Interference competition among burying beetles (Silphidae, Nicrophorus)

Abstract. 1. This study investigated the impact of intraspecific and interspecific competition on the reproductive success of a biparental burying beetle, Nicrophorus defodiens Mannerheim.
2. Marked pairs or single females were placed on small and large mouse carcasses in the field in 1985 and 1986. Carcasses were exhumed after 9–10 days to determine the identity of the resident adult(s) and the production of young.
3. Competition was assessed by the prevalence of takeovers by intruders (unmarked adults). For N.defodiens , after the initial colonization of the carcass, interspecific competition from larger N.orbicollis Say and N.sayi Laporte was substantial and more intense than intraspecific competition. Competition was also greater in the middle of the breeding season and on large as opposed to small carcasses.
4. Successful takeovers resulted in the expulsion of the prior resident(s), killing of any offspring present on the carcass, and oviposition of a new clutch by the intruder.
5. Females aided by males were more likely than single females to avoid takeovers but did not produce larger broods or larvae of larger mass.
6. An additional laboratory experiment in 1985 and a field experiment in 1986 suggest that N.defodiens is able to reproduce on very small carcasses despite intense heterospecific activity.     

Competition between the nymphs of two regionally co-occurring species of Notonecta (Hemiptera: Notonectidae)

1. Two species of freshwater invertebrate predator, Notonecta maculata and N. obliqua , showed a negative association in a series of small, man-made ponds in the Peak National Park, Derbyshire, U.K. The present study examines the potential role of interspecific interactions among nymphs on this regional distribution pattern.
2. The survival, development and feeding efficiency of nymphs were examined in laboratory and field mesocosm experiments with intra- and interspecific competition and contrasting environmental complexity.
3. Survival to adulthood and mean lifespan varied significantly in interspecific competition treatments in both laboratory and field experiments, with N . maculata showing higher survival in the simple environment and N. obliqua higher survival in the complex environment.
4. Variations in feeding efficiency were consistent with the survival trends: N. maculata had a higher efficiency in the simple environment, whereas N. obliqua had greater efficiency in the complex environment. There was evidence of a developmental response in feeding efficiency, with differences between species increasing with age.
5. These results suggest that the relative competitive abilities of the two species are affected by habitat complexity, and that competition between species may modify the species distribution where they co-occur.     

A chemically triggered transition from conflict to cooperation in burying beetles

Although interspecific competition has long been recognised as a major driver of trait divergence and adaptive evolution, relatively little effort has focused on how it influences the evolution of intraspecific cooperation. Here we identify the mechanism by which the perceived pressure of interspecific competition influences the transition from intraspecific conflict to cooperation in a facultative cooperatively breeding species, the Asian burying beetle Nicrophorus nepalensis. We not only found that beetles are more cooperative at carcasses when blowfly maggots have begun to digest the tissue, but that this social cooperation appears to be triggered by a single chemical cue – dimethyl disulfide (DMDS) – emitted from carcasses consumed by blowflies, but not from control carcasses lacking blowflies. Our results provide experimental evidence that interspecific competition promotes the transition from intraspecific conflict to cooperation in N. nepalensis via a surprisingly simple social chemical cue that is a reliable indicator of resource competition between species.     

Can the "challenge hypothesis" be applied to insects?

The challenge hypothesis, conceived for testosterone and vertebrates, has recently been applied to juvenile hormone (JH) and insects. Scott [Scott, M.P., 2006a. Resource defense and juvenile hormone: the "challenge hypothesis" extended to insects. Horm. Behav. 49, 276-281] found that JH in the burying beetle Nicrophorus orbicollis increased in response to a social challenge in the presence of a breeding resource, while there was no such JH response in the purported brood parasite Nicrophorus pustulatus. Two important implications of the challenge hypothesis that need to be tested are whether JH affects dominance and why there are interspecific differences in the JH response to a social challenge. The effect of JH on dominance in burying beetles was examined by topical application of JH III to one of two competing females at 24 h and again at 1 h prior to presentation of a breeding resource (mouse carcass). JH supplementation had no effect on dominance in intraspecific interactions in N. orbicollis, as measured by possession of the carcass on Day 1, 3 or 7. Similarly, JH did not alter carcass ownership during competition between N. orbicollis and N. pustulatus nor did it affect reproductive success. Irrespective of JH supplementation, N. pustulatus became increasingly dominant as the trials progressed, rarely occupying the carcass on Day 1, but excluding N. orbicollis in nearly half the trials by Day 7. These findings, and a brief review of the burying beetle literature, suggest that the challenge hypothesis, as applied to testosterone and vertebrates, does not yet have an analogous model for JH and insects.     

Resource control by territorial male cichlid fish in Lake Malawi

1. The rocky habitat communities of Lake Malawi contain a high diversity of ecologically similar, predominantly herbivorous, cichlid fish species. How so many species can coexist is still unknown.
2. Adult males of the majority of these species hold permanent territories which form dense multispecies mosaics across the shores.
3. The study examined six coexisting species of cichlids from the rocky habitat where males are known to control access to the algal food resources within their territorial space. These included four sibling species of the Pseudotropheus ( Tropheops ) species complex, which are specialized epilithic algal herbivores, and two sibling species of the trophically more generalist P . ( Maylandia ) species complex which feed upon both epilithic algae and plankton.
4. The hypotheses that both intraspecific and interspecific territoriality occur in these communities and that interspecific aggression initiated by territorial males is preferentially directed at species with the most similar diets to their own were tested.
5. It was found that males preferred to exclude conspecific intruders, while they tolerated intruders with different diets to their own. Also, male P . ( Tropheops ) preferentially excluded similar heterospecific dietary specialists, while male P. ( Maylandia ) tolerated similar heterospecific dietary generalists.
6. Based on this study, it is proposed that interspecific territoriality may be reducing competition between species with different diets and promoting their coexistence, while it may be enhancing competition between species with the same diet. Furthermore, it suggests that ecological generalism may be reducing the intensity of interspecific competition, while specializations for the same resources may be increasing competition.     

Host shift by the burying beetle, Nicrophorus pustulatus, a parasitoid of snake eggs

Recent work [Ecoscience (2000) vol. 7, 395-397] suggests that the burying beetle Nicrophorus pustulatus may have undergone a remarkable host shift, exploiting snake eggs rather than carrion as resources for breeding. We conducted behavioural and physiological experiments to examine the hypothesis of a host shift and to formulate hypotheses on its origin. Two congeners of N. pustulatus, Nicrophorus orbicollis and Nicrophorus defodiens did not respond to snake eggs with typical breeding behaviour. When N. pustulatus male-female pairs (n = 14) were presented with clutches of snake eggs, the number of offspring but not the mean size of offspring varied with snake egg mass, indicating effective regulation of brood size. When breeding on turtle eggs, N. pustulatus had a more variable response than when exploiting snake eggs, suggesting that turtle eggs are not a primary resource for breeding. Nicrophorus pustulatus presented with both snake eggs and a mouse carcass combined and exploited the two resources within the same nest (10 of 12 trials). Mouse carcasses and snake eggs were treated differently. Carcasses were moved, buried and stripped of hair in a manner characteristic of burying beetles, whereas snake eggs were not moved or buried. Females that discovered a mouse carcass also had a significantly greater juvenile hormone increase than did females discovering snake eggs. Some responses to the two resources, however, were similar. Female N. pustulatus oviposited rapidly in response to either a mouse carcass or snake eggs, and males elevated sex pheromone emission in response to either resource. The efficient use of snake eggs, the ability to regulate brood size and the different responses to snake eggs and carrion suggest that N. pustulatus is well adapted to exploiting snake eggs for breeding. The use of snake eggs by N. pustulatus has potential implications for conservation of oviparous reptiles.     

Study of community assembly patterns and interspecific interactions involved in insect succession on rat carcasses

Although many forensic entomological studies have described patterns of carrion insect succession and theoretical studies have explained interspecific interactions that drive succession, empirical studies on the quantitative and ecological aspects of carrion insect succession, such as the degree of historical contingency in community assembly and interspecific interactions during succession, are limited. In this study, I investigated variability in the successional pathways of carrion insect communities in rat carcasses and their decomposition processes, and examined the interspecific interactions involved in succession, such as the effects of carcass utilization by early‐arriving species on late‐arriving species. Members of the families Calliphoridae and Formicidae and the species Eusilpha japonica (Motschulsky) and Nicrophorus concolor Kraatz were chiefly observed. In almost all carcasses, formicid species arrived first, and calliphorid species and E. japonica arrived simultaneously or immediately after. Nicrophorus concolor arrived last, with its time of colonization occurring earlier in carcasses with greater E. japonica abundance. Meanwhile, the early‐arriving species decreased when N. concolor arrived. Nicrophorus concolor tended to reproduce on carcasses with lower cumulative abundance of early‐arriving species and tended to feed on carcasses with greater cumulative abundance of early‐arriving species. These results show that the successional pathways of the chief carrion insect species are highly consistent among carcasses. In contrast, early‐arriving species seem to influence the utilization patterns of carcasses by late‐arriving species, and therefore produce variability in the decomposition process. These results also show that succession could be driven by facilitation and interspecific competition between early‐arriving and late‐arriving species.     

Interspecific competition between two field populations of grass-feeding bugs

Abstract. 1. We tested the hypothesis that interspecific competition between two grass-feeding stenodemine bug species ( Notostira elongata Geoffroy and Megaloceraeu recticornis Geoffroy) was capable of causing population-scale mortality in the field.
2. N.elongata nymphs were added to two field enclosures in which the M.recti-cornis population was to hatch. In one of these enclosures the grass Arrhena-therum elatius L., on which only M.recticornis can feed, was also added. Population changes of the bug species were then monitored. An unmanipulated enclosure and an unfenced plot of grassland were used as controls.
3. The results demonstrated that N.elongata lowered the survival of M.recticornis and that this effect was removed by the addition of A.elatius . The results also suggested that N.elongata suffered less from the effects of competition with M.recticornis when the latter had access to the refuge foodplant.
4. Interspecific competition was rare among members of the grass-feeding stenodemine guild in the area studied. Only one out of fifteen possible pairs competed, and this competition occurred only under special circumstances.     

Inter- and intra-specific dominance relationships and feeding behaviour of Golden Eagles Aquila chrysaetos and Sea Eagles Haliatetus albicilla at carcasses

The behaviour of Golden Eagles Aquila chrysaetos and Sea Eagles Haliceetus albicilla scavenging on artificially laid out carcasses in coastal Sør-Trøndelag Province, Norway, was studied during two winters (totalling 640 h of observations) and the intervening summer (430 h). Neither species fed at carcasses in summer. Although smaller, Golden Eagles were strongly dominant over Sea Eagles in direct competition for carcass access. In Sea Eagles, females dominated males, while in Golden Eagles, few conflicts between birds of known sex were observed. Age effects were weak and not statistically significant in both species. Conflicts for carcass access tended to be most escalated between Golden Eagles and least escalated between Sea Eagles, with interspecific conflicts intermediate. Most conflicts were won by the aggressor, suggesting that birds were generally able to assess relative dominance before launching an attack. Young eagles fed longer at a carcass than older individuals in both species, suggesting that young eagles may have been hungrier or less efficient feeders. Sea Eagles waited longer than Golden Eagles between arrival in the immediate carcass area and feeding at the carcass. This effect was greater when the carcass was already occupied but also occurred when no other eagle was already present. While interspecific competition for carrion did not appear to have important consequences for the two species in coastal Norway, in western Scotland, where Sea Eagles are currently re-establishing, carrion is important in the diet of both species all year round. Interspecific competition for this resource may therefore play a role in determining the ultimate realized niche (and therefore numbers) of the two species in Scotland in the longer term.     

Brood discrimination, nest mate discrimination, and determinants of social behavior in facultatively quasisocial beetles (Nicrophorus spp.)

In this study we investigated ecological determinants of socialityin burying beetles (Nicrophorus spp.), potential conflicts ofinterest among reproductive females, and the effects of nestingfailure and costs of fighting on cooperation. Burying beetlesare known to form monogamous pairs when exploiting small vertebratecarcasses. More complex social behavior in this group is poorlyunderstood. We conducted experiments in which one or two females(N. defodiens, N. orbicollis) were provided small or large carcasseson which to breed. On large but not on small carcasses, twofemales often formed cooperative breeding associations (jointlyprepared a carcass and fed young). In N. defodiens, but notN. orbicollis, two females produced a larger brood than singlefemales on large carcasses. In both species, the reproductiveoutput per female was less for two than for one female. Thepresence of a second female did not decrease the preparationtime of a carcass (discovery of resource to egg hatch). Conflictwas evident between females. Trials employing females of similarsize were more likely to result in injury than trials usingfemales of dissimilar size (N. tomentosus, N. defodiens, N.orbicollis). In N. tomentosus, those associations that persistedthe longest resulted in the fewest injuries. After care of youngwas initiated, conflict among familiar nest mates was not observed.There was no evidence that breeding females could discriminatebetween brood; use of a genetic marker (N. orbicollis) demonstratedthat females fed related and unrelated young alike. Femalesof similar size (high potential cost of fighting for the dominantindividual) were not more likely to form cooperative breedingassociations than females of dissimilar size (low cost of fightingfor dominant). Females of a species subject to a high rate ofnest failure (N. defodiens) were more likely to cooperate thanfemales of a species with a low rate of nest failure (N. orbicollis).It is argued that limited reproductive opportunities, difficultyin controlling rivals' access to a large carcass, and the superabundantlarval food supply represented by a large carcass, but not kinselection, have contributed to the evolution of cooperativebehavior in this group. In addition, we hypothesize that beetlesmight initially tolerate consexual rivals on large carcasseswhen there is a high likelihood of nesting failure, therebyavoiding potentially costly conflicts.     

Patterns of intra- and interspecific association in leaf-mining insects on three oak host species

Abstract. 1. We determined mortality and distributional patterns of leaf miners on three oak host species (Quercus falcata, Q.nigra and Q.hemisphaerica) in northern Florida, U.S.A.
2. Patterns of intra- and interspecific occurrence within leaves, and mortality of five most abundant leaf miner species were analysed as a test of competition.
3. Miners co-occurred on leaves more often that expected by chance (P<0.05) in six of ten possible species combinations and log-linear model analysis showed no negative higher-order interactions.
4. All five miner species had highly clumped distributions between leaves (P<0.01).
5. Leaf miner survival was less than expected for four of five species when co-occurring on leaves with conspecifics than when mining with heterospecifics or alone (P<0.05).
6. We conclude that interspecific competition is unapparent within this leaf miner guild and that intraspecific competition occurs in four of the five major leaf miner species. We discuss leaf miner selection of common leaves, perhaps based on chemical/physical leaf characters, as a cause of intra- and interspecific aggregation.     

Patchiness, aggregation, and the coexistence of competitors for ephemeral resources

Abstract. 1. A laboratory experiment was conducted to determine the outcome of competition between two species of carrion-breeding Diptera for food at two levels of resource patchiness.
2. Adults of Fannia howardi Malloch (Muscidae) and Megaselia scalaris Loew (Phoridae) were released into a large population cage and were given access to three 2g sections of pork kidney in one treatment (low patchiness) and to twelve 0.5g sections in the other (high patchiness).
3. Increasing the patchiness of the resource resulted in reduced overlap in the distribution of the two species, and therefore less interspecific competition. However, this did not result in improved survival of the poorer competitor, F.howardi. Both species aggregated their eggs at high patchiness, and therefore suffered from increased levels of intraspecific competition.
4. The experiment shows that increasing patchiness in this way could lead to prolonged coexistence, as has been predicted in other studies.     

The potential for interspecific competition between two abundant defoliators on oak: leaf damage and habitat quality

Abstract. 1. The pedunculate oak, Quercus robur L., suffers high annual levels of spring defoliation in Wytham Woods, Oxon. The two major defoliators, Tortrix viridana L. and Operophtera brumata L., sometimes reach high enough densities to completely defoliate trees.
2. Experiments with larvae of both species enclosed on oak trees in Wytham Woods suggest considerable potential for intra- and interspecific competition between T. viridana and O. brumata.
3. T. viridana and O. brumata consume equivalent amounts of leaf tissue per unit caterpillar density. Despite this, O. brumata is the superior competitor, measured by mortality during the larval feeding period.
4. Evidence is presented that T. viridana larvae depend upon the protected microclimate of leaf rolls to maintain water balance, and that leaf damage may disrupt the construction of, and microclimate within, leaf rolls. O. brumata may therefore be competitively superior because it indirectly interferes with the water balance of T. viridana.
5. The importance of interspecific competition in the dynamics of T. viridana and O. brumata populations is discussed. Stochastic mortality factors acting on first instar larvae of both species probably make intense competition a rare event.     

施氮对空心莲子草(Alternanthera philoxeroides)和莲子草(Alternanthera sessilis)种间关系的影响

自然界的氮素释放总是呈现出空间和时间上的异质性,但关于异质性氮释放对于入侵植物和本地植物种间关系影响的研究相对较少。将入侵植物空心莲子草(Alternanthera philoxeroides)和同属本地植物莲子草(Alternanthera sessilis)分别进行单种种植(12株,无种间竞争)和混种种植(每种6株,有种间竞争),模拟大气氮湿沉降设置由两种不同施氮总量(15g N m~(-2)a~(-1)和30g N m~(-2)a~(-1))和两种不同施氮频率(每5天1次和每15天1次)交叉组成的4种施氮处理,并以不施氮为对照。施氮总量的增加显著促进了两种植物的生长,但对两种植物的种间竞争关系没有显著影响。施氮频率对两种植物的生长以及种间竞争关系都没有显著影响。两种植物在面对竞争时表现出不同的生物量分配策略,空心莲子草将更多的生物量分配到茎,而莲子草将更多的生物量分配到根。在全球变化的背景下,大气氮湿沉降可能会改变两种植物的种群结构和动态,但可能对这两种植物的种间关系影响较小。     

山西五鹿山白皮松群落乔灌层的种间分离

种间分离的研究对于揭示种间相互作用、群落组成与动态具有重要意义。为了探讨五鹿山自然保护区白皮松林内物种间的关系及其共存机制,运用ArcGIS软件、N×N最近邻体列联表及其2×2列联表截表的方法、Pielou的分离指数,Hegyi单木竞争指数对山西五鹿山白皮松群落55个样方内的15种乔木和灌木的种间分离规律及9种乔木的种内种间竞争关系进行研究,并根据种间分离情况以及物种对环境的适应方式,将样地中15个物种划分为3个生态种组。结果表明:(1)该群落中随机毗邻种对占绝大多数(80%),正分离种对较少(16.19%),负分离种对极少(3.81%),群落趋于稳定。(2)种间分离在不同种之间存在着一定的差异,正分离常发生在群落中的建群种或优势种之间;而负分离常发生在群落中的优势种和一些伴生种之间。(3)物种的种间分离状况与其种内种间竞争及物种的空间分布格局具有密切的关系,正分离常常发生在种内竞争大于种间竞争呈聚集分布的优势种或建群种之间;而负分离常常发生在种间竞争大于种内竞争的优势种和伴生种之间,且同一生态种组内的物种种间竞争较不同生态种组间激烈。(4)种间分离与群落的生境异质性和物种的生态需求也具有密切的关系,即两个具有不同生境要求的物种之间常常发生正分离;而具有相同或相似生境要求的物种之间常常发生负分离。(5)白皮松群落内物种的总体分离规律为全面不分离。(6)由生境异质性和物种间不同的生态适应性引起的种内种间关系是维持五鹿山自然保护区白皮松群落稳定共存的机制。     

Influences of host feeding-niche and foodplant type on generalist and specialist parasitoids

Abstract. 1. We classified the parasitoids of 185 British herbivorous insect species as being koinobionts (which should tend to be specialists) or idiobionts (potential generalists) to examine the influences of host feeding-niche and foodplant type on the numbers of parasitoid species attacking individual host species.
2. The majority of parasitoid species of exophytic hosts are koinobionts, whereas endophytic hosts support mainly idiobionts.
3. Parasitoid assemblage size increases with host foodplant size and complexity; for endophytic hosts this is due to an increase in idiobionts on hosts on large plants, but for exophytic hosts it is the number of koinobionts that increases with foodplant size.
4. Comparison of these patterns with those predicted under a competition hypothesis suggests that parasitoid communities associated with endophytic hosts may be commonly limited by interspecific competition, whereas those of exophytic hosts probably are not.     

The realized food niches of three species of stream-dwelling triclads (Turbellaria)

SUMMARY. 1. The diet of Phagocata vitta, Crenobia alpina and Polycelis felina in a Welsh stream was examined, using a serologicat technique, at monthly intervals over 18 months.
2. For each triclad species, dietary composition changed little with time or increasing body size, and overall feeding intensity was greatest in summer and least in winter.
3. Alt species fed most heavily on oligochaetes, chironomids and Gammarus. Phagocata vitta fed lightly on Plecoptera, Trichoptera and Ephemeroptera whilst the other two triclad species fed substantially on the first two of these prey categories.
4. Food niche breadth indices indicated a wide diet for all species with some specialization in some months. Electivity indices showed positive electivity for oligochaetes and chironomids by all species and for Plecoptera by C. alpina and P. felina .
5. Diets overlapped and none had a food refuge: therefore the potential for interspecific competition for food exists.     

Monogamy to communal breeding: exploitation of a broad resource base by burying beetles (Nicrophorus)

Abstract.

• 1 To investigate the range of resource size that burying beetles (Nicrophorus) exploit, small (21–33 g), medium (50–90 g) and large (120–210g) carcasses, were placed in the field and then exhumed after 1, 4, 8, 12 or 18 days.
• 2 Nicrophorus attempts to utilize carcasses over this entire size range but has greater success on smaller carcasses.
• 3 Larger carcasses were more difficult to exploit because: (a) they took longer to conceal beneath the leaf litter; (b) they were less likely to be rounded into brood balls; (c) they were more likely to be utilized by dipterans; and (d) they were occupied by greater numbers of congeners.
• 4 Larger carcasses, however, did support greater numbers of larvae and contained broods of greater total mass than smaller carcasses.
• 5 Beetles sometimes bred communally on larger carcasses in the field and same-sex adults were observed to feed young.
• 6 Two follow-up experiments were conducted in the laboratory. On a large carcass N.defodiens, N.tomentosus or N.orbicollis can raise a maximum of 35–50 young. Nicrophorus pustulatus, in contrast, appears to be unique among Nicrophorus in that it can raise nearly 200 young on larger carcasses.
• 7 Nicrophorus orbicollis and Nicrophorus sayi are extremely dependent on parental regurgitations and young fail to survive to the second instar if parents are removed. Young of Nicrophorus defodiens, N.tomentosus and N.pustulatus can develop normally without parental regurgitations.
• 8 I discuss these results in the context of reproduction on carcasses of different size and hypothesize that this breeding system is facultatively quasisocial.

     

Competition refuges and coexistence: an example from Serengeti carnivores

1. In the last two decades predator–prey models have shown that 'refuges', in which prey can seek respite from predation, are crucial for the persistence of prey and predator. This concept is equally applicable to interspecific competition and, in a heterogeneous environment, species with low competitive ability should seek out 'competition refuges' where competition is reduced.
2. Cheetahs have low competitive ability compared with their principal competitors, hyenas and lions, which are directly responsible for their low density. This study uses distribution data collected in the Serengeti National Park in Tanzania over a 4-year period to show that cheetahs are more strongly associated with each other than with their competitors and utilize areas with low-density prey.
3. Cheetahs exhibit local avoidance behaviour in both space and time with respect to lions and hyenas. This behaviour is facultative and is strongest when cheetahs are engaged in activities that might expose them to food loss or increase the risk of close interactions, such as when they are hunting or eating.
4. Lactating cheetahs, whose range is restricted, are more likely to have difficulties finding prey and come into more frequent contact with lions than free-ranging animals.
5. It is argued that although cheetahs always lose in direct competition, they persist in the ecosystem by seeking out 'competition refuges' with low densities of lions and hyenas and that their mobility is the key to their continued coexistence with these predators. This pattern of distribution may be generally applicable to other species which, although widely distributed, always occur at low densities.