural response.

Tandem running is a recruitment strategy whereby one ant leads a single naïve nest mate to a resource. While tandem running progresses towards the goal, the leader ant and the follower ant maintain contact mainly by tactile signals. In this paper, we investigated whether they also deposit chemical signals on the ground during tandem running. We filmed tandem-running ants and analysed the position of the gasters of leaders and followers. Our results show that leader ants are more likely to press their gasters down to the substrate compared to follower ants, single ants and transporter ants. Forward tandem-run leaders (those moving towards a new nest site) performed such trail-marking procedures three times more often than reverse tandem leaders (those moving towards an old nest site). That leader ants marked the trails more often during forward tandem runs may suggest that it is more important to maintain the bond with the follower ant on forward tandem runs than on reverse tandem runs. Marked trails on the ground may serve as a safety line that improves both the efficiency of tandem runs and their completion rates.

Monogeny, the production of unisexual broods by individual females, has been recognized for nearly 80 years. The genetic nature of gall midges' sex determination predicts an equal numbers of male-producing and female-producing females in the populations such that the overall sex ratio is expected to be nearly 1:1. However, observations of some strictly monogenous populations with biased sex ratio, mainly toward females, have raised the question of whether gall midges are able to adjust their offspring sex ratio in response to changes in environmental conditions, and some authors have even considered sex ratio regulation as a strong force in the course of the evolution of monogeny. In this paper, first, by studying the sex ratio variations of the predatory gall midge, Aphidoletes aphidimyza within a generation, we showed that adult males emerge up to 1 day earlier and have shorter life span than females (less than 4 days and up to 6 days, respectively). Although, the sex ratio of A. aphidimyza at the time of emergence was nearly 1:1 (52.41 % males), a simple population simulation indicated that the differential mortality of sexes can lead to a female-biased sex ratio estimation (57.88 % females) under random sampling in the natural environments. Our results imply that the primary sex ratio of monogenous gall midges is nearly 1:1 and that the arrhenogenic/thelygenic gall midges are not able to alter the number of their male/female progenies in response to changes in environmental conditions.

Cacops, one of the most distinctive Paleozoic amphibians, is part of a clade of dissorophoid temnospondyls that diversified in the equatorial region of Pangea during the Late Carboniferous and Early Permian, persisting into the Late Permian in Central Russia and China. Dissorophids were a successful group of fully terrestrial, often spectacularly armoured predators, the only amphibians apparently able to coexist with amniotes when the latter started to dominate terrestrial ecosystems. In this paper, we describe excellent new skulls from the Early Permian of Oklahoma attributed to Cacops, Cacops morrisi sp. nov. and provide for the first time detailed information about this iconic dissorophid. These specimens show anatomical and ontogenetic features that will impact on future studies on the evolution of terrestriality in tetrapods. For example, the large, posteriorly closed tympanic embayment has fine striations on an otherwise smooth surface, documenting the oldest known clear evidence for the presence of a tympanic membrane in the fossil record, a structure that is used for hearing airborne sound in extant tetrapods. The skull of C. morrisi also has several features associated with predatory behaviour, indicating that this dissorophid may have been one of the top terrestrial predators of its time.

The Southern river otter or 'huillin', Lontra provocax, is an endangered species endemic of the Andean Patagonian region of Argentina and Chile. It feeds almost exclusively on the genera of macro-crustacea: Aegla and Sammastacus. The aim of this study was to analyse the role of food availability on the huillin's distribution using a scale-dependent analysis of crustacean and otter distributions. We compared the distributions of otters and macro-crustaceans along a north-south regional gradient, between river basins of northern Patagonia, in an altitudinal gradient within a river basin, and between habitat types within a lake. We investigated the distribution of otters by sign surveys along lake shores, river banks and marine coasts, and of crustaceans using surveys in the water, undigested remains in mink (Mustela vison) scats, presence of external skeletons at the waterside and through interviews with local people. Our results show that there were heterogeneities in the distributions of macro-crustaceans at four scales and these were generally reflected in the distributions of freshwater otters. We conclude that the main factor limiting the distributions of L. provocax in freshwater environments is the availability of macro-crustaceans. This paper shows how scale-dependent type analyses of population distribution serves as a method for identifying key environmental factors for species for which the use of long-term demographies is unfeasible.

In recent years, three terrestrial bugs, Adomerus triguttulus and Sehirus cinctus (Cydnidae) and the closely related Parastrachia japonensis (Parastrachiidae), have been the focus of several fascinating studies because of the remarkable, extensive parental care they were found to display. This care includes egg and nymph guarding, production of trophic eggs, unfertilized, low cost eggs that are used as food by newly hatched nymphs, and progressive provisioning of the host seed. In this study, we have investigated yet a third related Asian cydnid, Canthophorus niveimarginatus, with regard to the possible occurrence of some or all of these complex traits in order to assess how widespread these maternal investment patterns are in this group of insects and to better understand the implications of their manifestations from an evolutionary context. Manipulative experiments were carried out in the lab to determine whether females provision nests. Observational and egg removal studies were carried out to determine whether trophic eggs are produced, and, if they are, their possible impact on nymphal success. The findings revealed that C. niveimarginatus does, in fact, progressively provision young, and this species also displays all of the other behaviors associated with extended parental care in subsocial insects. Moreover, unlike the other two related species, which place trophic eggs on the surface of the original egg mass, C. niveimarginatus produces both pre- and post-hatch trophic eggs. Nymphs deprived of access to post-hatch trophic eggs had significantly lower body weight and survival rate than those that fed on them. To our knowledge, this is the first time the production of both pre- and post-hatch trophic eggs has been demonstrated in insects outside the Hymenoptera. In this paper, we qualitatively and quantitatively demonstrate the provisioning behavior and patterns of trophic egg production in C. niveimarginatus. When and how trophic eggs are produced and delivered to young should have important correlations with the ecological and life history constraints under which a species has evolved. Thus, we also discuss the possible ecological and life history factors that favor the evolution of post-hatch trophic eggs.

Priscochelys hegnabrunnensis, a fragmentary piece of armour shell from the Muschelkalk of Germany (Upper Triassic) with few diagnostic morphological features, was recently proposed to represent the oldest known stem turtle. As such, the specimen is of high importance because it shifts the date of the first appearance of turtles back about 20 Ma, which equals about 10% of the total stratigraphic range of the group. In this paper, I present new morphologic, histologic and neutron tomographic (NT) data that relate to the microstructure of the bone of the specimen itself. In opposition to the previous morphologic descriptions, P. hegnabrunnensis was found to share several distinctive features (i.e. bone sutures congruent with scute sulci, absence of a diploe structure with interior cancellous bone, thin vascular canals radiating outwards from distinct centres in each field and rugose ventral bone surface texture consisting of mineralised fibre bundles) with cyamodontoid placodonts (Diapsida: Sauropterygia) and fewer with stem turtles (i.e. depth of sulci). Two aspects that were previously thought to be relevant for the assignment to the turtle stem (conical scutes and presence of foramina) are argued to be of dubious value. P. hegnabrunnensis is proposed to represent a fragmentary piece of cyamodontoid armour consisting of fused conical plates herein. The specimen is not a part of the turtle stem and thus does not represent the oldest turtle. Accordingly, P. hegnabrunnensis does not shorten the ghost lineage to the potential sister group of turtles.

Social insects can modulate body temperature to increase foraging efficiency; however, little is known about how the relative value of protein resources affects forager body temperature. Such regulation may be important given that colony growth is often limited by protein availability. In this paper, we present what are, to our knowledge, the first data for social insects showing that thoracic temperatures (T (th)) of foragers increase with the protein content of food resources. In an introduced population of western yellowjacket (Vespula pensylvanica), we measured T (th) of foragers collecting high-quality protein (100% canned chicken) and low-quality protein (50% canned chicken, 50% indigestible alpha-cellulose by volume) at different ambient air temperatures (T (a)). Wasps foraging on 100% chicken consistently exhibited higher T (th) compared to wasps foraging on 50% chicken. After correcting for T (a), the mean T (th) for wasps collecting 100% chicken were 1.98 degrees C higher than those of individuals collecting 50% chicken. We suggest that this mechanism may increase foraging efficiency in this and other social wasp species.

Feigning death is a method of self-defense employed among a wide range of prey species when threatened by predator species. This paper reports on death-feigning behavior by the fire ant, Solenopsis invicta, during intraspecific aggression among neighboring fire ant workers. Days-old workers responded to aggression by death feigning, weeks-old workers responded by fleeing and months-old workers responded by fighting back. By feigning death, days-old workers were four times more likely to survive aggression than older workers. From a proximate perspective, retaliation by young workers against aggressive older workers is certain to fail. With their relatively soft exoskeleton, young workers would be prone to injury and death and unable to execute an effective attack of biting or stinging older workers with harder exoskeletons. From an ultimate perspective, death feigning allows young workers to survive and contribute to brood care and colony growth, both of which are essential to queen survival and fitness.

The display of female traits by males is widespread in the animal kingdom. In several species, this phenomenon has been shown to function adaptively as a male mating strategy to deceive sexual rivals (female mimicry). Freshly emerged males of the parasitic wasp Lariophagus distinguendus (Hymenoptera: Pteromalidae) are perceived by other males as if they were females because of a very similar composition of cuticular hydrocarbons which function as a sex pheromone in this species inducing courtship behaviour in males. Within 32 h, however, males deactivate the pheromone and are no longer courted by other males. In this paper, behavioural experiments were performed to test hypotheses on potential costs and benefits associated with the female odour in young males. We did not find any benefits, but demonstrated that young males were significantly more often outrivaled in male-male contests when competing with two older males for a female. Also, young males were significantly more often mounted in homosexual courtship events during these contests. Thus, display of female traits by males is not necessarily beneficial, and in fact, can be disadvantageous. We suggest that these costs have favoured the evolution of the pheromone deactivation mechanism in L. distinguendus males. The function of cuticular hydrocarbons as a female courtship pheromone in L. distinguendus might have evolved secondarily from a primary function relevant for both genders, and the deactivation of the signal in males might have caused a shift of specificity of the chemical signal from the species level to the sex level.

Predation has received considerable theoretical and empirical support in population regulation. The effect of predators, however, could be achieved in direct (killing) or indirect effects (such as displacement). In this paper, we explored the relationship between Eurasian kestrels Falco tinnunculus and skylarks Alauda arvensis in Mediterranean grasslands. First, we analysed the presence of skylarks in the kestrel diet over 9 years. We also compared a grassland area of experimentally increased kestrel density and a second grassland as control area to evaluate the direct or indirect effect on skylark abundance. We also considered two different habitats, grazed and ungrazed plots. If skylark abundance decreased as the kestrel breeding season progressed in high-density kestrel area compared with the control area, it would suggest a direct effect (predator hypothesis). If skylark abundance remains constant in both areas of contrasting kestrel density, it would suggest that skylarks avoid kestrels (avoidance hypothesis). We found that skylark abundance decreased in the kestrel area from the beginning of kestrel nest-box installation to recent years. The rate of skylark consumption decreased in a 9-year period as kestrel abundance increased, although the total amount skylark consumption did not show a decreasing trend. In addition, skylarks were more abundant in the kestrel-free area than in the kestrel area. Finally, we found that skylark abundance did not change through the kestrel breeding period in relation to grazing. We suggest that an increased breeding density of kestrels during their breeding period may force the skylarks to breed in other areas, which may explain the decline of their abundance.

Decision making is of crucial importance in the lives of both animals and humans. How decisions of group members scale up to group decisions is of great interest. Accordingly, we gave homeless ant colonies (n = 67) in three experiments a choice between two nest sites (with small, big or mixed sand grains), each of which had to be excavated to be habitable. Among the colonies that chose only one of the new nest sites, all preferred the ones that could be excavated most easily and quickly. There are interesting parallels between the collective choice of mining sites and the ability of certain ants to select short cuts; both involve positive feedback. However, in this paper, we discuss a mechanism whereby collective co-ordination in the production of social infrastructure can occur in the absence of signalling.

There has been considerable recent interest in the biology of spiders that specialise on ants as prey, but previous studies have tended to envisage the level of adaptation as being to ants as a group. In this paper, we provide evidence that Zodarion germanicum is a spider that has dietary and venom adaptations by which it targets a particular subset of ants, the subfamily Formicinae. We reared spiders from first instar in the laboratory on three different diets: formicine ants only, myrmicine ants only and mixed (both formicine and myrmicine ants). Fitness-related life-history parameters were determined, and we found that the spiders on the formicine-only diet lived longer and grew at a faster rate. Lipid, carbon and nitrogen compositions of ants were analysed, but we found no evidence of formicines differing from myrmicines in macro-nutrient content. This suggests that effects on longevity and growth depended on more specific nutrients or on compounds the prey uses for defence. We investigated how efficient Z. germanicum was at paralysing different ants and our findings suggest that the spider's venom is especially effective against formicines. Taken together, our findings suggest that Z. germanicum has evolved specialisation at the level of targeting a particular ant subfamily, the Formicinae

More than one century after its original description by Marsh in 1877, we report in this paper the first uncontroversial evidence of a member of the genus Stegosaurus out of North America. The specimen consists of a partial skeleton from the Upper Jurassic of Portugal, herein considered as Stegosaurus cf. ungulatus. The pres- ence of this plated dinosaur in the upper Kimmeridgian- lower Tithonian Portuguese record and synchronic levels of the Morrison Formation of North America reinforces previous hypothesis of a close relationship between these two areas during the Late Jurassic. This relationship is also supported by geotectonic evidences indicating high proba- bility of an episodic corridor between the Newfoundland and Iberian landmasses. Together, Portuguese Stegosaurus dis- covery and geotectonic inferences could provide a scenario with episodical faunal contact among North Atlantic land- masses during the uppermost Kimmeridgian-lowermost Tithonian (ca. 148-153 Ma ago).

Aphids often form mutualistic associations with ants, in which the aphids provide the ants with honeydew and the ants defend the aphids from predators. In this paper, we report aphid egg protection by ants as a novel aspect of the deeply interdependent relationship between a tree-feeding aphid and its attendant ant. The ant Lasius productus harbours oviparous females, males, and eggs of the hinoki cypress-feeding aphid Stomaphis hirukawai in its nests in winter. We investigated the behaviour of ants kept with aphid eggs in petri dishes to examine whether the ants recognise the aphid eggs and tend them or only provide a refuge for the aphids. Workers carried almost all of the aphid eggs into the nest within 24 h. The ants indiscriminately tended aphid eggs collected from their own colonies and those from other ant colonies. The ants cleaned the eggs and piled them up in the nest, and egg tending by ants dramatically increased aphid egg survival rates. Starving the ants showed no significant effect on aphid egg survivorship. Without ants, aphid eggs were rapidly killed by fungi. These results suggested that grooming by the ants protected the aphid eggs, at least, against pathogenic fungi. This hygienic service afforded by the ants seems indispensable for egg survival of these aphids in an environment rich in potentially pathogenic microorganisms.

The geographically widespread Python reticulatus, the world's longest snake, has been largely neglected by taxonomists. Dwarfed individuals from Tanahjampea Island, Indonesia, differ strikingly in morphology. Phylogenetic relationships were analyzed using a 345-bp fragment of the cytochrome b gene for 12 specimens from different populations. Both genetic differences and morphological characters distinctly revealed two taxonomic subunits. The island populations of Tanahjampea and Selayar form two monophyletic lineages, supported by high bootstrap values, with distinct differences in color pattern and scalation. We consider these forms to represent two new subspecies. The Tanahjampea form is genetically related to populations of the Sunda Islands and mainland Southeast Asia, whereas the Selayar form is related to populations of Southwest Sulawesi. We conclude that, due to strong directional surface currents in this region, gene flow between Tanahjampea and Selayar is prevented. Sea-level changes during the Pleistocene probably contributed to the isolation of the two taxa described. Aspects of ecology and conservation status are briefly discussed. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00114-002-0320-4.

The Arnold Berliner Award is presented for the first time this year, as part of the 100-year birthday celebration of Naturwissenschaften (Fig. 1, Thatje 2013). It is my great pleasure to announce the first recipient of the award, Dr Mark Young (University of Edinburgh), in this editorial. Mark Young (Fig. 2) was nominated for his work on the iconic Diplodocus, which, as a sauropod dinosaur, belonged to the largest terrestrial herbivores to ever have roamed Earth. Their pure size may have pushed them to the evolutionary limits of vertebrate biomechanics. In his paper published in Naturwissenschaften (Young et al. 2012), Mark and his colleagues presented a compelling multidisciplinary approach demonstrating that the gigantic Diplodocus was unlikely to have employed its bizarre dentition for a previously suggested bark-stripping hypothesis feeding strategy, and that the rather unusual (aberrant) dentition of Diplodocus may have, instead, demonstrated an adaptation for food procurement rather than an adaptation to high mechanical bite force. This would suggest that Diplodocus had a skull which was seemingly over-engineered for muscle-driven biting alone; the reason for which may have been related to other behavioral or ecological traits. This paradigmchanging discovery was only possible by making use of modern technology, the computed tomographic scanning of a fossil specimen of a Diplodocus longus skull from museum collection, and subsequent modeling of its biomechanics of feeding. The Arnold Berliner Award was established in recognition of the founding editor of the journal. The award is sponsored by Springer Science+Business Media and is given annually for the best research article published in NAWI during the previous calendar year (Thatje 2012). Criteria are excellence in science, originality, and in particular interdisciplinarity

We report the results of a simple experiment to determine whether honeybees feed their small hive beetle nest parasites. Honeybees incarcerate the beetles in cells constructed of plant resins and continually guard them. The longevity of incarcerated beetles greatly exceeds their metabolic reserves. We show that survival of small hive beetles derives from behavioural mimicry by which the beetles induce the bees to feed them trophallactically. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at htpp://dx.doi.org/10.1007/s00114-002-0326-y.

This paper describes experiments on the locomotor activity rhythm of queens of the ant species Camponotus compressus, which were performed to investigate the consequences of mating on circadian clocks. Locomotor activity rhythm of virgin and mated queens was monitored individually under constant conditions of the laboratory. The locomotor activity rhythm of virgin queens entrained to a 24 h (12:12 h) laboratory light/dark (LD) cycle and free-ran under constant dim red light (RR) with a free-running period ( tau) of approximately 24 h. The locomotor activity of the mated queens on the other hand was arrhythmic during the period when they were laying eggs, and robust rhythmicity appeared soon after the egg-laying phase was over. The tau of the locomotor activity rhythm of mated queens was significantly greater than that of virgin queens. These results are contrary to the commonly held belief that the role of circadian clocks in ant queens ceases after mating flights, thus suggesting that circadian clocks of ant queens are adaptively plastic and display activity patterns, perhaps depending on their physiological state and tasks in the colony.

The ceratopsian dinosaur Turanoceratops tardabilis, from the Upper Cretaceous of Uzbekistan, has been assigned variously as a member of Ceratopsidae (the clade including Triceratops and relatives; Nessov et al. 1989), sister taxon to Ceratopsidae (e.g., Sereno 1997), or a nomen dubium (e.g., Dodson et al. 2004). Cladistic affiliation is of considerable importance for this problematic taxon because all other members of the diverse Ceratopsidae are restricted to western North America. In a recent paper published in Naturwissenschaften, (Sues and Averianov 2009) described newly recovered specimens of Turanoceratops and conducted a cladistic analysis of Ceratopsia. Based on their study, the authors concluded that Turanoceratops is indeed a member of the less inclusive Ceratopsidae and, thus, represents “the first definite ceratopsid dinosaur recorded from Asia.” This result has important implications for the biogeography, origin, and evolution of horned dinosaurs. The ingroup taxa used for their phylogenetic analysis of Turanoceratops (Sues and Averianov 2009) included a thorough representation of non-ceratopsid ceratopsians, but ceratopsids were restricted to two genera (Centrosaurus and Triceratops). Additionally, the source matrix (Makovicky and Norell 2006) was not focused on Ceratopsidae, so many informative phylogenetic characters pertaining to the clade were not included. In order to better evaluate the phylogenetic relationships of Turanoceratops relative to Ceratopsidae, we scored the taxon for the matrix previously published by Dodson et al. (2004). This matrix includes Protoceratops as an outgroup, Zuniceratops, and 13 ceratopsid genera. Eight out of 73 characters were scored for Turanoceratops as follows (character number followed by state in parentheses): 21(1), 22(0), 25(0), 29(0), 57(0), 60(1), 61(1), and 62(0). The matrix was analyzed in PAUP* (Swofford 2003), and three equally parsimonious trees (length=85, consistency index=0.9059, retention index=0.9437) were recovered (Fig. 1). In all three trees, Turanoceratops fell outside of Ceratopsidae (defined as all taxa descended from the most recent common ancestor of Triceratops and Centrosaurus; Dodson et al. 2004), and an additional two steps were required to move Turanoceratops within Ceratopsidae. Turanoceratops lacks at least two features shared by all ceratopsids in this data set: a true supracranial sinus complex and reduced subsidiary ridges on the teeth. The postorbital is remarkably similar to that of the derived neoceratopsian Zuniceratops, particularly in the presence of Naturwissenschaften (2009) 96:869–870 DOI 10.1007/s00114-009-0543-8

[Extract] Studies of the organization of search in ants suggest that individual and colonylevel patterns are related to foraging ecology and may be adaptive [1, 2]. The organization of search in termites has received far less attention, in part because of their cryptic, subterranean foraging habits. Search in subterranean termites involves the construction of a gallery system, a series of branching tunnels in the soil that brings termites into contact with food. The organization of the gallery system might reflect the efficiency of colony search. In this paper we describe the nonrandom colony- level search pattern of the subterranean termite Reticulitermes ftavipes (Kollar), and suggest that it may be organized to minimize search redundancy.

Proteinase inhibitors (PI) occur ubiquitously across microorganisms, plants, and animals [1-3] and have been shown to inhibit "unwanted or foreign" proteolysis and to defend the host tissues against herbivore predation [4, 5]. In plants, PIs are found at very high levels in storage tissues such as tubers and fruits and are argued to serve as defense against herbivores [3, 5]. Recently, Pearce et al. [6] showed that PI accumulation in wild tomato followed a parabolic pattern with age of the fruit; the PI level was low at early (2-3 weeks after anthesis) and late (ripened) stages of fruit growth, but very high at peak fruit growth. They suggested that such a pattern of accumulation may help deter premature dispersal of seeds by dispersers. In this paper, we extend this idea and argue that the dispersal mode of a spedes may strongly influence both the level and the temporal pattern of PI accumulation in its fruit. Analyzing the trypsin inhibitors, a major class of PI in 47 plant species representing various dispersal modes, we offer a test of this hypothesis. Fruits of animal-dispersed compared to those of wind/water or passively (explosively) dispersed species are mostly modified into rich storage organs containing a relatively high level of soluble compounds. A characteristic feature of fruits is the temporal separation of the growth of the mesocarp/pericarp (fleshy part) and the embryo [7]. In most species the embryo initiates growth much later than that of the mesocarp. Such a pattern has important implications for the successful dispersal of fl'uit in animaldispersed species. If dispersers remove fruits when much of the mesocarp growth is complete, but the embryo is still immature (usually coinciding with the peak growth stage of the fruit), plants do not gain any dispersal advantage. Selection in these species should favor the preferential removal of fruits by dispersers at a time when the embryo is mature (usually at the ripened stage of fruit) to gain effective dispersal of their seeds. Accordingly, in species where fruits are dispersed by animals, selection can be expected to favor a high accumulation of PI at the peak growth stage of the fruit (to deter dispersers from removing premature fruits) and low or negligible accumulation at the ripened stage of fruit (to encourage dispersers to remove the fruits). In contrast to fleshy fruits, nonfleshy fruits dispersed passively or by wind, do not run the risk of being removed prematurely by dispersers because, unlike fleshy fruits, their pericarp is not generally modified into rich storage organs. Thus, in these species, selection should not favor the accumulation of PI in fruits. Further, for the same reason, though the temporal pattern of growth of pericarp and embryo in nonfleshy fruit is also separated as in the fleshy fruits, selection should not favor any temporal pattern of PI accumulation in fruits dispersed passively or by wind. Randomly collected fruits of species representing the three dispersal modes, animal, wind/water, and passive (Table 1), available at the Botanical Garden, University of Agricultural Sciences, Bangalore (12 ° 58'N, 77 ° 35'E), were assessed for a major class of PI, namely, trypsin inhibitors (TI). The TI levels were determined in the rues©or pericarp during the peak growth phase of the fruits, when the growth of the fruit tissue was nearly complete. To examine the temporal pattern of TI accumulation for two randomly selected species of each dispersal mode, the TI levels of the fruit tissue were estimated at three stages of growth, namely, immediately following fertilization; at the peak growth stage; and at the ripened stage. Five hundred mg of the fruit tissue (mesocarp or pericarp) from randomly collected fruits of each species was macerated in 2.5 ml of 0.01 M potassium phosphate buffer (pH 7) containing 1 M KC1, and centrifuged at 18000 g for 10 min at 4 °C. The supernatant was depigmented by gel filtering through a Sephadex G-25 column (1 x 10 cm), equilibrated with 0.01 M phosphate buffer containing 0.15 M NaC1 (pH 7.6); 1-ml fractions were eluted with the same buffer. The fractious containing proteins, as detected by the dye-binding method [8], were pooled and analyzed for the presence of trypsin inhibitors. The trypsin inhibitors in the samples were estimated according to [9]. The data were expressed as trypsin inhibitory units (TIU) per gram fresh weight as

Several types of nesting associations, termed quasisocial (females more or less identical, no division of labor), semisocial (females of the same generation, with division of labor) and matrifilial (overlap of generations, with division of labor) are described in many primitively social hymenopteran species. In the latter two types, at least one potentially reproductive female forgoes her own reproduction in favor of that of a dominant female. Quasisociality is generally of a temporary nature, because dominance relations develop. It turns then into semisociality [11. The large carpenter bee Xylocopa sulcatipes in southern Israel has been described as forming both solitary and social nests [2, 3]. This paper shows that in certain associative nests founded by two females an extreme example of reproductive competition, oophagy, occurs. A X. sulcatipes egg measures approximately 11 mm in length and 2.2 mm in diameter; for an adult female of 22 mm body length, egg cannibalism therefore represents a significant loss of investment. X. su/catipes typically nests in twigs o f dead branches where the female excavates a straight tunnel. Females also use pre-existing cavities in stems of Arundo, Ferula and other plants existing naturally in the habitat. The female builds a linear array of cells [4]. Each is provisioned with pollen and nectar, and an egg is deposited upon the loaf-like food mass. Finally the cell is sealed with masticated wood. As part of a study on Xylocopa at Hazeva Field Centre, Arava Valley

Distinctive cues are predicted to evolve when the benefits obtained by the recognition process overcome its costs. When individual recognition is particularly beneficial for both senders and receivers, the expression of strongly distinctive signals is predicted to evolve. On the other hand, it could be predicted that each individual should show a very stable individual signature. In the same perspective, a great stability of the individual signatures could be expected. Lemur catta is the first non-human primate in which olfactory individual recognition has been demonstrated on the basis of the specialized brachial gland secretions. In this paper, I performed gas chromatograph analyses of right and left gland samples collected in two different periods (breeding and non-breeding seasons) from seven males. The aim was to verify if a diversification in such cues, already demonstrated at the inter-individual level, also occurs at the intra-individual level between left and right glands. I verified, by discriminant analysis and chemical distance comparisons, that each gland of each lemur has its particular signature that is maintained through time. Moreover, such diversification resulted so marked to make the overall intra-individual chemical differences similar to/as strong as the inter-individual ones. Since in rodents several odors from different glands may be integrated in individual recognition, I suggest that bilateral diversification in L. catta scents may offer an enhanced distinctiveness that could provide benefits in mate choice and social relationships.