Assessing natural resource use by forest-reliant communities in Madagascar using functional diversity and functional redundancy metrics

Biodiversity plays an integral role in the livelihoods of subsistence-based forest-dwelling communities and as a consequence it is increasingly important to develop quantitative approaches that capture not only changes in taxonomic diversity, but also variation in natural resources and provisioning services. We apply a functional diversity metric originally developed for addressing questions in community ecology to assess utilitarian diversity of 56 forest plots in Madagascar. The use categories for utilitarian plants were determined using expert knowledge and household questionnaires. We used a null model approach to examine the utilitarian (functional) diversity and utilitarian redundancy present within ecological communities. Additionally, variables that might influence fluctuations in utilitarian diversity and redundancy--specifically number of felled trees, number of trails, basal area, canopy height, elevation, distance from village--were analyzed using Generalized Linear Models (GLMs). Eighteen of the 56 plots showed utilitarian diversity values significantly higher than expected. This result indicates that these habitats exhibited a low degree of utilitarian redundancy and were therefore comprised of plants with relatively distinct utilitarian properties. One implication of this finding is that minor losses in species richness may result in reductions in utilitarian diversity and redundancy, which may limit local residents' ability to switch between alternative choices. The GLM analysis showed that the most predictive model included basal area, canopy height and distance from village, which suggests that variation in utilitarian redundancy may be a result of local residents harvesting resources from the protected area. Our approach permits an assessment of the diversity of provisioning services available to local communities, offering unique insights that would not be possible using traditional taxonomic diversity measures. These analyses introduce another tool available to conservation biologists for assessing how future losses in biodiversity will lead to a reduction in natural resources and provisioning services from forests.     

Different competitive potential in two coexisting mouse lemur species in northwestern Madagascar

Interspecific competition has been suggested to influence the biogeographic distribution patterns of species. A high competitive potential could entail species-specific advantages during resource acquisition that could translate into a higher potential for range expansion. We investigated whether differences in the competitive potential of the morphologically similar and partially sympatric gray mouse lemur (Microcebus murinus) and golden-brown mouse lemur (Microcebus ravelobensis) may help to explain differences in their geographic range sizes. We carried out encounter experiments with 14 pairs of captured female mouse lemurs of both species. The experimental dyads were tested in a two-cage arrangement, with individuals being separated from each other outside the experiments. Two days of habituation and four subsequent days of 1-h encounter experiments were conducted, before releasing the animals again in the wild. In general, the M. murinus individuals won significantly more conflicts than their partners. In eight of 14 tested pairs, there was a significant species bias in winning conflicts, and in 87.5% of these dyads, M. murinus was the "dyad winner". A high competitive potential did not depend on body mass. Furthermore, "dyad winners" spent more time feeding (P < 0.05) and were less spatially restricted than "dyad losers". To conclude, our results suggest that the widely distributed M. murinus may indeed have a higher competitive potential than the regional endemic M. ravelobensis, which may, among other possible factors, have enabled this species to expand geographically, despite the presence of other competing congeners.     

Seasonal changes of functional groups in coleopteran communities in pine forests

Fauna assemblages reflect their habitat relating to ecological function in an ecosystem. The functional groups are concerned with how a resource is processed by different species to provide a specific ecosystem service or function. We elucidated seasonal changes of coleopteran functional groups in forests, and evaluated their ecological roles related to available food resources. Coleopteran communities were collected weekly or biweekly using Malaise traps at nine study sites in Japanese red pine forests in Korea from late June to September 2005. Compositions of the functional groups were compared at the different sites and at sampling times with respect to taxa richness and abundance. Cluster analysis and non-metric multidimensional scaling were used to characterize spatial and temporal changes of functional groups. Herbivores and dead/live wood feeders regulating primary production in the pine forests were the dominant coleopteran groups in July, followed by detritivores and predators that dominated from July to August, resulting from the accumulation of detritus. Then, fungivores became dominant due to increased fungal biomass in the forest. Seasonal changes of coleopteran functional groups shifted from regulators of primary production to regulators of decomposition, reflecting their available food resources. In addition, abundance of detritivores and predators were dependent on total abundance of coleopterans, suggesting that these two groups reflect their habitat condition.     

Analysis of avian communities in Lake Guri,Venezuela, using multiple assembly rule models

This study analyzed the distribution of resident, forest-interior bird species nesting on islands in Lake Guri, Venezuela using several different community assembly rule models. The models that were tested included Diamond's Assembly Rules, Size Structure, Guild Proportionality, Favored States, and Nestedness. It was determined that the species composition of the study communities was only weakly influenced by competition, but that competition did appear to limit the size similarity which is permissible for co-occurring species. There was no tendency for the relative proportion of species within guilds (i.e. insectivore, omnivore, nectivore and frugivore) to remain stable among the islands. When only the insectivorous and omnivorous species were analyzed (using feeding strata as the functional groups) there was some support for the guild proportionality hypothesis. This study found no support for Fox's Favored State hypothesis, possibly due to the overrepresentation of insectivores and omnivores in the species pool. The island communities exhibit a highly nested structure. This high degree of nestedness supports the hypothesis that the assemblages are more strongly determined by differential extinction vulnerability and selective species loss than by interspecific or inter-guild competition. Understanding patterns of community assembly and their underlying forces has important implications for conservation ecology and reserve design.     

Sociality in the gray mouse lemur (Microcebus murinus) in northwestern Madagascar

The mating system and social organization of the gray mouse lemur (Microcebus murinus) was investigated in two three-month field studies (covering a period before, during, and after the first mating season) in Ampijoroa, northwestern Madagascar. The spatial and temporal distribution of the sexes within a population was studied using mark/recapture techniques and radiotelemetry to assess possible contest or scramble competition between the males. Sociality was inferred from the occurrence and probability of nocturnal social encounters, the temporal stability of daily sleeping groups, and nocturnal ranging patterns of co-sleepers. Males and females were evenly distributed in the study area within a network of highly overlapping home ranges. No indications were found for the spatial monopolization of the females by certain dominant males. Males and females had spatial access to several potential mates; the mating system is therefore characterized as a multi-male/multi-female system. Male home range sizes increased during the first mating season, which was interpreted as an indicator for scramble competition between the males. Competitive mate searching, sperm competition, and temporary mate guarding as well as female mate choice are suggested as the most probable reproductive strategies. Over the course of the study the animals lived continually within the study area, and most females formed stable individualized sleeping associations. Females that slept together shared a higher percentage of their home range than did females that slept at different sites. It is suggested that this network of social relationships should be described as a dispersed but individualized neighborhood.     

The assembly of ecological communities inferred from taxonomic and functional composition

Among-site variation in metacommunities (beta diversity) is typically correlated with the distance separating the sites (spatial lag). This distance decay in similarity pattern has been linked to both niche-based and dispersal-based community assembly hypotheses. Here we show that beta diversity patterns in community composition, when supplemented with functional-trait information, can be used to diagnose assembly processes. First, using simulated data, we show how the relationship between distance decay patterns in taxonomic and functional measures of community composition can be used to predict the influence of a given trait on community assembly. We then use the patterns generated by the simulation as a template to show that the sorting of benthic macroinvertebrate metacommunities in headwater streams is likely influenced by different sets of functional traits at regional and local scales. We suggest that functional-trait databases and spatially referenced taxonomic surveys can be used to predict the spatial scales at which different aspects of interspecific functional variation are involved in niche-based community assembly while accounting for the influence of dispersal-based community assembly processes.     

Global climate cycles and cyclones: consequences for rainfall patterns and lemur reproduction in southeastern Madagascar

Most studies that examine the influence of climatic change on flora and fauna have focused on northern latitudes; however, there is increasing recognition that tropical regions are also being affected. Despite this, regions such as Madagascar, which are rich in endemic biodiversity but may have low adaptive capacity to climatic change, are poorly represented in studies examining the effects of climate variability on biota. We investigated how El Niño Southern Oscillations (ENSO) influence precipitation patterns in the rainforest region of southeastern Madagascar (1962–2006) and then constructed models to assess the potential contribution of climatic variables on the reproductive parameters of the Milne Edward's sifaka, a threatened lemur species (Propithecus edwardsi), over a 20‐year period. The Southern Oscillation Index of sea surface temperature (SST) anomalies in the tropical Pacific was associated with precipitation patterns including wetter wet seasons during warmer phases and drier dry seasons following cooler phases. The best‐supported models of lemur fecundity (female offspring per female that survive to 1 year of age per year) included cyclone presence during gestation and ENSO phase before conception and during the first 6 months of life. Models also suggested that heavy rains during gestation may limit birth rates and that prolonged drought during female lactation may limit first year offspring survival; although these variables were given little importance for predicting overall fecundity relative to ENSO phases and cyclone presence. Our results linking lemur reproduction with climatic variability suggest that climatic changes may be an additional threat to Madagascar's unique and already endangered flora and fauna. The association between precipitation in southeastern Madagascar and SST anomalies in the tropical Pacific suggests that dynamics of wildlife populations even in tropical areas such as Madagascar can be affected by global climate cycles making them potentially vulnerable to global climate change.     

The evolution of primate communities and societies in Madagascar

During their 120 to 165 million years of isolation, the flora and fauna of Madagascar evolved, to a large extent, independently of the African mainland.¹ In contrast to other oceanic islands, Madagascar is large enough to house the major components of tropical ecosystems, allowing tests of evolutionary hypotheses on the level of complete communities. Taking lemurs, the primates of Madagascar, as an example, evolutionary hypotheses correctly predict the organization of their community structure with respect to ecological correlates. Lemur social systems and their morphological correlates, on the other hand, deviate in some respects from those of other primates. Apparently, lemur social systems are influenced by several selection pressures that are weak or rare in other primates. These include variable activity patterns and avoidance of infanticide. The interspecific variation in lemur social systems therefore offers a unique opportunity for a comprehensive study of the determinants of primate social systems.     

Light interception in species with different functional groups coexisting in moorland plant communities

Competition for light is one of the most essential mechanisms affecting species composition. It has been suggested that similar light acquisition efficiency (Φ_mass, absorbed photon flux per unit aboveground mass) may contribute to species coexistence in multi-species communities. On the other hand, it is known that traits related with light acquisition vary among functional groups. We studied whether Φ_mass was similar among species with different functional groups coexisting in moorland communities. We conducted stratified clipping in midsummer when the stand biomass reached a maximum. Light partitioning among species was estimated using a model accounting for both direct and diffuse light. Evergreen species were found to have a significantly lower Φ_mass than deciduous species, which resulted from their lower absorbed photon flux per unit leaf area and lower specific leaf area. Shrubs had a smaller leaf mass fraction, but their Φ_mass was not lower than that of herbs because they had a higher leaf position due to the presence of wintering stems. Species with vertical leaves had a higher Φ_mass than those with horizontal leaves despite vertical leaves being a decided disadvantage in terms of light absorption. This higher Φ_mass was achieved by a greater leaf height in species with vertical leaves. Our results clearly demonstrate that light acquisition efficiency was different among the functional groups. However, the trend observed is not necessarily the same as that expected based on prior knowledge, suggesting that disadvantages in some traits for light acquisition efficiency are partly compensated for by other traits.     

N-Glycans mutations rule oligomeric assembly and functional expression of P2X3 receptor for extracellular ATP

N-Glycosylation affects the function of ion channels at the level of multisubunit assembly, protein trafficking, ligand binding and channel opening. Like the majority of membrane proteins, ionotropic P2X receptors for extracellular ATP are glycosylated in their extracellular moiety. Here, we used site-directed mutagenesis to the four predicted N-glycosylation sites of P2X(3) receptor (Asn(139), Asn(170), Asn(194) and Asn(290)) and performed comparative analysis of the role of N-glycans on protein stability, plasma membrane delivery, trimer formation and inward currents. We have found that in transiently transfected HEK293 cells, Asn(170) is apparently the most important site for receptor stability, since its mutation causes a primary loss in protein content and indirect failure in membrane expression, oligomeric association and inward current responses. Even stronger effects are obtained when mutating Thr(172) in the same glycosylation consensus. Asn(194) and Asn(290) are the most dispensable, since even their simultaneous mutation does not affect any tested receptor feature. All double mutants containing Asn(170) mutation or the Asn(139)/Asn(290) double mutant are instead almost unable to assemble into a functional trimeric structure. The main emerging finding is that the inability to assemble into trimers might account for the impaired function in P2X(3) mutants where residue Asn(170) is replaced. These results improve our knowledge about the role of N-glycosylation in proper folding and oligomeric association of P2X(3) receptor.     

Neutral assembly of bacterial communities

Two recent, independent advances in ecology have generated interest and controversy: the development of neutral community models (NCMs) and the extension of biogeographical relationships into the microbial world. Here these two advances are linked by predicting an observed microbial taxa-volume relationship using an NCM and provide the strongest evidence so far for neutral community assembly in any group of organisms, macro or micro. Previously, NCMs have only ever been fitted using species-abundance distributions of macroorganisms at a single site or at one scale and parameter values have been calibrated on a case-by-case basis. Because NCMs predict a malleable two-parameter taxa-abundance distribution, this is a weak test of neutral community assembly and, hence, of the predictive power of NCMs. Here the two parameters of an NCM are calibrated using the taxa-abundance distribution observed in a small waterborne bacterial community housed in a bark-lined tree-hole in a beech tree. Using these parameters, unchanged, the taxa-abundance distributions and taxa-volume relationship observed in 26 other beech tree communities whose sizes span three orders of magnitude could be predicted. In doing so, a simple quantitative ecological mechanism to explain observations in microbial ecology is simultaneously offered and the predictive power of NCMs is demonstrated.     

Population and social dynamics changes in ring-tailed lemur troops at Berenty,Madagascar between 1989 - 1999

In the present study, we recorded all births, immigrations, deaths, and emigrations for a population of ring-tailed lemurs at Berenty Reserve, Madagascar, between September 1989 and August 1999. In September 1989, three troops (C, B, and T) inhabited the study area of 14.2 ha. During the 10-year period, eight troop divisions, six evictions of females, and three troop takeovers of ranges by other troops occurred in and around the study area. Consequently, in August 1999, the number of troops in the same area increased to six (CX, C1, C2A, C2B, T1, and T2). The number of lemurs aged >1 year increased from 63 to 82, which resulted from 204 births, 58 immigrations, 125 deaths, and 118 emigrations. Of the 204 newborn lemurs during the study period, 103 died, 44 emigrated outside the study area, and 57 remained within the study area. The total number of lemurs that emigrated from natal troops was 69 (54 males and 15 females). Natal males left their troops around the age of 3. Non-natal males changed troops after a tenure varying from 1 to 7 years. Survival curves showed a fall in survival rates of both sexes to < 0.5 between the ages of 2 and 3. For females, the survival rate gradually decreased to < 0.2 at the age of 9. On the other hand, due to emigration, the survival rate of males could not be determined after the age of 5 yr. Since some males attained high-rank at the age of 6 – 10 yr, the prime age for male ring-tailed lemurs is thought to be around 7 – 10 yr. Ring-tailed lemurs are essentially female philopatric, because all cases of females leaving natal troops resulted from troop divisions or forced evictions. Such social changes may have resulted from competition among females. All cases of troop divisions or evictions occurred in larger troops consisting of ≥20 lemurs, and only a few females could rejoin their troops. When males joined such a female-group, a new troop was formed. Although promoted by an increase in population, frequent emigrations of females from original troops are the characteristics of ring-tailed lemurs at Berenty.     

Unraveling assembly of stream biofilm communities

Microbial biofilms assemble from cells that attach to a surface, where they develop into matrix-enclosed communities. Mechanistic insights into community assembly are crucial to better understand the functioning of natural biofilms, which drive key ecosystem processes in numerous aquatic habitats. We studied the role of the suspended microbial community as the source of the biofilm community in three streams using terminal-restriction fragment length polymorphism and 454 pyrosequencing of the 16S ribosomal RNA (rRNA) and the 16S rRNA gene (as a measure for the active and the bulk community, respectively). Diversity was consistently lower in the biofilm communities than in the suspended stream water communities. We propose that the higher diversity in the suspended communities is supported by continuous inflow from various sources within the catchment. Community composition clearly differed between biofilms and suspended communities, whereas biofilm communities were similar in all three streams. This suggests that biofilm assembly did not simply reflect differences in the source communities, but that certain microbial groups from the source community proliferate in the biofilm. We compared the biofilm communities with random samples of the respective community suspended in the stream water. This analysis confirmed that stochastic dispersal from the source community was unlikely to shape the observed community composition of the biofilms, in support of species sorting as a major biofilm assembly mechanism. Bulk and active populations generated comparable patterns of community composition in the biofilms and the suspended communities, which suggests similar assembly controls on these populations.     

Adaptive and phylogenetic significance of ontogenetic sequences in Archaeolemur, subfossil lemur from Madagascar

Among the best known of recently extinct Malagasy lemurs is Archaeolemur, which is represented by many hundreds of specimens. The phylogenetic affinities of this taxon are unclear, especially in light of recent preliminary analysis of ancient DNA which does not support its previously accepted close relationship with the living Indridae. We examined the nearly complete skeletons of two adults and one juvenile and other less complete specimens to reconstruct aspects of the ontogeny of Archaeolemur. To compare the development of Archaeolemur to that of living strepsirrhines we collected data on Propithecus verreauxi, Eulemur fulvus, and Lemur catta. Additionally, because Archaeolemur exhibits some morphological convergences with distantly related papionins, we tested for convergence in the developmental patterns of Archaeolemur and Macaca fascicularis. Data include the status of tooth eruption, craniofacial sutural closure, and postcranial epiphyseal fusion, as well as linear measurements. We used discriminant function analysis and other tools to explore ontogenetic similarities and differences. The adaptive and phylogenetic significance of ontogenetic information is discussed. Our analysis shows that Archaeolemur displays a clear strepsirrhine pattern of development with only minor macaque convergences. Among the Strepsirrhini, Archaeolemur is slightly more similar developmentally to E. fulvus and L. catta than to P. verreauxi. Some of the distinctive features of the ontogeny of Archaeolemur may be related to diet, while others bear apparent testimony to a relatively rapid absolute pace of growth and development.     

Seasonal changes in general activity, body mass and reproduction of two small nocturnal primates: a comparison of the golden brown mouse lemur ( Microcebus ravelobensis) in Northwestern Madagascar and the brown mouse lemur ( Microcebus rufus) in Eastern Madagascar

To investigate for the first time the relationship between contrasting patterns of seasonal changes of the environment and activity, body mass and reproduction for small nocturnal primates in nature, we compared a population of golden brown mouse lemur (Microcebus ravelobensis) in a dry deciduous forest of northwestern Madagascar and of the brown mouse lemur (Microcebus rufus) in an evergreen rain forest of eastern Madagascar. Both species live under similar photoperiodic conditions. Golden brown mouse lemurs (GBML) were active during the whole period (May to December) irrespective of changing environmental conditions. In contrast, a part of the population of brown mouse lemurs (BML) showed prolonged seasonal torpor, related to body mass during periods of short day length and low ambient temperatures. Differences between species might be due to differences in ambient temperature and food supply. Body weight and tail thickness (adipose tissue reserve) did not show prominent differences between short and long photoperiods in GBML, whereas both differ significantly in BML, suggesting species-specific differences in the photoperiodically driven control of metabolism. Both species showed a seasonal reproduction. The rate of growth and size of the testes were similar and preceded estrous onset in both species suggesting a photoperiodic control of reproduction in males. The estrous onset in females occurred earlier in GBML than in BML. Estrous females were observed over at least 4 months in the former, but in only 1 month in the latter species. Intraspecific variation of estrous onset in GBML may be explained by body mass. Interspecific variation of female reproduction indicates species-specific differences in the control of reproduction. Thus, environmentally related differences in annual rhythms between closely related small nocturnal lemurs emerged that allow them to cope with contrasting patterns of seasonal changes in their habitats.     

Daily torpor in the gray mouse lemur (Microcebus murinus) in Madagascar: energetic consequences and biological significance

Patterns and energetic consequences of spontaneous daily torpor were measured in the gray mouse lemur (Microcebus murinus) under natural conditions of ambient temperature and photoperiod in a dry deciduous forest in western Madagascar. Over a period of two consecutive dry seasons, oxygen consumption (VO₂) and body temperature (T _b) were measured on ten individuals kept in outdoor enclosures. In all animals, spontaneous daily torpor occurred on a daily basis with torpor bouts lasting from 3.6 to 17.6 h, with a mean torpor bout duration of 9.3 h. On average, body temperatures in torpor were 17.3±4.9°C with a recorded minimum value of 7.8°C. Torpor was not restricted to the mouse lemurs’ diurnal resting phase: entries occurred throughout the night and arousals mainly around midday, coinciding with the daily ambient temperature maximum. Arousal from torpor was a two-phase process with a first passive, exogenous heating where the T _b of animals increased from the torpor T _b minimum to a mean value of 27.1°C before the second, endogenous heat production commenced to further raise T _b to normothermic values. Metabolic rate during torpor (28.6±13.2 ml O₂ h^–1) was significantly reduced by about 76% compared to resting metabolic rate (132.6±50.5 ml O₂ h^–1). On average, for all M. murinus individuals measured, hypometabolism during daily torpor reduced daily energy expenditure by about 38%. In conclusion, all these energy-conserving mechanisms of the nocturnal mouse lemurs, with passive exogenous heating during arousal from torpor, low minimum torpor T _bs, and extended torpor bouts into the activity phase, comprise an important and highly adapted mechanism to minimize energetic costs in response to unfavorable environmental conditions and may play a crucial role for individual fitness. Received: 8 July 1999 / Accepted: 3 December 1999     

Arrival order among native plant functional groups does not affect invasibility of constructed dune communities

Different arrival order scenarios of native functional groups to a site may influence both resource use during development and final community structure. Arrival order may then indirectly influence community resistance to invasion. We present a mesocosm experiment of constructed coastal dune communities that monitored biotic and abiotic responses to different arrival orders of native functional groups. Constructed communities were compared with unplanted mesocosms. We then simulated a single invasion event by bitou (Chrysanthemoides monilifera ssp. rotundata), a dominant exotic shrub of coastal communities. We evaluated the hypothesis that plantings with simultaneous representation of grass, herb and shrub functional groups at the beginning of the experiment would more completely sequester resources and limit invasion than staggered plantings. Staggered plantings in turn would offer greater resource use and invasion resistance than unplanted mesocosms. Contrary to our expectations, there were few effects of arrival order on abiotic variables for the duration of the experiment and arrival order was unimportant in final community invasibility. All planted mesocosms supported significantly more invader germinants and significantly less invader abundance than unplanted mesocosms. Native functional group plantings may have a nurse effect during the invader germination and establishment phase and a competitive function during the invader juvenile and adult phase. Arrival order per se did not affect resource use and community invasibility in our mesocosm experiment. While grass, herb and shrub functional group plantings will not prevent invasion success in restored communities, they may limit final invader biomass.     

Reptile, amphibian, and lemur diversity of the Malahelo Forest, a biogeographical transition zone in southeastern Madagascar

Ambatorongorongo Mountain lies at the historical intersection betweenhumid, spiny, and littoral forests in southeastern Madagascar. We report theresults of surveys of the herpetofauna and lemurs occurring in Malahelo Forest,a small (<25 ha) forest fragment lying on the western slope ofAmbatorongorongo Mountain. There are at least 41 reptile, 11 amphibian, and 7lemur species in this forest, including several that are endemic to southeasternMadagascar and are at severe risk of extinction. The species richness of theMalahelo fauna is comparable to that of even the largest forest reserves in theregion. We also evaluate the similarity of the Malahelo herpetofauna to that ofnearby humid, spiny, and littoral forests to assess the biogeographic affinitiesof its amphibians and reptile assemblages. Both groups contain speciescharacteristic of each of the three surrounding forest types, but thebiogeographic patterns appear to differ for amphibians and reptiles. Overall,the herpetofauna and lemurs of the Malahelo Forest indicate that it is a remnantof a biogeographic transition zone between the major forest types ofsoutheastern Madagascar. The combination of high species richness, regionalendemics, and unique herpetofaunal and lemur assemblages should make MalaheloForest a high conservation priority, and we give recommendations for protectingwhat remains of this important transitional forest.     

A simple rule for the evolution of contingent cooperation in large groups

Humans cooperate in large groups of unrelated individuals, and many authors have argued that such cooperation is sustained by contingent reward and punishment. However, such sanctioning systems can also stabilize a wide range of behaviours, including mutually deleterious behaviours. Moreover, it is very likely that large-scale cooperation is derived in the human lineage. Thus, understanding the evolution of mutually beneficial cooperative behaviour requires knowledge of when strategies that support such behaviour can increase when rare. Here, we derive a simple formula that gives the relatedness necessary for contingent cooperation in n-person iterated games to increase when rare. This rule applies to a wide range of pay-off functions and assumes that the strategies supporting cooperation are based on the presence of a threshold fraction of cooperators. This rule suggests that modest levels of relatedness are sufficient for invasion by strategies that make cooperation contingent on previous cooperation by a small fraction of group members. In contrast, only high levels of relatedness allow the invasion by strategies that require near universal cooperation. In order to derive this formula, we introduce a novel methodology for studying evolution in group structured populations including local and global group-size regulation and fluctuations in group size.