Experimentally induced polygyny in a monogamous bird species: prothonotary warblers and the polygyny threshold

Summary Hypotheses regarding evolution of polygyny were tested using prothonotary warblers (Protonotaria citrea), through experimental manipulations of habitat quality. Two experiments were performed. Experiment 1 involved supplying nest boxes in different breeding habitats: flooded riparian areas (high food availability) and adjacent dry bottomland (low food availability). Nine bigamous matings were induced in this experiment, all occurring in flooded habitats even though monogamous mating opportunities existed in dry areas. Costs of polygyny for secondary females in flooded habitat were similar to costs incurred by monogamous females in dry habitat. Thus, a polygyny threshold apparently existed, possibly based on differences in food abundance in the two habitats. However, male quality covaried with habitat quality, as males in flooded areas were older and larger than males in dry habitat. Secondary females may have chosen polygyny in the best breeding situation, a combination of male and territory quality. In Experiment 2 nest box density was varied within flooded habitat to allow males to monopolize different numbers of nest-sites. Monogamous females settled earlier on territories with a large number of nest-sites, and males that defended many nest-sites were more likely to become polygynous. Male physical characteristics were not related to occurrence of polygyny in flooded habitat. Limitation of suitable nest-holes ultimately constrains occurrence of polygyny in prothonotary warblers.     

Resource defense polygyny shifts to female defense polygyny over the course of the reproductive season of a Neotropical harvestman

Although studies classify the polygynous mating system of a given species into female defense polygyny (FDP) or resource defense polygyny (RDP), the boundary between these two categories is often slight. Males of some species may even shift between these two types of polygyny in response to temporal variation in social and environmental conditions. Here, we examine the mating system of the Neotropical harvestman Acutisoma proximum and, in order to assess if mate acquisition in males corresponds to FDP or RDP, we tested four contrasting predictions derived from the mating system theory. At the beginning of the reproductive season, males fight with other males for the possession of territories on the vegetation where females will later oviposit, as expected in RDP. Females present a marked preference for specific host plant species, and males establish their territories in areas where these host plants are specially abundant, which is also expected in RDP. Later in the reproductive season, males reduce their patrolling activity and focus on defending individual females that are ovipositing inside their territories, as what occurs in FDP. This is the first described case of an arachnid that exhibits a shift in mating system over the reproductive season, revealing that we should be cautious when defining the mating system of a species based on few observations concentrated in a brief period. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Compensating for the costs of polygyny in hen harriers Circus cyaneus

In polygynous species, the adults are faced with a dilemma during chick rearing. Males must decide how to distribute food between their females and food allocation patterns are often highly unequal. In turn, the females that receive less food from males have to decide how much time to invest in additional hunting. If they spend more time hunting, then they leave their young exposed to weather and predators. However, if they stay at the nest, they increase the risk of their chicks starving. One way that birds may compensate for reduced provisioning is by increasing the size of prey caught. We tested this hypothesis by comparing prey deliveries to nests of hen harriers, Circus cyaneus, with females of different breeding status. As expected, male harriers delivered less food items to the nests of polygynous females, and especially their secondary, or β females. However, both sexes were able to compensate by delivering larger items and there was no difference in the overall mass of food delivered to nests. Moreover, females spent a similar amount of time at the nest, irrespective of status, and there were no overall differences in breeding success. Our results show that polygynous female harriers can compensate for the costs of polygyny, but we suggest that their ability to do so will vary according to the abundance of both large prey and predators.     

Towards the definition of budget models for the evolution of deltas

Global climatic change is likely to take place and could eventually affect Mediterranean deltas and other lowlying coastal regions. This would have serious implications for the natural resources of these deltaic areas, as well as for human settlements and related economic activities. To achieve sound decision making, to prevent damages and to avoid risky investments, it is necessary to understand the integral functioning of deltaic areas and to determine their vulnerability and response to large-scale change phenomena. Optimal use of the available knowledge will require that existing and new field measurements are combined and that integrated (physical/ecological) conceptual models of deltaic behaviour are developed with socio-economics scenarios as boundary conditions. This paper illustrates the methodological effort towards organizing a modeling framework to conduct budget computations at various scales with reference to the most significant ‘physiogrpahic units’ and to the most significant deltaic processes. The final objective is to handle the problem of evaluating possible changes under different scenarios.     

Colony foundation and polygyny in the ant Formica podzolica

We document the variation in number of queens occurring naturally in founding, immature and mature nests of the ant Formica podzolica, and compare development of colonies and survivorship of queens in experimental nests started with 1–16 foundresses. Number of queens per nest was associated with stage of colony development. Most nests were monogynous, but 20% of immature nests (n = 66) and 25% of mature nests (n = 92) were oligogynous or polygynous. Colonies were usually established by single queens (i.e., haplometrosis), but colony establishment by multiple queens (i.e., pleometrotis) was also common, occurring in 27% of founding nests (n = 492). Foundress groups in the field were small ( = 1.47 ± 0.04 queens/nest), and large groups experienced high mortality and low productivity in artificial nests. Therefore, the many queens (up to 140) in some immature and mature colonies were probably secondarily pleometrotic. Experimental nests started with 1–4 queens were more successful than those initiated by 8 or 16 queens. Small groups (2–4 queens) produced more pupae before the first nests reared workers than single foundresses or larger groups (8 or 16 queens). Although single foundresses were less productive than queens in small groups, they experienced greater survivorship and less weight loss than queens in pleometrotic associations. Besides low productivity, queen mortality and weight loss were greatest in large groups.     

Nest site limitation and facultative polygyny in the ant Leptothorax longispinosus

Summary The ant L. longispinosus displays geographic variation in its pattern of facultative polygyny (Fig. 2). In nature, nest density and frequency of multiple queening are positively associated over three sites. A putative causal relation between availability of vacant nest sites and polygyny was examined in New York, where a plot was seeded with additional nest sites and monitored for 24 months. Both queen number and worker number per nest on the experimental plot were reduced relative to controls (Fig. 4, Fig. 5), indicating that scarcity of available nest sites influences the pattern of polygyny in this species. The observed demographic changes resulted from fractionation of existing colonies; adding nest sites induced polydomy. Although numbers of adult ants changed with addition of nest sites, the numbers of immatures were no different after 2 years (Table 1), suggesting that the population was undergoing growth to expand into the additional sites. These results are the first direct experimental evidence linking polygyny to an ecological parameter for any ant species.     

Multiple mating and facultative polygyny in the Panamanian leafcutter ant Acromyrmex echinatior

Queen mating frequency of the facultatively polygynous ant Acromyrmex echinatior was investigated by analysing genetic variation at an (AG)_n repeat microsatellite locus in workers and sexuals of 20 colonies from a single Panamanian population. Thirteen colonies were found to be monogynous, 5 colonies contained multiple queens, whereas the queen number of 2 colonies remained unresolved. Microsatellite genotypes indicated that 12 out of 13 queens were inseminated by multiple males (polyandry). The mean queen mating frequency was 2.53 and the mean genetically effective paternity frequency was 2.23. These values range among the highest found in ants, and the results are in keeping with the high mating frequencies reported for other species of leafcutter ants. Consistent skew in the proportional representation of different patrilines within colonies was found, and this remained constant in two consecutive samples of offspring. Dissections showed that all examined queens from multiple-queen colonies were mated egg-layers. The mean relatedness value among nestmate workers in polygynous colonies was lower than that for monogynous colonies. No diploid males were detected in a sample of 70 genotyped males. Worker production of males was detected in one queenless colony. We discuss our findings in relation to known patterns of multiple maternity and paternity in other eusocial Hymenoptera. Received: 2 September 1998 / Received in revised form: 3 February 1999 / Accepted: 7 February 1999     

Secondary polygyny by inbred wingless sexuals in the dolichoderine ant Technomyrmex albipes

Summary Technomyrmex albipes makes huge polydomous colonies which consist of up to several millions of adults. In field colonies, dealate queens are rare or absent, though winged males and winged females emerge annually (synchronously) in large numbers from late may to mid June. Field and laboratory observations showed that the reproduction of established colonies was performed by wingless females inseminated by wingless males from the same colony. Dissections and morphological examinations revealed that wingless females are workers with no spermatheca and intercastes with a spermatheca. Most intercastes were inseminated, had developed ovaries, and seemed to reproduce, while workers did not seem to reproduce. Extranidal tasks were performed only by workers. Approximately half of the adult population were intercastes, and wingless males represented only a small portion of all adults, the rest being nonreproductive workers. Intercastes and wingless males were produced throughout the year except in winter. The winged females and males copulate outside the nest only after the nuptial flight and the dealate females are able to perform independent founding, but they are also eventually supplanted by intercastes. The adoption of dealate queens by an established natal colony did not seem to occur. Thus we infer that in this species the winged reproductives disperse and found new colonies, while inbred wingless reproductives allow the enlargement and budding of colonies. This species has a special trophic-flow system. There is no trophallaxis among adults, and nutrient transfer from adults to other colony members is achieved exclusively by specialized trophic eggs. All females (dealate queens, intercastes, and workers) seem to produce trophic eggs. This aphid-like life cycle, i.e., the occurrence of both winged and wingless reproductive forms, may have evolved as an adaptation supporting the development of secondary polygyny and polydomy.Offprint requests to: K. Yamauchi     

Serial polygyny and colony genetic structure in the monogynous queenless ant Diacamma cyaneiventre

Serial polygyny, defined as the temporal succession of several reproductive females in a colony, occurs in some monogynous social insects and has so far attracted little attention. Diacamma cyaneiventre is a queenless ponerine ant found in the south of India. Colonies are headed by one singly mated worker, the gamergate. After the death of the gamergate or her absence following colony fission, the gamergate is replaced by a newly eclosed nestmate worker. After a replacement, colonies go through short-lived periods in which two matrilines of sisters co-occur. This is a situation which can be described as serial polygyny. To measure the consequences of serial polygyny, a genetic analysis was performed on 449 workers from 46 colonies of D. cyaneiventre using five microsatellite loci. The presence of more than one matriline among workers of the same nest was detected in 19% of colonies, indicating a recent change of gamergate. The average genetic relatedness among nestmate workers was 0.751 and did not significantly differ from the theoretical expectation under strict monogyny and monandry (0.75). A simple analytical model of the temporal dynamics of serial polygyny was developed in order to interpret these results. We show that the rate of gamergate turnover relative to the rate of worker turnover is the crucial parameter determining the level of serial polygyny and its effect on the genetic structure of colonies. This parameter, estimated from our data, confirms that serial polygyny occurs in D. cyaneiventre but is not strong enough to influence significantly the average genetic relatedness among workers.     

Intrasexual selection in Mirza coquereli : evidence for scramble competition polygyny in a solitary primate

The primates of Madagascar (Lemuriformes) deviate from fundamental predictions of sexual selection theory in that polygynous species lack sexual dimorphism, have even adult sex ratios and often live in female-dominated societies. It has been hypothesized that intrasexual selection in these species is either reduced or primarily focused on traits related to scramble competition. The goal of this study was to examine these hypotheses by studying the mating system of a solitary nocturnal species, Mirzacoquereli. During a 4-year field study in western Madagascar, I captured and followed 88 individually marked animals. I found that adult males were significantly larger than females, providing the first evidence for sexual size dimorphism in lemurs. In addition, the adult sex ratio was biased in favour of females in 3 out of 4 years. There was no significant sex difference in canine size, however. Males showed pronounced seasonal variation in testis size with a 5-fold increase before and during the short annual mating season. During the mating season, males had more injuries than females and more than quadrupled their home ranges, overlapping with those of more than ten females, but also with about the same number of rivals. Only about one social interaction per 10 h of observation was recorded, but none of them were matings. Together, these results indicate that these solitary lemurs are clearly subject to intrasexual selection and that male-male competition is primarily, but not exclusively, of the scramble type. In addition, they suggest that the above-mentioned idiosyncracies may be limited to group-living lemurs, that social systems of solitary primates are more diverse than previously thought, and that the temporal distribution of receptive females is responsible for this particular male mating strategy. Received: 11 January 1997 / Accepted after revision: 18 April 1997     

Female defence polygyny in the bumblebee wolf,Philanthus bicinctus (Hymenoptera: Sphecidae)

Summary In the digger wasp Philanthus bicinctus the first adult males emerge about a week before the first females. The sex ratio of emerging wasps is near unity. Females appear to mate once and do so when they first start to nest, not when they first emerge.Males defend mating territories in a large area of female ground nests. Territoriality begins when females first start to nest and throughout the season coincides with the digging activities of females initiating rests. During the day territoriality occurs only when females are engaged in digging activities.Males compete for certain territory sites within the resting area; the most frequently occupied male territories are in areas where the density of nests is highest. About half of the males are excluded from nesting area territories. Some of these males defend areas among the flowers of snakeweed where females occasionally forage; this appears to be a loser tactic of males unable to obtain nesting area territories.Although there was no significant difference in the sizes of males on nesting area and snakeweed territories, within the nesting area larger males usually occupy territories with the higher nest densities.     

Extra-pair paternity in willow ptarmigan broods: measuring costs of polygyny to males

Willow ptarmigan are one of only three monogamous grouse species in North America. However, in some populations between 5 and 20% of individuals pair polygynously. It has been suggested that monogamy may be maintained by the high cost of polygyny to males. We have used DNA fingerprinting to assess the actual reproductive success of both monogamous and polygynous adults. We determined whether or not the putative parents were the biological parents of the chicks from 38 broods. Of these clutches 30 were from monogamous matings, and 8 were from bigamous matings. Of the 207 chicks from monogamous matings 96% were within-pair offspring, compared to 67% of the 49 chicks from bigamous matings. All extra-pair offspring chicks resulted from extra-pair fertilizations (EPFs), and there were no instances of intraspecific nest parasitism. Mate guarding by monogamous males seems to be a highly effective method for maintaining genetic monogamy, as the only cases in which EPFs occurred were when the resident female left the territory for a few days or when a second female visited the territory. Our results support the notion that certainty of parentage may be one factor constraining willow ptarmigan males to be monogamous.     

Testing hotspot models of lek evolution: data from three species of ungulates

Summary Hotspot models propose that leks have evolved because certain sites are associated with extremely high female encounter rates. In this paper we test five predictions of hotspot models, using data from studies of three species of lekking ungulates. As predicted, we found that lekking species generally had large female home ranges. However, in contrast to other predictions of hotspot models, ungulate leks did not occur at sites of maximum adult female density, lekking was not associated with low overall female density, and the adult sex ratio and the proportion of females that were in oestrus both differed substantially between leks and off-lek areas. Our empirical results therefore confirm recent theoretical assessments of the hotspot process, suggesting that while hotspots may help explain broad patterns of male dispersion, further mechanisms are needed to generate the extent of territory clustering seen at leks. Correspondence to: A. Balmford     

Scramble competition polygyny in thirteen-lined ground squirrels: the relative contributions of overt conflict and competitive mate searching

Summary The scramble competition polygyny of male thirteen-lined ground squirrels involves overt conflict and sperm competition as well as competitive mate searching. The overt conflict component of intrasexual competition occurs sporadically: males sometimes are able to mate in the complete absence of competitors, while on other occasions up to five other males may be present. We examined the immediate effects of conflict on male mating behavior, and found that it increases the time spent in pursuit of an estrous female, reduces the efficiency of copulating, but carries only a small risk of exclusion from mating. In contrast to some other sciurid species, dominance appears to be a poor predictor of the mating success of male thirteen-lined ground squirrels. However, skill at locating females is very important. Because overt conflict seldom results in exclusion from mating opportunities, and because females of this species accept multiple mates, success in copulating conforms closely with success in mate searching: the sheer number of estrous females located is a consistently strong predictor of male mating success. Furthermore, individual differences in mate location ability seem to generate levels of variance in male copulatory success that are comparable to those of more combative species.     

Advances in the study of directed evolution for cellulases

If cellulose can be effectively hydrolyzed into glucose by cellulase, the production costs of hydrogen, ethanol or other chemicals from cellulosic materials will be greatly decreased, and economically viable production of biohydrogen and bioethanol will become feasible. Cellulose is degraded into glucoses by multi-component enzyme systems. Nowadays cellulases are widely used in brewing, food, bioenergy, fodder, textiles, paper, pharmaceuticals, environmental protection and other industries. However, existing cellulases have several problems that limit their wider applications, including the low turnover number for solid cellulosic materials, and low stability in adapting to various application conditions. For example, high temperature, low pH, and so on. Application of directed evolution technology may be one of the most effective ways for improving the characteristics of cellulases. This paper presents a brief review of the cellulose hydrolysis mechanism by cellulase, advances in cellulases (endoglucanase and β-glucosidase) improvement by directed evolution for several characteristics (for instance, thermal stability, pH adaptability and enzyme activity), limitations of directed evolution for cellulases, and the outlook for directed evolution for cellulase.     

Persistence models for mark-recapture

The stable of models available for analyzing mark-recapture data (Otis et al. Wild Momogr 66:135, 1978) includes those having behavioral characteristics, time variation, heterogeneity, along with combinations of those characteristics. This paper proposes use of a series of models based on the persistence model of Ramsey and Usner (Biometrics 59:331–339, 2003). We show that persistence can be modeled in combination with behavior and with time variation. We apply the persistence model to situations in which capture occasions are not equally-spaced in time. Two case studies illustrate the use of these extended persistence models.     

Sex differences in fitness following a group take-over among Toque macaques: testing models of social evolution

Summary A group of toque macaques took-over the home range of one of its subordinate neighboring groups and fused with it to form a larger cohesive group. In the 7 years before the take-over, the dominant group had consistently won all contests at common feeding sites, yet the fitnesses of the females of these two groups did not differ significantly (Fig. 2A). After the take-over the females of the subjugated group occupied the lowest ranks in the combined dominance hierarchy of the merged groups (Fig. 1) and thereby lost the advantages of an own home range, such as priority of access to food. Consequently, in the merged group, survivorship and reproductive success among the subjugated females were significantly less than among the females of the dominant subgroup (Table 2, 4). The dominant matrilines grew numerically and replaced all of the subjugated females, and all but one of their offspring, within 8 years after the take-over (Fig. 2B). These data support the hypothesis that cooperation among female kin in defending resources against strange females is important in the evolution of female-bonded groups. Before the merger all 5 natal males of the subordinate group had transferred to the dominant group, where they occupied high and mid-level dominance ranks (Fig. 1). These males survived at a significantly greater rate than their subordinate female kin. Thus, the cost of group transfer seems to be greater for females than for males, and this may be one reason that females generally do not emigrate or that groups do not fuse. The data suggested three hypotheses. First, since large body size and other adaptations for fighting, giving males an advantage in male-male competition for mates, are also of advantage in resource competition with males and females, such male characters may also be favored by non-sexual selection, especially where male reproductive strategy involves group transfer. Second, female bonded groups evolved as female defensive coalitions against not only female but also male resource competitors, there having been a mutual influence in the coevolution of large-sized males and female gregariousness. Third, female defensive coalitions against large-sized aggressive males are also advantageous out-side the context of food competition, or, independent of foraging strategy.     

Using the maximum principle of impulse control for ecology-economical models

In this work we present mathematical models for population of single cohort and homogeneous animals. Investigating these mathematical models, we determine structure of optimal impulsive control which used maximum principle for optimal processes with impulse control.     

A reliability index for models

Since the prediction of environmental changes depends heavily on mathematical models, environmental planners have a definite need for a general test that can be easily applied to determine the reliability of a model or to choose between available models. In this paper a ‘reliability index’ for a model is defined. This index is a number k ? 1 which is determined from a set x₁,x₂,…,x_n of model predictions and a corresponding set y₁,y₂,…,y_n of observations. One interpretation of the index k is that the model predictions agree with observations, in some sense, within a factor of k. The index k is defined using an intuitive, geometric approach and is justified through agreement with another index developed using statistical techniques. Examples are given for a model which predicts the atmospheric movement of radon-222 originating from a concentrated source of radium-226, such as a uranium tailings pile.