Allometric growth in juvenile marine turtles: possible role as an antipredator adaptation

Female marine turtles produce hundreds of offspring during their lifetime but few survive because small turtles have limited defenses and are vulnerable to many predators. Little is known about how small turtles improve their survival probabilities with growth though it is assumed that they do. We reared green turtles (Chelonia mydas) and loggerheads (Caretta caretta) from hatchlings to 13 weeks of age and documented that they grew wider faster than they grew longer. This pattern of allometric growth might enable small turtles to more quickly achieve protection from gape-limited predators, such as the dolphinfish (Coryphaena hippurus). As a test of that hypothesis, we measured how dolphinfish gape increased with length, reviewed the literature to determine how dolphinfish populations were size/age structured in nearby waters, and then determined the probability that a small turtle would encounter a fish large enough to consume it if it grew by allometry vs. by isometry (in which case it retained its hatchling proportions). Allometric growth more quickly reduced the probability of a lethal encounter than did isometric growth. On that basis, we suggest that allometry during early ontogeny may have evolved because it provides a survival benefit for small turtles.     

Alternative routes to the leader male role in a multi-level society: follower vs. solitary male strategies and outcomes in hamadryas baboons

The nested one-male units (OMUs) of the hamadryas baboon are part of a complex social system in which "leader" males achieve near exclusive mating access by forcibly herding females into permanent consortships. Within this multi-level social system (troops, bands, clans and OMUs) are two types of prereproductive males--the follower and solitary male--whose different trajectories converge on the leader role. Here we compare OMU formation strategies of followers, who associate with a particular OMU and may have social access to females, with those of solitary males, who move freely within the band and do not associate regularly with OMUs. Data were derived from 42 OMU formations (16 by followers and 26 by solitary males) occurring over 8 years in a hamadryas baboon band at the Filoha site in Ethiopia. "Initial units" (IUs) with sexually immature females (IU strategy) were formed by 44% of followers and 46% of solitary males. The remaining followers took over mature females when their leader was deposed (challenge strategy) or disappeared (opportunistic strategy), or via a seemingly peaceful transfer (inheritance strategy). Solitary males took over mature females from other clans and bands, but mainly from old, injured or vanished leaders within their clan (via both the challenge and opportunistic strategies). Former followers of an OMU were more successful at taking over females from those OMUs than any other category of male. Despite this advantage enjoyed by ex-follower leaders, ex-solitary leaders were equally capable of increasing their OMU size at a comparable rate in their first 2 years as a leader. These results demonstrate the potential for males to employ both multiple roles (follower vs. solitary male) and multiple routes (IU, inheritance, challenge, opportunistic) to acquire females and become a leader male in a mating system characterized by female defense polygyny in a competitive arena.     

Variation in social behavior within a spider mite genus, Stigmaeopsis (Acari: Tetranychidae)

The spider mites belonging to the genus Stigmaeopsis constructextremely dense oval woven roofs (web-nests) over depressionson the lower surface of host leaves and are known to have akind of sociality. The four species that occur on bamboo plantsin Japan show different nest areas. The nest area of Stigmaeopsislongus is the largest, followed by that of S. celarius, S. takahashii,and S. saharai in decreasing order. Smaller nests effectivelyprevent adults of several predator mite species from intruding.We hypothesized that variation in nest size reflects differentanti-intruder adaptations of this mite group in relation totheir sociality. The larger nest makers may adopt an alternativeanti-intruder strategy, namely, counterattack by a large group,so as to compensate for the disadvantage of large nests. S.longus and S. celarius adults effectively defended their largenests against potential predators, and the effects of nest defenseincreased with the number of individuals in a nest. S. takahashiiand S. saharai revealed no counterattack effect. Counterattackabilities that increase with the adult density, and thus, socialitymay compensate for the vulnerability of larger nests.     

Ecological immunology: life history trade-offs and immune defense in birds

There has been considerable recent interest in the effects oflife-history decisions on immunocompetence in birds. If immunocompetenceis limited by available resources, then trade-offs between investmentin life-history components and investment in immunocompetencecould be important in determining optimal life-history traits.For this to be true: (1) immunocompetence must be limited byresources, (2) investment in life-history components must benegatively correlated with immunocompetence, and (3) immunocompetencemust be positively correlated with fitness. To gather such empiricaldata, ecologists need to be able to measure immunocompetence.We review techniques used to measure immunocompetence and howthey are applied by ecologists. We also consider the componentsof the immune system that constitute immunocompetence and evaluatethe possible consequences of measuring immunocompetence in differentways. We then review the empirical evidence for life-historytrade-offs involving immune defense. We conclude that thereis some evidence suggesting that immunocompetence is limitedby resources and that investment in certain life-history componentsreduces immunocompetence. However, the evidence that immunocompetenceis related to fitness is circumstantial at present, althoughconsistent with the hypothesis that immunocompetence and fitnessare positively correlated. We argue that future work needs toexamine the fitness effects of variation in immunocompetenceand suggest that artificial selection experiments offer a potentiallyimportant tool for addressing this issue.     

Glow-worm larvae bioluminescence (Coleoptera: Lampyridae) operates as an aposematic signal upon toads (Bufo bufo)

It is an established fact that the spectacular bioluminescentdisplays of adult fireflies and glow-worms are used as courtshipsignals; however, the survival value of the glowing behaviorof their larvae remained the subject of speculation for manyyears. Our study is the first that demonstrates that lampyridlarvae use luminescence to signal unpalatability to nocturnal,visually guided predators. Wild-caught toads (Bufo bufo) weremore reluctant to attack luminescent artificial prey, and weshow that avoidance learning increased this reluctance. Afterbeing exposed to glow-worm larvae (Lampyris noctiluca), whichthe toads experienced as disagreeable, attack latencies to luminescentprey increased, but not those to nonglowing prey. Not all toadsshowed avoidance learning to the same extent, because of eitherdifferences in previous experience with glow-worms or differencesin memory.     

Antagonistic antiparasite defenses: nest defense and egg rejection in the magpie host of the great spotted cuckoo

Brood parasites dramatically reduce the reproductive successof their hosts, which therefore have developed defenses againstbrood parasites. The first line of defense is protecting thenest against adult parasites. When the parasite has successfullyparasitized a host nest, some hosts are able to recognize andreject the eggs of the brood parasite, which constitutes the secondline of defense. Both defense tactics are costly and would be counteractedby brood parasites. While a failure in nest defense implies successfulparasitism and therefore great reduction of reproductive successof hosts, a host that recognizes parasitic eggs has the opportunityto reduce the effect of parasitism by removing the parasiticegg. We hypothesized that, when nest defense is counteractedby the brood parasite, hosts that recognize cuckoo eggs shoulddefend their nests at a lower level than nonrecognizers becausethe former also recognize adult cuckoos. Magpie (Pica pica) hoststhat rejected model eggs of the brood parasitic great spottedcuckoo (Clamator glandarius) showed lower levels of nest defensewhen exposed to a great spotted cuckoo than when exposed toa nest predator (a carrion crow Corvus corone). Moreover, magpiesrejecting cuckoo eggs showed lower levels of nest defense againstgreat spotted cuckoos than nonrecognizer magpies, whereas differencesin levels of defense disappeared when exposed to a carrion crow.These results suggest that hosts specialize in antiparasitedefense and that different kinds of defense are antagonistically expressed.We suggest that nest-defense mechanisms are ancestral, whereasegg recognition and rejection is a subsequent stage in the coevolutionaryprocess. However, host recognition ability will not be expressedwhen brood parasites break this second line of defense.     

Exploiting new terrain: an advantage to sociality in the slime mold Dictyostelium discoideum

Understanding the ecological benefits of social actions is centralto explaining the evolution of social behavior. The social amoebaDictyostelium discoideum has been well studied and is a modelfor social evolution and development, but surprisingly littleis known about its ecology. When starving, thousands of thenormally solitary amoebae aggregate to form a differentiatedmulticellular organism known as a slug. The slug migrates towardthe soil surface where it metamorphoses into a fruiting bodyof hardy spores held up by a dead stalk comprising about one-fifthof the cells. Multicellularity in D. discoideum is thought tohave evolved to lift the spores above the hazards of the soilwhere spores can be picked up for long-distance dispersal. Here,we show that multicellularity has another advantage: local dispersalto new food sources. We find that cells shed by D. discoideumslugs during migration consume and remove bacteria in the pathof the slug, although slugs themselves do not breakup. We alsoshow that slugs are adept at local dispersal by comparing migrationof slugs with migration of individual cells of the mutant, CAP2,which cannot aggregate and so rely only on cellular movement.In particular, the solitary cells of the aggregation mutantare unable to cross a soil barrier, easily crossed by slugs.We propose that the exploitation of local food patches is animportant selective benefit favoring multicellular cooperationin D. discoideum.     

Paternity and paternal effort in the pumpkinseed sunfish

Theoretical models suggest that males should adjust their parentaleffort according to paternity when parental effort is costly,paternity varies among clutches, and males have a cue to assesspaternity. To date, nearly all tests of this theory have beenconducted using birds as model organisms. In this study we examinedthese three factors and the relationship between paternity andmale parental care in a fish system. In the pumpkinseed sunfish(Lepomis gibbosus), parental care is provided exclusively bymales (parentals), but some males (sneakers) parasitize othersby sneaking fertilizations. Parental males significantly lostweight during the parental care period. Clutch size and amountof parental effort did not affect a male's probability of obtainingmore eggs. Paternity was variable among broods. The proportionof young sired by a parental male was not associated with frequencyof fanning eggs or defense of hatched young, but was positivelycorrelated with levels of nest defense during the egg stage.Egg survivorship might restrict an adjustment of fanning behavior,and a general decline in parental behavior (with brood age)might explain the lack of adjustment once the eggs hatch. Parentalmales did not adjust their care when we experimentally manipulatedone possible cue of paternity. Together, these results indicatethat male pumpkinseeds do adjust their care in relation to paternity,but the cues used to assess paternity are not clear.     

Parasitic spawning in sand gobies: an experimental assessment of nest-opening size, sneaker male cues, paternity, and filial cannibalism

Sneaking is common in nest-building fish with paternal care,but the role of nest-opening size in protecting against entryby sneaker males has never been tested before. Using the sandgoby (Pomatoschistus minutus), a fish with exclusive paternalcare, experimental manipulations of nest openings provided nosupport for the hypothesis that nest openings serve as physicalor visual defense or that sneaker males prefer to parasitizenests with wide openings. Female mating preference was alsonot influenced by nest-opening size. However, female courtshipbehavior and visibility were important cues for sneaker males.Most sneak entries occurred when the nest holder was occupiedwith courtship, chasing another sneaker male or nest building.In half the cases of observed sneak entry, the sneaker malefertilized eggs, also when sneaking only occurred before spawning.Sneak entry and its duration were good estimates of stolen paternity,but neither sneak entries nor degree of fertilizations werecorrelated with filial cannibalistic behavior. Testes size didnot explain parasitic spawning success in replicates with geneticallydetermined sneaking. However, all sneaker males without breedingcoloration had huge testes and small sperm duct glands, whereasnest-holding males had small testes and large sperm duct glands,and sneaker males with breeding coloration were intermediate.     

Cornicle secretions of Uroleucon nigrotuberculatum (Homoptera: Aphididae) as the last bullet against lady beetle larvae

Aphids have evolved various defense strategies against natural enemies, including secretions from their cornicles. We assessed the defensive function of cornicle secretions by the goldenrod aphid, Uroleucon nigrotuberculatum (Olive), against larvae of the lady beetles Coccinella septempunctata bruckii Mulsant and Propylea japonica (Thunberg) (Coleoptera: Coccinellidae). The aphid secreted red droplets from its cornicles when attacked by the larvae. Two‐thirds of the C. septempunctata bruckii larvae and 46.7% of the P. japonica larvae that preyed on the aphids died before reaching the pre‐pupal stage. The secretions caused molting failure when smeared on the larvae's heads or glued to the larvae's mouthparts, killing 56.7% of C. septempunctata bruckii larvae and 36.7% of P. japonica larvae. Second instar larvae were affected most. About 40% of third and fourth instar larvae of C. septempunctata bruckii vomited soon after ingesting the aphids. In the field, up to 40% of first and second instar larvae were smeared with red secretions. Our results show that these cornicle secretions are an effective and active defense against earlier instars of coccinellid larvae.     

Hosts are ahead in a marine host-parasite coevolutionary arms race: innate immune system adaptation in pipefish syngnathus typhle against Vibrio phylotypes

Microparasites have a higher evolutionary potential than their hosts due to an increased mutation rate and a shorter generation time that usually results in parasites being locally adapted to their sympatric hosts. This pattern may not apply to generalist pathogens as adaptation to sympatric host genotypes is disadvantageous due to a narrowing of the host range, in particular under strong gene flow among host populations. Under this scenario, we predict that the immune defense of hosts reveals adaptation to locally common pathogen phylotypes. This was tested in four host populations of the pipefish Syngnathus typhle and associated bacteria of the genus Vibrio. We investigated the population divergence among host and bacteria populations and verified that gene flow is higher among host populations than among parasite populations. Next, we experimentally assessed the strength of innate immune defense of pipefish hosts using in vitro assays that measured antimicrobial activity of blood plasma against sympatric and allopatric Vibrio phylotypes. Pipefish plasma displays stronger antimicrobial activity against sympatric Vibrio phylotypes compared to allopatric ones. This suggests that host defense is genetically adapted against local bacteria with a broad and unspecialized host spectrum, a situation that is typical for marine systems with weak host population structure.     

Burkholderia multivorans acts as an antagonist against the growth of Burkholderia pseudomallei in soil

In this study, it was demonstrated, by using agar diffusion tests and a Transwell system, that Burkholderia multivorans NKI379 has an antagonistic effect against the growth of B. pseudomallei. Bacterial representatives were isolated from agricultural crop soil and mixed to construct a partial bacterial community structure that was based on the results of reproducible patterns following PCR-denaturing gradient gel electrophoresis analysis of total soil chromosomes. The antagonistic effect of B. multivorans on B. pseudomallei was observed in this imitate community. In a field study of agricultural crop soil, the presence of B. pseudomallei was inversely related to the presence of the antagonistic strains B. multivorans or B. cenocepacia. B. multivorans NKI379 can survive in a broader range of pH, temperatures and salt concentrations than B. pseudomallei, suggesting that B. multivorans can adapt to extreme environmental changes and therefore predominates over B. pseudomallei in natural environments.     

Microhabitat selection as an antipredator strategy in the aquatic insect Pachydiplax longipennis Burmeister (Odonata: Libellulidae)

Summary An investigation of the larval dragonfly fauna associated with the plant, Sagittaria platyphylla, was conducted in a small pond. Despite the presence of several larval anisopteran species in the pond, only Pachydiplax longipennis larvae were found on Sagittaria plants. A study of the microspatial distribution of P. longipennis larvae on S. platyphylla indicated that larvae use the various regions of a plant in a highly non-random fashion. Larvae show a strong preference for the leaf axil area. A generalized predator, the bluegill sunfish (Lepomis macrochirus), was allowed to selectively eat either of two larvae placed in various plant regions. This experiment indicated that larvae in a leaf axil area were significantly less susceptible to bluegill predation than larvae positioned in other plant regions. The microspatial distribution of starved larvae revealed that larvae with high hunger levels occupied the leaf axil area significantly less than well fed larvae, suggesting 1) larvae do not use these regions as feeding sites, and 2) high hunger levels may induce a behavioral shift in habitat use, with starved larvae forced into areas of high predation risk by the need to fulfill nutritional requirements.     

Alternative models of vertebrate speciation in Amazonia: an overview

The main hypotheses proposed to explain barrier formation separating populations and causing the differentiation of vertebrate species in Amazonia are based on different (mostly historical) factors, as follows. (1) Changes in the distribution of land and sea or in the landscape due to tectonic movements or sea-level fluctuations (Paleogeography hypothesis). (2) The barrier effect of Amazonian rivers (River hypothesis). (3) A combination of the barrier effect of broad rivers and vegetational changes in Northern and Southern Amazonia (River-refuge hypothesis). (4) The isolation of forest blocks near areas of surface relief in the periphery of Amazonia during dry climatic periods of the Tertiary and Quaternary (Refuge theory). (5) Competitive species interactions and local species isolations in peripheral regions of Amazonia due to invasion and counterinvasion during cold/warm periods of the Pleistocene (Disturbance-vicariance hypothesis). (6) Parapatric speciation across steep environmental gradients without separation of the representative populations (Gradient hypothesis). Several of these hypotheses are probably relevant to a different degree for the speciation processes in different faunal groups or during different geological periods. The paleogeography hypothesis refers mainly to faunal differentiation during the Tertiary and in combination with the Refuge hypothesis; Milankovitch cycles leading to global climatic-vegetational changes affected the biomes of the world not only during the Pleistocene but also during the Tertiary and earlier geological periods. New geoscientific evidence for the effect of dry climatic periods in Amazonia supports the predictions of the Refuge theory.     

Stable group size in cooperative breeders: the role of inheritance and reproductive skew

Recent studies of reproductive skew have revealed great variationin the distribution of direct fitness among group members, yetthere have been surprisingly few attempts to explore the consequencesof such variation for stable group size, and none that takeinto account the future benefits of group membership to nonbreeders.This means that the existing theory is not suited to explainthe group size of most cooperatively breeding vertebrates andprimitively social insects in which group membership involvessubstantial future benefits. Here we model the group size ofsuch species as social queues in which nonbreeders can inherita breeding position if they outlive those ahead of them in thequeue. We demonstrate, however, that the results can be generalizedto systems in which inheritance occurs via scramble competition,rather than via a strict queue. The model predicts that stablegroup size will depend on the number of breeding positions inthe group and the mortality rates of breeders and nonbreeders,but not on the distribution of reproduction among the pool ofbreeders. This is because deaths occur at random, so that eachindividual has the same chance of surviving to reach each breedingposition. We tested a specific prediction of the model usingdata on ovarian development in the paper wasp, Polistes dominulus.We found a positive correlation between group size and the proportionof females with fully developed eggs, as predicted. Our resultsclarify the interaction between the dominance structure andsize of animal groups and add to the growing recognition ofthe potential for inheritance as a major determinant of bothindividual behavior and group-level characteristics of animalsocieties.     

Effect of soil hardness on aggression in the solitary wasp Mellinus arvensis

1. Two alternative nesting strategies are exhibited by soil‐nesting Mellinus arvensis females, digging a new nest (diggers) and searching for an old unoccupied burrow (searchers). Wasps appear unable to distinguish between occupied and unoccupied nests, and aggressive interactions between searchers and nest owners at nest entrances are frequent. 2. In aggressive encounters, there is an advantage in size and residency status. 3. The costs associated with the two nesting strategies vary across geographically separated populations: nest digging incurs costs in terms of time, and these vary according to the hardness of the soil substrate; nest searching is variably costly in terms of risk of injury in aggressive encounters with nest‐owning females. 4. Individual female wasps can switch between nesting strategies, and thus soil hardness, by affecting the cost of nest construction, affects the relative frequencies of the two nesting strategies within a population, favouring an increase in the searching strategy. This, in turn, affects the frequency and intensity of aggression between females at a nesting aggregation. 5. Female body size is correlated with soil hardness. As aggressive encounters are more frequent in sites with hard soil substrates, there is increased selective advantage in having large body size at these sites. 6. Body size is determined primarily by the availability of food resources during larval development, which is, to a degree, a function of the size of the adult female. There is a trade‐off between provisioning a few cells with many provisions in each, leading to the development of few but large adults, as opposed to many cells with few provisions, leading to many small offspring. The relative advantage of these two provisioning strategies is, at least in part, a function of the hardness of the soil substrate.     

Breeding suppression in the bank vole as antipredatory adaptation in a predictable environment

Summary In northern Fennoscandia, microtine rodent populations fluctuate cyclically. The environment of an individual vole can be considered to be predictable when the risks of predation and intra- and interspecific competition change with the cycle, such that both are high during the population highs of voles. The risk of predation is also high during the vole crash. After the crash, the vole population is characterized by low intra- and interspecific competition and low predation pressure. The main predators affecting voles during the crash are the small mustelids, least weasel and stoat. The density of these specialist predators declines drastically during the winter after the vole crash. We studied experimentally the impact of the perceived presence of stoats on the breeding and mating behaviour of voles. In a series of breeding experiments with bank voles,Clethrionomys glareolus, both old and young females suppressed breeding when exposed to the odour of stoats,Mustela erminea. The weights of females decreased in both experimental and control groups, but more among the voles under odour exposition. It seems that females actively avoided copulations under high predation risk and that breeding suppression is mediated by a change in female mating behaviour. There was no change in male behaviour or physical condition between the experimental and control treatments. An alternative mechanism for the observed breeding suppression could be the one caused by decreased feeding in females mediated with low energy intake which does not allow breeding. Regardless of its mechanism, delay of breeding should increase the probability of non-breeding females to survive to the next breeding season. The females surviving the crash should gain a strong selective advantage in a predator-free environment of the subsequent breeding season, which could explain the adaptive function of this antipredatory strategy.     

Enemy release but no evolutionary loss of defence in a plant invasion: an inter-continental reciprocal transplant experiment

Plant chemical defenses and escape from natural enemies have been postulated to select for dietary specialization in herbivorous insects. In field and laboratory bioassays, we evaluated the effectiveness of intact and chemically modified larval shield defenses of the generalist Chelymorpha alternans and the specialists Acromis sparsa and Stolas plagiata (Chrysomelidae: Cassidinae) against three natural predators, using larvae reared on two morning glory (Convolvulaceae) species. We assessed whether: (1) specialists were better defended than generalists when both were fed and assayed on the same plant; (2) larval shield defenses were chemical, physical, or both; and (3) specialists exploit chemistry better than generalists. Live specialist larvae survived at higher rates than did generalists in predator bioassays with the bug Montina nigripes (Reduviidae), but there were no differences among groups against two species of Azteca ants (Hymenoptera: Dolichoderinae). Solvent leaching by H₂O or MeOH significantly reduced shield efficacy for all species compared to larvae with intact shields. In contrast, freshly killed specialist larvae exhibited significantly lower capture rates and frequencies than the generalists. Although solvent leaching significantly reduced overall shield efficacy for freshly killed larvae of all species, the pattern of leaching effects differed between specialists and generalists, with H₂O-leaching having a greater impact on the specialists. The overall vulnerability of the generalists appears due to lower chemical protection, which is ameliorated by increased escape behaviors, suggesting a selective trade-off between these defensive components. These experiments indicate that shield defenses are essential for larval survival and that specialists are superior at exploiting plant compounds residing in the aqueous fraction. Our results support the hypothesis that diet-specialized herbivorous insects have more effective defenses than generalists when both feed on the same plant due to the differential ability to exploit defensive precursors obtained from the host. The evolution of dietary specialization may therefore confer the advantage of enhanced enemy-free space.