Why are metapopulations so rare?

Roughly 40 years after its introduction, the metapopulation concept is central to population ecology. The notion that local populations and their dynamics may be coupled by dispersal is without any doubt of great importance for our understanding of population-level processes. A metapopulation describes a set of subpopulations linked by (rare) dispersal events in a dynamic equilibrium of extinctions and recolonizations. In the large body of literature that has accumulated, the term "metapopulation" is often used in a very broad sense; most of the time it simply implies spatial heterogeneity. A number of reviews have recently addressed this problem and have pointed out that, despite the large and still growing popularity of the metapopulation concept, there are only very few empirical examples that conform with the strict classical metapopulation (CM) definition. In order to understand this discrepancy between theory and observation, we use an individual-based modeling approach that allows us to pinpoint the environmental conditions and the life-history attributes required for the emergence of a CM structure. We find that CM dynamics are restricted to a specific parameter range at the border between spatially structured but completely occupied and globally extinct populations. Considering general life-history attributes, our simulations suggest that CMs are more likely to occur in arthropod species than in (large) vertebrates. Since the specific type of spatial population structure determines conservation concepts, our findings have important implications for conservation biology. Our model suggests that most spatially structured populations are panmictic, patchy, or of mainland-island type, which makes efforts spent on increasing connectivity (e.g., corridors) questionable. If one does observe a true CM structure, this means that the focal metapopulation is on the brink of extinction and that drastic conservation measures are needed.     

Does cadence of Iguana iguana displays facilitate individual recognition?

Iguanid lizards communicate threat, courtship and territorial advertisement through stereotyped headbob displays. It has been hypothesized that slight interindividual differences in these displays might enhance individual recognition and maintain dominance relationships within populations, but the precise display elements responsible have not been determined. The purpose of this study was to examine if experimentally induced differences in displays would decrease recognition of individual conspecifics. Specifically, social interactions between adult male green iguanas (Iguana iguana) were examined at normothermic and hypothermic body temperatures. Whereas the cadence (rate of muscle contraction) of stereotypic displays of individual lizards was slowed at hypothermic body temperature (Q ₁₀ 1.6), the amplitude (maximal muscle contraction) of hypothermic displays remained constant relative to normothermic displays. Normothermic lizards viewing a display from a hypothermic lizard responded to that display as if it were performed at a normothermic cadence. During paired encounters between lizards (one hypothermic, one normothermic), dominant/subordinate dyads within the group were unusually maintained regardless of which individual was hypothermic. The exception occurred when dominant individuals at the lowest experimental body temperatures retreated from subordinate individuals. The results suggest that individual lizards recognized their relative social status even when cadence of displays was radically altered, indicating that cadence alone probably does not function in individual recognition. Other morphological, display elements, or scent cues may provide information useful in individual recognition.Communicated by G.M. Klump     

Why do honey bees dance?

The honey bee dance language, used to recruit nestmates to food sources, is regarded by many as one of the most intriguing communication systems in animals. What were the ecological circumstances that favoured its evolution? We examined this question by creating experimental phenotypes in which the location information of the dances was obscured. Surprisingly, in two temperate habitats, these colonies performed only insignificantly worse than colonies which were able to communicate normally. However, foraging efficiency was substantially impaired in an Asian tropical forest following this manipulation. This indicates that dance language communication about food source locations may be important in some habitats, but not in others. Combining published data and our own, we assessed the clustering of bee forage sites in a variety of habitats by evaluating the bees’ dances. We found that the indicated sites are more clustered in tropical than in temperate habitats. This supports the hypothesis that in the context of foraging, the dance language is an adaptation to the particular habitats in which the honey bees evolved. We discuss our findings in relation to spatial aggregation patterns of floral food in temperate and tropical habitats.     

Where are the SVHCs?

Background

The European chemicals regulation REACH includes the legal duty for suppliers to inform consumers on request about the presence of substances of very high concern (SVHCs) in articles. Since this requirement has been in force now for 10 years, the intention of this study was to find out whether information on SVHCs is adequately communicated to the consumer today. Data on the presence of SVHCs in articles were collected as a prerequisite for the subsequent requests for a targeted choice of articles to examine the operability of the ‘right to know.’

Results

Literature data show that SVHCs have been measured and described in a large variety of commodities. 32% of 334 information requests for articles which were suspected to contain SVHCs were answered by suppliers and a minor number of these answers were of good quality. Only two respondents indicated the presence of SVHCs in their articles. Suppliers are not legally obliged to respond to requests if their articles are free of SVHCs. Therefore, the absence of a response might be interpreted as an indication that SVHCs are present below 0.1% in the articles in question. However, there are certain doubts that only two out of 334 articles suspected contain SVHCs.

Conclusions

The data question whether the ambitious aims of the SVHC regime can be achieved under the present conditions. Measures are proposed on how to improve implementation of the information requirement and to amend the legal criteria in the upcoming REACH revision.

     

Why do females have so few extra-pair offspring?

It is generally accepted that if a female can improve her offspring’s genetics via extra-pair copulations (EPC), it is by copulating with extra-pair males whose phenotypes are more superior or whose genes are more compatible to hers than those of her bonded male. Here, we present a model that puts together uncertainties about the male genetic quality, a postcopulatory sperm bias in favor of the better or the more compatible genes, and costs that females pay by being choosy about extra-pair male quality. The model’s conclusions challenge traditional views of good genes explanations of EPC. When phenotypes give incomplete information about genotypes, a female choosing a phenotypically superior extra-pair male, may nevertheless find herself trading good genes of a bonded male for poor genes of an extra-pair male. Such “unfortunate sperm replacements” can limit the female involvement in EPC even when EPC are otherwise cost-free. The model also shows that even a female bonded to a phenotypically superior male may benefit by EPC, provided that sperm competition is biased toward sperm with more fit or more compatible genes. Furthermore, if choosiness is sufficiently costly, a female may even do best by copulating with a random extra-pair male.     

Why do female mice mate with multiple males?

Females often show multi-male mating (MMM), but the adaptive functions are unclear. We tested whether female house mice (Mus musculus musculus) show MMM when they can choose their mates without male coercion. We released 32 females into separate enclosures where they could choose to mate with two neighboring males that were restricted to their own territories. We also tested whether females increase MMM when the available males appeared unable to exclude intruders from their territories. To manipulate territorial intrusion, we introduced scent-marked tiles from the neighboring males into males' territories, or we rearranged tiles within males' own territories as a control. Each female was tested in treatment and control conditions and we conducted paternity analyses on the 57 litters produced. We found that 46 % of litters were multiply sired, indicating that multiple paternity is common when females can choose their mates. Intrusion did not increase multiple paternity, though multiple paternity was significantly greater in the first trial when the males were virgins compared to the second trial. Since virgin male mice are highly infanticidal, this finding is consistent with the infanticide avoidance hypothesis. We also found that multiple paternity was higher when competing males showed small differences in their amount of scent marking, suggesting that females reduce MMM when they can detect differences in males' quality. Finally, multiple paternity was associated with increased litter size but only in the intrusion treatment, which suggests that the effect of multiple paternity on offspring number is dependent on male–male interactions.     

Why do male Callosobruchus maculatus beetles inseminate so many sperm?

Male Callosobruchus maculatus F. (Coleoptera: Bruchidae) inseminate more sperm than females can effectively store in their spermathecae. This study examines the adaptive significance of excess sperm transfer by measuring components of male and female reproductive success in response to manipulating the number of sperm inseminated. The number of sperm transferred during copulation was reduced from 56,000 ±4,462 to 8,700±1,194 by sequentially mating males to virgin females. Reducing the number of sperm inseminated by the first male to mate had no effect on the extent of sperm precedence, but reducing the number of sperm inseminated by the second male resulted in a significant reduction in the extent of sperm precedence. When large numbers of sperm are inseminated the remating refractory period of females is increased. These results indicate that males transferring large numbers of sperm during copulation have a two-fold advantage at fertilization; they are more effective at preempting previously stored sperm and they are likely to father more offspring by delaying the time of female remating. The transfer of excess sperm does not appear to serve as nonpromiscuous male mating effort; the number of eggs laid, their fertility and the subsequent survival of zygotes were unaffected by manipulating the number of sperm inseminated. The underlying mechanisms of sperm precedence were also examined. Simple models of sperm displacement failed to accurately predict the patterns of sperm precedence observed in this species. However, the results do not provide conclusive evidence against the models but rather serve to highlight our limited understanding of the movement of sperm within the female's reproductive tract.     

Why do nocturnal orb-web spiders (Araneidae) search for light?

The nocturnal orb-web spider Larinioides sclopetarius lives near water and frequently builds webs on bridges. In Vienna, Austria, this species is particularly abundant along the artificially lit handrails of a footbridge. Fewer individuals placed their webs on structurally identical but unlit handrails of the same footbridge. A census of the potential prey available to the spiders and the actual prey captured in the webs revealed that insect activity was significantly greater and consequently webs captured significantly more prey in the lit habitat compared to the unlit habitat. A laboratory experiment showed that adult female spiders actively choose artificially lit sites for web construction. Furthermore, this behaviour appears to be genetically predetermined rather than learned, as laboratory-reared individuals which had previously never foraged in artificial light exhibited the same preference. This orb-web spider seems to have evolved a foraging behaviour that exploits the attraction of insects to artificial lights. Received: 8 June 1998 / Received in revised form: 18 January 1999 / Accepted: 19 January 1999     

Why is the coral-reef fish Valenciennea strigata (Gobiidae) monogamous?

Parental-investment theory predicts that monogamy should be rare, and empirical evidence supports this prediction. Monogamy has generally been explained by either (1) a need for biparental care, or (2) a uniform distribution of limiting resources. By contrast, monogamy has evolved in several coral-reef fishes without biparental care, and many coral-reef fishes may not be limited by resources. Monogamy in these fishes might be explained by either (3) a low population density that favors mate fidelity, or (4) an abundance of resources that allows all males to breed and thus lowers the cost-benefit ratio for females that mate guard. We tested predictions of these hypotheses in the coral-reef fish, Valenciennea strigata, except that (1) biparental care had previously been rejected. We found no evidence of resource limitation (2a): food densities within territories did not differ from the density of food in unused habitat adjacent to territories; potential nest sites also existed in this unused habitat. Similarly, (2b) if resources limit the population, territory defense might require the coordinated efforts of a pair; however, widowed fish maintained their territories and (3) remated rapidly. Finally, (4) all males maintained a nest, both sexes enforced monogamy by mate guarding, and females benefited from guarding a high-quality (large) mate. Females mated to large males fed more than when they paired with small males. The results support the hypothesis that abundant and uniformly distributed resources can lead to monogamy by lowering the cost of guarding a mate when mate guarding provides benefits. Received: 25 November 1997 / Accepted after revision: 29 December 1997     

Understanding and predicting ecological dynamics: are major surprises inevitable?

Ecological surprises, substantial and unanticipated changes in the abundance of one or more species that result from previously unsuspected processes, are a common outcome of both experiments and observations in community and population ecology. Here, we give examples of such surprises along with the results of a survey of well-established field ecologists, most of whom have encountered one or more surprises over the course of their careers. Truly surprising results are common enough to require their consideration in any reasonable effort to characterize nature and manage natural resources. We classify surprises as dynamic-, pattern-, or intervention-based, and we speculate on the common processes that cause ecological systems to so often surprise us. A long-standing and still growing concern in the ecological literature is how best to make predictions of future population and community dynamics. Although most work on this subject involves statistical aspects of data analysis and modeling, the frequency and nature of ecological surprises imply that uncertainty cannot be easily tamed through improved analytical procedures, and that prudent management of both exploited and conserved communities will require precautionary and adaptive management approaches.     

When are no-take zones an economically optimal fishery management strategy?

Discussions on the use of marine reserves (no-take zones) and, more generally, spatial management of fisheries are, for the most part, devoid of analyses that consider the ecological and economic effects simultaneously. To fill this gap, we develop a two-patch ecological-economic model to investigate the effects of spatial management on fishery profits. Because the fishery effects of spatial management depend critically on the nature of the ecological connectivity, our model includes both juvenile and adult movement, with density dependence in settlement differentiating the two types of dispersal. Rather than imposing a reserve on our system and measuring its effect on profits, we ask: "When does setting catch levels to maximize system-wide profits imply that a reserve should be created?" Closing areas to fishing is an economically optimal solution when the value derived from spillover from the reserve outweighs the value of fishing in the patch. The condition, while simple to state in summary form, is complex to interpret because it depends on the settlement success of the dispersing organisms, the nature of the costs of the fishing, the economic and ecological heterogeneity of the system, the discount rate, and growth characteristics of the fish population. The condition is more likely to be satisfied when the closed area is a net exporter of biomass and has higher costs of fishing, and for fish populations with density-independent settlement ("adult movement") than with density-dependent settlement ("larval dispersal"). Rather surprisingly, there are circumstances whereby closing low biological productivity areas, and even sometimes low cost areas to fish, can result in greater fishing profits than when both areas are open to fishing.     

Antipredator strategies of house finches: are urban habitats safe spots from predators even when humans are around?

Urbanization decreases species diversity, but it increases the abundance of certain species with high tolerance to human activities. The safe-habitat hypothesis explains this pattern through a decrease in the abundance of native predators, which reduces predation risk in urban habitats. However, this hypothesis does not consider the potential negative effects of human-associated disturbance (e.g., pedestrians, dogs, cats). Our goal was to assess the degree of perceived predation risk in house finches (Carpodacus mexicanus) through field studies and semi-natural experiments in areas with different levels of urbanization using multiple indicators of risk (flock size, flight initiation distance, vigilance, and foraging behavior). Field studies showed that house finches in more urbanized habitats had a greater tendency to flock with an increase in population density and flushed at larger distances than in less urbanized habitats. In the semi-natural experiment, we found that individuals spent a greater proportion of time in the refuge patch and increased the instantaneous pecking rate in the more urbanized habitat with pedestrians probably to compensate for the lower amount of foraging time. Vigilance parameters were influenced in different ways depending on habitat type and distance to flock mates. Our results suggest that house finches may perceive highly urbanized habitats as more dangerous, despite the lower number of native predators. This could be due to the presence of human activities, which could increase risk or modify the ability to detect predators. House finches seem to adapt to the urban environment through different behavioral strategies that minimize risk.     

Why is dominance hierarchy age-related in social insects? The relative longevity hypothesis

In temperate regions, older eusocial hymenopteran females with annual life cycles (annual-temperate) tend to dominate younger females, a behavior demonstrated by many Polistes. However, in queenless ponerine ants and primitively eusocial tropical wasps (perennial/tropical), a younger female can be dominant and occasionally takes over from the older, most dominant reproductive female, the alpha. We investigated these patterns using an inclusive fitness model. The most important difference between the above two cases lay in the length of individual life compared with colony life. Colonies dissolve before winter in the annual-temperate case, so the expected future tenure of the replacement alpha is never longer than that of the original alpha. This makes the non-reproductive subordinate tactic more advantageous for individuals that emerge later in the season because of the fitness cost of superseding. The perennial/tropical case does not have a clear upper limit for colony longevity, so the model predicts that late-born younger daughters are more likely to challenge their mother-alpha because of the expected long future tenure of the new alpha compared with the small indirect cost of the mothers reproductive failure. To switch tactics from being a subordinate to being the new alpha is only optimal in some situations. The high mortality rate of subordinate workers does not qualitatively alter the above pattern. The specific sexual production schedule of the colony sometimes affects the optimal behavior of females in the perennial/tropical case, and older individuals can dominate younger ones when the end of current round of sexual production is imminent.Electronic Supplementary Material Supplementary material is available for this article at     

Why are larvae of the social parasite wasp <Emphasis Type="Italic">Polistes sulcifer</Emphasis> not removed from the host nest?

A challenge for parasites is how to evade the sophisticated detection and rejection abilities of potential hosts. Many studies have shown how insect social parasites overcome host recognition systems and successfully enter host colonies. However, once a social parasite has successfully usurped an alien nest, its brood still face the challenge of avoiding host recognition. How immature stages of parasites fool the hosts has been little studied in social insects, though this has been deeply investigated in birds. We look at how larvae of the paper wasp obligate social parasite Polistes sulcifer fool their hosts. We focus on cuticular hydrocarbons (CHCs), which are keys for adult recognition, and use behavioral recognition assays. Parasite larvae might camouflage themselves either by underproducing CHCs (odorless hypothesis) or by acquiring a chemical profile that matches that of their hosts. GC/MS analyses show that parasite larvae do not have lower levels of CHCs and that their CHCs profile is similar to the host larval profile but shows a reduced colony specificity. Behavioral tests show that the hosts discriminate against alien conspecific larvae from different colonies but are more tolerant towards parasite larvae. Our results demonstrate that parasite larvae have evolved a host larval profile, which overcomes the host colony recognition system probably because of the lower proportion of branched compounds compared to host larvae. In some ways, this is a similar hypothesis to the odorless hypothesis, but it assumes that the parasite larvae are covered by a chemical blend that is not meaningful to the host.     

Whirligig beetle aggregations: what are the costs and the benefits?

Laboratory experiments investigated the benefits and costs of aggregation formation in the whirligig beetles, Gyrinus marinus and G. substriatus (Coleoptera: Gyrinidae). Different sized groups of whirligigs were exposed to fish predators, and capture rate per group and per individual were estimated. Attack rate per group increased with group size, suggesting that these aggregations behave as selfish herds. In another series of experiments in which whirligigs were exposed to top- and side-attacking predators, large groups were found to detect predators when they were further away than small groups. Video analysis of these groups showed that beetles tended to increase their swimming speed after exposure to a predator but did not show an increased tendency to circle. In natural aggregations, filmed in the field, swimming speed was found to be related to group size. Body condition of field collected beetles, as estimated from the regression of body mass on body length, was not found to be related to group size, implying that large groups do not suffer reduced condition. Received: 31 January 1997 / Accepted after revision: 24 November 1997     

Consumer resource matching in urbanizing landscapes: are synanthropic species over-matching?

Population responses of synanthropic species to urbanization may be explained by the resource-matching rule, which postulates that individuals should distribute themselves according to resource availability. According to the resource-matching rule, urban habitats will contain greater densities if they provide better resources than rural habitats. However, because resource availability is density dependent, individuals in urban areas would ultimately achieve fitness levels comparable to, but no better than, individuals in less urban areas. Some ecologists suggest that synanthropic birds may not conform to the resource-matching rule and may instead overmatch (i.e., overexploit) in urban habitats, ultimately leading to lower fitness despite greater resource levels. Using the Northern Cardinal (Cardinalis cardinalis) as a focal species, we evaluated if Cardinal populations in urban and rural habitats were consistent with predictions of consumer resource matching. During 2003-2006 we documented population density, adult body condition, apparent survival, and annual reproductive productivity of Cardinals in riparian forest stands within urban (n = 8 stands) and rural (n = 6 stands) landscapes in Ohio, USA. Density of Cardinals in urban forests was four times that found in more rural forests. Mark-resight data from 147 males and 125 females over four years indicated that apparent survival rates were similar between urban and rural landscapes (phi = 0.64, SE = 0.039 for males and phi = 0.57, SE = 0.04 for females). Similarly, body condition indices of 168 males, 142 females, and 118 nestlings did not differ significantly between landscapes. Annual reproductive productivity (mean number of fledglings per pair over breeding season) of 294 pairs was comparable for urban (2.4 +/- 0.18 [mean +/- SE] and rural (2.1 +/- 0.18) young birds. Thus, contrary to recent suggestions, we find that high densities of certain synanthropic species in urban landscapes are consistent with expectations of consumer resource matching.     

Setbacks to shoot growth are common in woody plants, so how are shoots of some species safer than others?

Tissue turnover is a critical facet of plant life history variation. This study quantifies losses from setbacks to growth of terminal woody shoots 1.2m long, across 83 species and seven sites in eastern Australia. Setbacks, where the leading meristem had been removed or died and a new leader had emerged, were common (median three per shoot). Shoots had lost an average of 0.25 m of lead-stem length for 1.2 m net shoot-length gain. Insects like girdlers and borers were prominent causes of large setbacks. The sites spanned tropical to temperate and humid to semiarid climates, but variation in stem loss was much greater across species than across sites. We measured 17 plant functional traits related to growth form, mechanics, hydraulics, and economics. Only four traits were correlated with variation across species in stem losses: stem diameter, stem nitrogen content, bark thickness, and maximum photosynthetic rate. The correlations were weak. Stem specific gravity (wood density) showed no correlation with risk. Our results suggest a pattern similar to the growth risk trade-off known for herbaceous plants, where traits associated with fast growth increase tissue turnover and herbivory, but the weak correlations leave ample scope for other influences that remain to be identified.     

Why do most tropical animals reproduce seasonally? Testing hypotheses on an Australian snake

Most species reproduce seasonally, even in the tropics where activity occurs year-round. Squamate reptiles provide ideal model organisms to clarify the ultimate (adaptive) reasons for the restriction of reproduction to specific times of year. Females of almost all temperate-zone reptile species produce their eggs or offspring in the warmest time of the year, thereby synchronizing embryogenesis with high ambient temperatures. However, although tropical reptiles are freed from this thermal constraint, most do not reproduce year-round. Seasonal reproduction in tropical reptiles might be driven by biotic factors (e.g., peak periods of predation on eggs or hatchlings, or food for hatchlings) or abiotic factors (e.g., seasonal availability of suitably moist incubation conditions). Keelback snakes (Tropidonophis mairii, Colubridae) in tropical Australia reproduce from April to November, but with a major peak in May-June. Our field studies falsify hypotheses that invoke biotic factors as explanations for this pattern: the timing of nesting does not minimize predation on eggs, nor maximize food availability or survival rates for hatchlings. Instead, our data implicate abiotic factors: female keelbacks nest most intensely soon after the cessation of monsoonal rains when soils are moist enough to sustain optimal embryogenesis (wetter nests produce larger hatchlings, that are more likely to survive) but are unlikely to become waterlogged (which is lethal to eggs). Thus, abiotic factors may favor seasonal reproduction in tropical as well as temperate-zone animals.     

Avian predators are less abundant during periodical cicada emergences, but why?

Despite a substantial resource pulse, numerous avian insectivores known to depredate periodical cicadas (Magicicada spp.) are detected less commonly during emergence years than in either the previous or following years. We used data on periodical cicada calls collected by volunteers conducting North American Breeding Bird Surveys within the range of cicada Brood X to test three hypotheses for this observation: lower detection rates could be caused by bird calls being obscured by cicada calls ("detectability" hypothesis), by birds avoiding areas with cicadas ("repel" hypothesis), or because bird abundances are generally lower during emergence years for some reason unrelated to the current emergence event ("true decline" hypothesis). We tested these hypotheses by comparing bird detections at stations coincident with calling cicadas vs. those without calling cicadas in the year prior to and during cicada emergences. At four distinct levels (stop, route, range, and season), parallel declines of birds in groups exposed and not exposed to cicada calls supported the true decline hypothesis. We discuss several potential mechanisms for this pattern, including the possibility that it is a consequence of the ecological and evolutionary interactions between predators of this extraordinary group of insects.     

Surplus territory contenders in male red-winged blackbirds: where are the desperados?

Summary We tested whether payoff asymmetries could explain why floater red-winged blackbirds seldom escalate contests to acquire territories. We removed territorial male \ldowners\rd and allowed floater replacements to claim territories. We then released owners to see how three currencies of payoff asymmetries (replacement occupancy) owners' likelihood and speed of reclaiming their territories (owner success). Neither the duration of an owner's internment nor the amount of time that a floater replacement had held the territory affected owner success (Figs. 1 and 2). Owner success was also not affected by the number of neighbors that they had (Fig. 3). Finally, owner success was the same irrespective of whether or not he was the likely sire of offspring on the territory, or the size of his harem (Fig. 4). Although these results are consistent with the proposal that payoff asymmetries will be irrelevant in contests for valuable resources, they are inconsistent with the proposal's corollary that excluded individuals should become \lddesperados\rd and escalate or even fight to the death in contests for those resources. Expected payoffs for passive acquisition of territories in this species may be higher than from a more aggressive desperado strategy.Correspondence to: P.J. Weatherhead