The relative influence of habitat loss and fragmentation: do tropical mammals meet the temperate paradigm?

The relative influence of habitat loss vs. habitat fragmentation per se (the breaking apart of habitat) on species distribution and abundance is a topic of debate. Although some theoretical studies predict a strong negative effect of fragmentation, consensus from empirical studies is that habitat fragmentation has weak effects compared with habitat loss and that these effects are as likely to be positive as negative. However, few empirical investigations of this issue have been conducted on tropical or wide-ranging species that may be strongly influenced by changes in patch size and edge that occur with increasing fragmentation. We tested the relative influence of habitat loss and fragmentation by examining occupancy of forest patches by 20 mid- and large-sized Neotropical mammal species in a fragmented landscape of northern Guatemala. We related patch occupancy of mammals to measures of habitat loss and fragmentation and compared the influence of these two factors while controlling for patch-level variables. Species responded strongly to both fragmentation and loss, and response to fragmentation generally was negative. Our findings support previous assumptions that conservation of large mammals in the tropics will require conservation strategies that go beyond prevention of habitat loss to also consider forest cohesion or other aspects of landscape configuration.     

The pursuit of extra-pair copulations by monogamous female razorbills: how do females benefit?

Summary Monogamous female razorbills Alca torda actively sought extra-pair copulations (EPCs) in mating arenas outside their nesting colony. Females showed marked variation in promiscuity, defined as the number of EPCs they accepted (0–7), and in receptivity, defined as the percentage of EPC opportunities accepted (0%–100%). The opportunity of females to encounter males for EPCs was measured by (a) the time spent in the mating arena, and (b) the guarding effectiveness of their mates. Females appeared to encounter males and accept EPCs independently of opportunity, which suggests that variation in promiscuity and receptivity is caused by differences in the degree to which individual females might benefit from EPCs. I tested predictions of four hypotheses which propose benefits which females could obtain from EPCs: good genes, genetic diversity of offspring, insurance against infertility of mate, and acquisition of a new mate. No evidence was found to support two predictions made exclusively by the good genes hypothesis: (a) paired males did not achieve greater EPC success than unpaired males, and (b) males who were cuckolded did not obtain fewer EPCs than males whose mates avoided EPCs, suggesting that a male's attractiveness to other females does not ensure his own mate's fidelity. A prediction of both the insurance and good genes hypotheses was supported: females (who in this species retain their mates between years) showed similar receptivity to EPCs between years. This finding contradicts the genetic diversity hypothesis which predicts that in species that lay one egg per year, females who refuse EPCs in one year will accept them in the next. The good genes and genetic diversity hypotheses both predict that females who accept EPCs will behave as if to maximize their chances of being fertilized by extra-pair males. Results contradicted this prediction: females who accepted EPCs were more, rather than less receptive to their mates' copulation attempts than were females who did not accept EPCs. This finding is compatible with the insurance hypothesis which only requires females to store the sperm of extra-pair males, without necessarily being fertilized by it. Insurance against male infertility was the only hypothesis whose predictions were not contradicted by any of the results.     

Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship?

The relationship between plant diversity and productivity has largely been attributed to niche complementarity, assuming that plant species are complementary in their resource use. In this context, we conducted an 15N field study in three different grasslands, testing complementarity nitrogen (N) uptake patterns in terms of space, time, and chemical form as well as N strategies such as soil N use, symbiotic N fixation, or internal N recycling for different plant species. The relative contribution of different spatial, temporal, and chemical soil N pools to total soil N uptake of plants varied significantly among the investigated plant species, within and across functional groups. This suggests that plants occupy distinct niches with respect to their relative N uptake. However, when the absolute N uptake from the different soil N pools was analyzed, no spatial, temporal, or chemical variability was detected, but plants, and in particular functional groups, differed significantly with respect to their total soil N uptake irrespective of treatment. Consequently, our data suggest that absolute N exploitation on the ecosystem level is determined by species or functional group identity and thus by community composition rather than by complementary biodiversity effects. Across functional groups, total N uptake from the soil was negatively correlated with leaf N concentrations, suggesting that these functional groups follow different N use strategies to meet their N demands. While our findings give no evidence for a biodiversity effect on the quantitative exploitation of different soil N pools, there is evidence for different and complementary N strategies and thus a potentially beneficial effect of functional group diversity on ecosystem functioning.     

The relationship between maternal phenotype and offspring quality: do older mothers really produce the best offspring?

Maternal effects are increasingly recognized as important drivers of population dynamics and determinants of evolutionary trajectories. Recently, there has been a proliferation of studies finding or citing a positive relationship between maternal size/age and offspring size or offspring quality. The relationship between maternal phenotype and offspring size is intriguing in that it is unclear why young mothers should produce offspring of inferior quality or fitness. Here we evaluate the underlying evolutionary pressures that may lead to a maternal size/age-offspring size correlation and consider the likelihood that such a correlation results in a positive relationship between the age or size of mothers and the fitness of their offspring. We find that, while there are a number of reasons why selection may favor the production of larger offspring by larger mothers, this change in size is more likely due to associated changes in the maternal phenotype that affect the offspring size-performance relationship. We did not find evidence that the offspring of older females should have intrinsically higher fitness. When we explored this issue theoretically, the only instance in which smaller mothers produce suboptimal offspring sizes is when a (largely unsupported) constraint on maximum offspring size is introduced into the model. It is clear that larger offspring fare better than smaller offspring when reared in the same environment, but this misses a critical point: different environments elicit selection for different optimal sizes of young. We suggest that caution should be exercised when interpreting the outcome of offspring-size experiments when offspring from different mothers are reared in a common environment, because this approach may remove the source of selection (e.g., reproducing in different context) that induced a shift in offspring size in the first place. It has been suggested that fish stocks should be managed to preserve these older age classes because larger mothers produce offspring with a greater chance of survival and subsequent recruitment. Overall, we suggest that, while there are clear and compelling reasons for preserving older females in exploited populations, there is little theoretical justification or evidence that older mothers produce offspring with higher per capita fitness than do younger mothers.     

Diversity and host range of foliar fungal endophytes: are tropical leaves biodiversity hotspots?

Fungal endophytes are found in asymptomatic photosynthetic tissues of all major lineages of land plants. The ubiquity of these cryptic symbionts is clear, but the scale of their diversity, host range, and geographic distributions are unknown. To explore the putative hyperdiversity of tropical leaf endophytes, we compared endophyte communities along a broad latitudinal gradient from the Canadian arctic to the lowland tropical forest of central Panama. Here, we use molecular sequence data from 1403 endophyte strains to show that endophytes increase in incidence, diversity, and host breadth from arctic to tropical sites. Endophyte communities from higher latitudes are characterized by relatively few species from many different classes of Ascomycota, whereas tropical endophyte assemblages are dominated by a small number of classes with a very large number of endophytic species. The most easily cultivated endophytes from tropical plants have wide host ranges, but communities are dominated by a large number of rare species whose host range is unclear. Even when only the most easily cultured species are considered, leaves of tropical trees represent hotspots of fungal species diversity, containing numerous species not yet recovered from other biomes. The challenge remains to recover and identify those elusive and rarely cultured taxa with narrower host ranges, and to elucidate the ecological roles of these little-known symbionts in tropical forests.     

The rarity of twins: a result of an evolutionary battle between mothers and daughters—or do they agree?

The aim of this study was to test the hypothesis that parent–offspring conflict over parental investment might have led to the rarity of dizygotic twins in humans. We explored the theoretical prediction that twins maximize their inclusive fitness by the death of a co-twin, while mothers of twins maximize fitness by raising both twins to independence. We used life history data (1700–1900) from two parishes in Central Norway to compare differences in inclusive fitness (measured as number of children reared to the age of 16 years, using Hamilton's rule) between twins and mothers of twins. Our results show that twins maximize their inclusive fitness by the death of a co-twin, while mothers of twins raise more children by rearing both twins to adulthood. However, because twins growing up as singletons may produce higher or at least equal number of offspring than the sum of the two twins growing up together, mothers might gain more grandchildren by allowing twins to grow up as singletons. To conclude, both selfish twins and their mothers might benefit by the death of a co-twin, indicating that there is no parent–offspring conflict responsible for the rareness of twins in these human populations. Finally, we discuss the results in the light of “The Insurance Egg Hypothesis” and “The Natural Selection Hypothesis”.     

Nestmate recognition in the ant Cataglyphis niger : do queens matter?

This study compares two basic models for the origin and maintenance of colony gestalt odor in the polygynous ant species Cataglyphis niger. In the first model, queens are centers of de novo biosynthesis and distribution of recognition odors (“queen-centered” model); in the second, colony odors are primarily synthesized and distributed by workers (“worker-centered” model). We tested the behavioral patterns that are predicted from each model, and verified by biochemical means the distributional directionality of these signals. Encounters between nestmates originating from split colonies were as amicable as between nestmates from non-split colonies; queenless ants were as aggressive as their queenright nestmates, and both were equally aggressed by alien ants. These results indicate that queens have little impact on the recognition system of this species, and lend credence to the worker-centered model. The queen-centered model predicts that unique queen substances should be produced in appreciable quantities and that, in this respect, queens should be more metabolically active than workers. Analysis of the chemical composition of postpharyngeal glands (PPGs) or cuticular extracts of queens and workers revealed high similarity. Quantitatively, queens possessed significantly greater amounts of hydrocarbons in the PPG than workers, but the amount on the thoracic epicuticle was the same. Queens, however, possess a lower hydrocarbon biosynthesis capability than workers. The biochemical evidence thus refutes the queen-centered model and supports a worker-centered model. To elucidate the directionality of cue distribution, we investigated exchange of hydrocarbons between the castes in dyadic or group encounters in which selective participants were prelabeled. Queens tended to receive more and give less PPG content, whereas transfer to the epicuticle was low and similar in all encounters, as predicted from the worker-centered hypothesis. In the group encounters, workers transferred, in most cases, more hydrocarbons to the queen than to a worker. This slight preference for the queen is presumably amplified in a whole colony and can explain their copious PPG content. We hypothesize that preferential transfer to the queen may reflect selection to maintain her individual odor as close to the average colony odor as possible. Received: 4 November 1997 / Accepted after revision: 5 February 1998     

Navigational strategies of homing pigeons at familiar sites: do landmarks reduce the deflections induced by clock-shifting?

To analyze the navigational strategy of homing pigeons at familiar sites in view of a possible role of local landmarks, two groups of pigeons—one familiar to the release site, the other unfamiliar—were released with their internal clock shifted 6 h fast, with untreated birds of both groups serving as controls. The two groups showed median deflections of 67% and 57%, respectively, of the expected size, with no consistent difference in the size of the deflection between familiar and unfamiliar birds. This clearly shows that familiarity with the release site and with the local landscape features does not affect the size of the deflections induced by clock-shifting. Obviously, pigeons familiar with the release site do not change their navigational strategy, but still continue to determine their home course solely as a compass course. General problems with orientation by landmarks are discussed; however, landmarks may help birds to recognize a site and recall the respective course.

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Darwin's special difficulty: the evolution of “neuter insects” and current theory

In the Origin of Species, Darwin discussed several challenges that worker insects presented to his theory of natural selection. Complex instincts such as building of combs of hexagonal cells were one problem and were explained by showing plausible intermediate stages. A more serious challenge was posed by the multiple worker castes seen in many ants. How could sterile individuals continue to evolve? A careful reading of the Origin suggests that Darwin was not primarily concerned by the evolution of worker sterility itself, which he considered a minor difficulty. Some modern commentaries on Darwin and insect workers seem to be cases of present interests interfering with the interpretation of the past. From a modern perspective, the evolution of a worker caste, and its corollary altruism, are evolutionary puzzles inasmuch as natural selection normally favors greater, not lesser, individual reproduction. These puzzles were resolved by Hamilton's theory of inclusive fitness. We now have a good functional understanding of how natural selection can cause both the origin of workers and their elaboration into greater levels of sterility and multiple morphological castes. Mechanistic understanding of morphological castes is also increasing via research into alternative developmental pathways. When the Origin was written, genetics did not exist and it would have been virtually impossible for Darwin to elaborate such ideas. However, the Origin probably addressed the main questions in the minds of Victorian readers in relation to insect workers. Darwin was prescient in having insights with close relationships to modern-day interests and the key principles involved, including kinship and benefits to the colony, even if these are not exact precursors to modern thinking.     

Timber Certification: Where Is the Voice of the Biologist?

Editor's note: This Conservation Forum begins with a reprinted editorial by Elizabeth Bennett from the August 2000 issue of Conservation Biology . Several papers were submitted in repsonse to that piece, and it is reprinted here to start this forum.     

Interacting effects of habitat destruction and changing disturbance rates on biodiversity: Who is going to survive?

Changes in disturbance rates due to climate change may increase or decrease diversity, whereas permanent loss of habitat is generally believed to decrease diversity. It is, however, very likely that the effects of disturbances and habitat destruction interact. Understanding such combined effects is essential to predict the response of communities to global changes and in particular which functional types of species are most endangered. Using an individual-based spatially explicit community model, we investigate (1) whether diversity-disturbance curves alter when spatially uncorrelated or autocorrelated habitat destruction is added, and (2) which functional types of species are able to survive under these altered conditions. Model communities consisted of four functional types of species trading off between colonisation ability and competition strength. We found that habitat destruction may alter both height and shape of diversity-disturbance curves: maximum diversity at intermediate disturbance rates may shift to other disturbance rates or even split into two peaks giving rise to bimodal diversity-disturbance relationships with different sub-communities persisting at low and high disturbance rates. Diversity responded differentially depending on how the colonisation-competition trade-off was represented. Our results suggest that, for trade-offs in seed production rate, generally the best coloniser will better withstand the interacting effects of habitat destruction and changing disturbance rates; however, for trade-offs in mean dispersal distances, functional types characterized by intermediate abilities will perform best. We conclude that predictions of the impacts of changing disturbance rates on biodiversity depend on community structure and cannot be made without knowledge of concurrent permanent habitat destruction.     

How do simple energy activities comprise complex behavior of life systems?: A conceptual synthesis and decomposition of the energy structure of life systems

Reduction and synthesis has always been Science’s way to learn about complex systems. The problem is, of course, that after studying the behavior of the system components, the reconstruction of system behavior cannot be any simple summation of the behavior of the individual components. How should we go about with the reconstruction is an umbrella question addressed in the paper. In particular, one should ask what is the contribution of each component to the system, and what kind and quantitative relations do the connections among the components signify in the total system? Answering these questions is necessary to understand the internal mechanism of a life system, the regularity of its dynamics, and the nature of the control mechanisms.The general approach of the paper is based on the assumption that the life energy transmission structure of the system is the fundamental principle of reduction and synthesis.The paper generalizes the energy transmission in terms of two basic types, based upon the mode of linkage in energy activities, such as sequential or parallel. The basic mathematical concept is an eigenparameter analysis in a model called Life Energy System Model (LESM). Simple examples illustrate the models application. It is true that the energy transmission structure of a natural life system is usually very complex, in which case the analysis has to incorporate more details than shown in the paper. Yet, the analysis still follows the same fundamental principle as outlined.     

Visions for the future of transport: Approaching consensus?

Many cities and national authorities have made substantial progress in developing transport strategies to reconcile environmental pressures with the increasing demand for travel. However, there is still some doubt as to what are the key ingredients of a successful strategy and progress in implementing such changes has been slow. Research is needed to better understand what are the essential ingredients to a balanced transport policy, the constraints, which are preventing these from being implemented, and the acceptability of such changes. This paper reports on the outcomes of a project funded under the Rees Jeffreys Road Fund research programme 'From Realism to Reality', which aims to identify the elements necessary for a successful transport policy in the UK into the 21st century, The project seeks to develop a vision for the transport system of the future through an understanding of the extent of the current problems in the transport sector and an examination of the range and effectiveness of options and instruments available to resolve them. The paper examines a range of visions that are currently in the public domain. Results show that there is a degree of consistency in the general aims and objectives of such visions, but that there is rather less consensus in the means proposed to achieve those aims.     

Evaluating animal personalities: do observer assessments and experimental tests measure the same thing?

The animal personality literature uses three approaches to assess personality. However, two of these methods, personality ratings and experimentation, have been little compared in captivity and never compared in the wild. We assessed the boldness of wild chacma baboons Papio ursinus using both ratings and experimental methods. Boldness was experimentally assessed when individuals were presented with a novel food item during natural foraging. The boldness of the same individuals was rated on a five-point scale by experienced observers. The ratings and experimental assessments of boldness were found to correlate positively and in a linear fashion. When considered categorically the two approaches showed variable agreement depending on the number of categories assigned and the cut-off criteria adopted. We suggest that the variation between approaches arises because each method captures different aspects of personality; ratings consider personality in absolute terms (using predefined criteria) and multiple contexts, while experimental assessments consider personality in relative terms (using experimental scores relative to the population average) and in limited contexts. We encourage animal personality researchers to consider adopting both methodologies in future studies. We also propose that future studies restrict their analyses to continuous data, since the greatest comparability between methods was found with these data. However, if individuals must be categorised, we suggest that researchers either (a) analyse only those individuals categorised as bold or shy by both ratings and experimental approaches or, if these methods cannot be employed simultaneously, (b) do not use approach-specific criteria but choose a cut-off that can be compared by both approaches.     

The role of water in the development of The Netherlands— a historical perspective

In this paper an overview is presented of the landscape development of the low-lying parts of The Netherlands. It is shown that water has played an important role in this development. Since prehistoric times, but especially since about 800 AD, man has gradually occupied the low lands along the sea and the main rivers. Through these settlements a chain of actions and reactions of man and his physical environment was set into motion. This chain of events is briefly outlined. It is stated and also illustrated that not only the characteristics of the landscape of the western part of The Netherlands, but also the organization of the Dutch society has been influenced strongly by manÕs need to cope with the omnipresence of water and the threats and opportunities posed by that. It is concluded that the outlined events can only be understood properly if interdisciplinary research takes a more prominent role than it does now.     

How do nutrient conditions and species identity influence the impact of mesograzers in eelgrass-epiphyte systems?

Coastal eutrophication is thought to cause excessive growth of epiphytes in eelgrass beds, threatening the health and survival of these ecologically and economically valuable ecosystems worldwide. Mesograzers, small crustacean and gastropod grazers, have the potential to prevent seagrass loss by grazing preferentially and efficiently on epiphytes. We tested the impact of three mesograzers on epiphyte biomass and eelgrass productivity under threefold enriched nutrient concentrations in experimental indoor mesocosm systems under summer conditions. We compared the results with earlier identical experiments that were performed under ambient nutrient supply. The isopod Idotea baltica, the periwinkle Littorina littorea, and the small gastropod Rissoa membranacea significantly reduced epiphyte load under high nutrient supply with Rissoa being the most efficient grazer, but only high densities of Littorina and Rissoa had a significant positive effect on eelgrass productivity. Although all mesograzers increased epiphyte ingestion with higher nutrient load, most likely as a functional response to the quantitatively and qualitatively better food supply, the promotion of eelgrass growth by Idotea and Rissoa was diminished compared to the study performed under ambient nutrient supply. Littorina maintained the level of its positive impact on eelgrass productivity regardless of nutrient concentrations.