Colony size and reproduction in the western harvester ant, Pogonomyrmex occidentalis

Summary: We report data from a four-year field study on the relationship between colony size and reproduction in the western harvester ant, Pogonomyrmex occidentalis. In all years, the likelihood of reproduction significantly increased with increasing size in both field censuses during naturally-occurring mating flights and experimentally-watered colonies whose entire reproductive output was collected. However, the total amount of reproductive biomass was unrelated to colony size. We describe the size threshold for reproduction in P. occidentalis and show that it varies across years. Once colonies become reproductively mature, they reproduce consistently although not in every year. We describe a method for collecting the entire reproductive output for desert ants whose reproductive flights are cued by rainfall.     

The tissue composition of<Emphasis Type="Italic"> Montastraea franksi</Emphasis> during a natural bleaching event in the Florida Keys

In this study, we used a correlative approach to (1) test for an association between bleaching and host tissue composition during a natural bleaching event, and (2) assess whether bleaching susceptibility varies between years. In August 1997, Montastraea franksi at 15-m depth on Conch Reef, Florida, bleached and the severity of the response varied among individuals. Seventy-five randomly selected colonies were quantified for bleaching using both an ordinal scale, assigned by eye, and a continuous scale, assessed using red, green, and blue (RGB) spectral analysis of photographs. Zooxanthella density and chlorophyll a content were evaluated as measures of bleaching, and coral tissue was analyzed for glycerol, free amino acids (FAA), protein, and mycosporine-like amino acids (MAAs); collectively, these are described as the "tissue composition." In 1998, most of the same coral colonies were analyzed for color and zooxanthella density to determine whether colony color in 1997 was correlated with color in 1998. In 1997, colonies of M. franksi that were ranked by color differed significantly in RGB brightness, zooxanthella density, and chlorophyll a content, but not in tissue composition. Similarly, a multivariate test for a linear relationship between color and tissue composition did not reveal a significant association. Analyses of corals in 1997 and 1998 revealed a significant positive relationship between color in both years (i.e., the same colonies were similarly colored in each year). These results are discussed in the context of the temporal scale of the sampling regime, the nature of the measured traits, and the adaptive bleaching hypothesis.     

Factors limiting rosette recruitment in scarlet gilia, Ipomopsis aggregata: seed and disturbance limitation

We performed a factorial combination of seed additions, surface soil disturbance, and protection from potential seed predation in experimental plots to elucidate the relative importance of each factor in the recruitment of scarlet gilia seedlings. We detected a significant interaction between seed addition and surface soil disturbance on seedling recruitment over 2 years of establishment. Plots that received both supplemental seed addition and disturbance established 10.5 times as many rosettes as control plots. We did not detect main or interactive effects of caging on seedling establishment. We explored the importance of density-dependent recruitment by investigating the number of rosettes per experimental plot through time. In addition, we used planned contrasts to compare the absolute and percent changes in control and treatment plot rosette densities between the study years. We found significant downward concave curvature to the seedling recruitment curve, suggesting negative density-dependent recruitment across the experimental range of rosette densities. We found a significant difference between the control plots and the seed addition-disturbance plots in terms of absolute changes in plot density from 1997 to 1998. Greater net mortality occurred in plots that received both seed addition and disturbance. However, we found no differences among treatments in the percent change in rosette density across the study years. Our study demonstrates that seedling recruitment in natural plant populations may be limited by the interaction of seed and microsite availability, and that seedling recruitment in scarlet gilia may be negatively density dependent. Our results suggest that the fecundity effects of particular plant-animal interactions (e.g., pollination, herbivory) may affect scarlet gilia population dynamics, particularly under conditions of high soil disturbance. Received: 11 June 1999 / Accepted: 24 November 1999     

Influence of Edge Exposure on Tree Seedling Species Recruitment in Tropical Rain Forest Fragments1

Edge creation has a pronounced influence on the understory vegetation, but the effects of edges on seedling species recruitment are still poorly understood. In Central Amazonia, 9–19 years after fragmentation, we recorded species richness and net seedling recruitment rate in 1 ha blocks exposed to none, one, or multiple edges within forest fragments. One‐hectare blocks were located in the center (no edge), the edge (one edge), the corners (two edges) of 10 and 100 ha fragments, and in a 1 ha fragment (four edges). In 1991, we counted all tree seedlings 5–100 cm tall found within permanent 1 m² plots located within the 1 ha blocks. In May 1993, we manually removed all seedlings that were smaller than 1 m tall from the permanent plots. Six years and five months later (October 1999), all new seedlings recruited into the plots were counted and classified into distinct morphospecies. Species richness of recruited seedlings, scaled by total seedling density, declined from the center to the edge, the corner blocks, and then to the 1 ha fragment. Overall, the four‐edged, 1 ha fragment had the poorest species richness and the non‐edged 100 ha central block the highest. The total number of recruited individuals was 40 percent less than that previously present, with the 100 ha corner having the lowest recruitment. Pairwise comparisons showed that species similarity was related to edge number for the 100 and 1 ha fragments. Species rank/abundance curves showed that a subset of species was common in all blocks within the fragments, and that the 100 ha center held more rare species than any other 1 ha block. This study demonstrated that, in a given fragment patch, the number of tree seedling species recruited varied inversely with the number of edges.     

Reproductive schedule and factors affecting soldier production in the eusocial bamboo aphid Pseudoregma bambucicola (Homoptera, Aphididae)

Summary: The reproductive characteristics of the soldier-producing aphid Pseudoregma bambucicola were studied in Kagoshima, Southern Japan, to know the factors affecting soldier production of eusocial aphids. The soldier proportion in aphid colonies was highest from October to November. In some large colonies, soldiers were observed in all seasons except in July when colony size was relatively small. Multiple regression analysis showed that the colony size was a principal factor affecting soldier proportion throughout a year. Other social or environmental factors such as aphid composition, host plant conditions and predator abundance were not always significant. Rearing experiments revealed that large colonies (̿,000 individuals) produced soldiers in almost all seasons while small colonies (<1,000) never produced any soldiers. The caste-production schedule of adult females was examined in the field. When solitary females produced both castes, they usually produced normal nymphs first and then soldiers. Females from large colonies tended to produce more soldiers in the earlier period of their lifetime, whereas females from newly established small colonies produced no or only a few soldiers at later times. The average number of soldiers and normal nymphs produced consecutively by a single female was >10 and >20, respectively. Because they have a small number of ovarioles (<15 on average), females should alter caste production within the same ovarioles according to changes in environmental conditions. Artificial removal or introduction of predators and reduction of colony size did not affect soldier production over two successive generations, revealing maternal effects on soldier production. Females cannot shift caste production quickly in response to changes in predator abundance and colony size. This is probably due to early developmental determination of castes within the mother's body.     

Coffee farm diversity and landscape features influence density of colonies of Atta cephalotes (Hymenoptera: Formicidae)

The density of colonies of leaf-cutting ants, Atta cephalotes L. (Hymenoptera: Formicidae), was measured and compared among coffee (Coffea arabica L.) plantations in five management categories: monoculture conventional, diversified conventional, diversified organic, highly diversified conventional, and highly diversified organic. Twenty-four small farms (<4 ha) in Turrialba, Costa Rica, were included in this study. Within-farm and off-farm (landscape) variables were measured and tested for their relationship with A. cephalotes colony density. Total ant colony density (colonies per ha) and density of new colonies shortly after a nuptial flight were significantly greater in the coffee monoculture conventional system, compared with all other systems. Total ant colony density and density of new colonies were inversely related to percentage of shade within the farms. Within farms, colony density was greater near edges adjacent to riparian forest than those adjacent to nonforested land. Regardless of edge type, plots closer to the edge (0-30 m) had greater colony density than those furthest from the edge. At the landscape scale, density of new colonies was positively related to fallow land use coverage within a 2,000-m buffer radius and to forest coverage within a 500-m radius. Results indicate that coffee farm management practices and landscape level factors can affect A. cephalotes colony densities. Understanding such practices and factors could assist in the development of better management methods of these injurious insects in coffee farms. Increased diversification in coffee farms, possibly due to the greater shade associated with it, may reduce colonization by the ants, which are considered forest gap specialists.     

Community interactions between ants and arboreal-nesting termites in New Guinea coconut plantations

Summary: We investigated the incidence of inquiline ants and of arboreal-nesting ants on a community of three arboreal-nesting termites living in New Guinea coconut plantations. Inquiline ants were present in 10 % of Microcerotermes biroi nests and in 4 % of Nasutitermes princeps nests. Live termite nests inhabited by the most common inquiline ant, Camponotus sp. A, were generally left by the ant after several months. In some nests, Camponotus sp. A was observed coexisting with its host during the whole observation period (3 years). Therefore, Camponotus sp. A was apparently an opportunistic inquiline which did not affect significantly the mortality of termite colonies. The arboreal-nesting ant, Crematogaster irritabilis, was locally found occupying up to 99 % of the trees present in 1 ha plots. In such hot spots, the overall abundance of termites was approximately half that of plots devoid of Crematogaster irritabilis. The high density of Crematogaster irritabilis may be an important limiting factor for the termite assemblage, by hastening the death or hindering the establishment of arboreal termite colonies.     

Suburban sprawl: environmental features affect colony social and spatial structure in the black carpenter ant,Camponotus pennsylvanicus

1. In social insects, the number of nests that a colony inhabits may have important consequences for colony genetic structure, the number of queens, sex allocation, foraging efficiency, and nestmate recognition. Within the ants, colonies may either occupy a single nest (monodomy) or may be organised into a complex network of nests and trails, a condition known as polydomy. 2. The current study is a large‐scale, long‐term, comprehensive field examination of various features of colony social and spatial structure in the facultatively polydomous black carpenter ant, Camponotus pennsylvanicus (DeGeer). The study examined the density, persistence, and the spatiotemporal distribution of colonies across a gradient of land disturbance associated with urban development. The temporal and spatial pattern of nest use was compared between fragmented landscapes where nesting sites were interspersed among human‐built structures (urban plots) and less disturbed landscapes with higher tree density (suburban plots). In addition, nesting site fidelity and changes in colony spatial structure were monitored over 7 years. 3. Long‐term monitoring and extensive sampling over a large spatial area allowed the first comprehensive insight into the spatiotemporal dynamics of colony and population structure in C. pennsylvanicus. A total of 1113 trees were inspected over 233 ha. Camponotus pennsylvanicus were active on 348 of the 1113 trees (31%) and these represented 182 distinct colonies. The colonisation rate remained relatively stable over 7 years suggesting that an equilibrium point had been reached. Relative to the suburban plots, tree density was 65% lower in the urban plots. The proportion of trees colonised by C. pennsylvanicus was significantly higher in the urban plots suggesting that intraspecific competition for nesting sites may be especially high in areas with lower tree density. Colony spatial structure also differed significantly between habitats and a higher incidence of monodomy was observed in the urban environment. The average number of trees per colony across all subplots was 1.95 (range 1–4) indicating that C. pennsylvanicus are weakly polydomous. 4. The composite picture that emerges for C. pennsylvanicus colonies in the urban habitat is a chain reaction of events: (i) the urban habitat has a lower tree density, (ii) lower tree density results in higher tree colonisation rate, (iii) higher tree colonisation rate results in simpler colony spatial structure (i.e. higher incidence of monodomy), and (iv) simpler colony spatial structure results in numerically smaller colonies. Long‐term monitoring of the spatiotemporal pattern of nest site use in selected colonies revealed a unique trend. While worker counts in selected colonies remained relatively stable throughout the course of the study, colony spatial structure changed considerably with 28% of colonies experiencing a change. Furthermore, the likelihood of detecting a change in colony spatial structure increased with the amount of time passing from the initial inspection. 5. In conclusion, tree density has a significant effect on a number of important colony features in C. pennsylvanicus. Besides tree density, other environmental features such as human‐built structures cause habitat fragmentation and may act as natural barriers to worker dispersal and/or foraging. Such barriers may ultimately affect the social and/or spatial structure at both the colony and the population level.     

Size and composition of swarming colonies in Provespa anomala (Hymenoptera, Vespidae), a nocturnal social wasp

Summary: Ninety-eight nests of various growing stages of Provespa anomala were found in the lowlands of Sumatra, Indonesia, from August to January. Its colony cycle is considered to be annual, with nests built throughout the year. There were two types of early nests before the emergence of workers, one with a worker number of less than 63 and the other of more than 231. The former were new nests, each containing a new queen and were built by reproductive swarming, whereas each of the latter was possibly re-built by an old queen and an absconding swarm. Ten swarming colonies were attracted by an oil-lamp light; these consisted of a copulated queen with immature oocytes in the ovary and 55 workers on average. No males, or workers stylopized by Xenos provesparum were observed either in colonies before worker emergence or in swarms coming flying into the light. Differences between swarming in Provespa, polygyny and nest relocation in Vespa, and those in swarm-founding Polistinae are discussed.     

Collective exploration and area marking in the ant Lasius niger

Summary: The aphid tending ant species, Lasius niger collectively explore new areas. Scouts are mobilised within the first five minutes of exploration with activity thereafter decreasing and stabilising after 40 minutes. Collective path choices of colonies underline the existence of chemical recruitment and area marking during exploration. To identify the territorial or home-range role of exploratory marking, we examined its influence on agonistic behaviour during intraspecific encounters. Ten nestmates and ten alien conspecifics were confronted: (i) on a paper marked by their own colony, (ii) on a paper marked by the alien colony or (iii) on a paper not chemically marked by ants. Residents and intruders showed a weak level of aggression and behaved similarly whatever the type of area marking. As no colony-specificity was observed in the odds of eliciting aggression, exploratory area marking in Lasius niger seems to be rather a home-range than a true territorial marking. We discuss the role of collective exploration, home-range marking and intraspecific interactions in the structuring and use of foraging space by neighbour Lasius niger colonies.     

Effects of logging and recruitment on community phylogenetic structure in 32 permanent forest plots of Kampong Thom,Cambodia

Ecological communities including tropical rainforest are rapidly changing under various disturbances caused by increasing human activities. Recently in Cambodia, illegal logging and clear-felling for agriculture have been increasing. Here, we study the effects of logging, mortality and recruitment of plot trees on phylogenetic community structure in 32 plots in Kampong Thom, Cambodia. Each plot was 0.25 ha; 28 plots were established in primary evergreen forests and four were established in secondary dry deciduous forests. Measurements were made in 1998, 2000, 2004 and 2010, and logging, recruitment and mortality of each tree were recorded. We estimated phylogeny using rbcL and matK gene sequences and quantified phylogenetic α and β diversity. Within communities, logging decreased phylogenetic diversity, and increased overall phylogenetic clustering and terminal phylogenetic evenness. Between communities, logging increased phylogenetic similarity between evergreen and deciduous plots. On the other hand, recruitment had opposite effects both within and between communities. The observed patterns can be explained by environmental homogenization under logging. Logging is biased to particular species and larger diameter at breast height, and forest patrol has been effective in decreasing logging.     

Species Composition and Inter-annual Dynamics of a Freshwater Tidal Plant Community Following Removal of the Invasive Grass, Phragmites australis

We document the regeneration of native freshwater wetland plant assemblages following removal of the common reed, Phragmites australis (Cav.) Trin. ex Steudel from two sites at Chapman Pond, East Haddam, Connecticut, USA. We gathered field data on composition of the vegetation 1 year before and for each of the 3 years after the removal in fall 1995/spring 1996 of Phragmites by two slightly different methods (hand-removal and herbiciding in one area, mowing/mulching and herbiciding in another). An area where Phragmites was left intact was similarly monitored. Our goals for this monitoring were: (1) to document plant species composition and richness before and after Phragmites removal and (2) to examine temporal and spatial variability in patterns of plant recruitment. Phragmites declined in both density and extent in both plots where removal treatments were applied. Richness, evenness, and density of non-Phragmites species increased steadily from 1996 to 1997 in all removal and intact plots. However, the species composition of the removal plots was richer than that of the intact plot, and more closely resembled that of comparable, uninvaded freshwater tidal wetlands. Rates of recovery of species richness in the removal plots declined from 1997 to 1998, potentially reflecting saturation of available colonization space, or the return of Phragmites. Phragmites has expanded its range in both of the removal plots since 1997. A model of its colonization indicates that Phragmites occupies space through localized proliferation of dense rhizomes rather than diffusely foraging with long tillers. Vigilance in monitoring is needed to document the spread of invasives, to evaluate the multi-faceted ecological effects of eradication efforts on both the invader and the regenerating community, and develop strategies for preventing re-invasion.     

Population ecology of Acacia tortilis in the semi-arid region of the Sudan

Abstract. The population ecology of Acacia tortilis, one of the important woody species in the arid and semi-arid regions of northern Africa, was studied in the Sudan: Ten sample plots were established and all individuals in the plots were mapped and their basal diameter, stem height, crown height and cover measured annually for three years. Individuals were grouped into different height-classes, ‘life-forms’: trees >1.65 m, shrubs > 0.65 m and < 1.65 m, saplings < 0.65 m and seedlings (saplings less than one year old). The population density was 510 ind./ ha in 1985, 432 ind. / ha in 1986 and 218 ind. / ha in 1987. The life-form distribution of the population showed clear differences between plots and years. Mortality in Acacia tortilis, expressed as loss in individuals per year, amounted to 167 ind. / ha / yr. Drought and lethal cutting were found to be the main mortality factors. The population declined by 15.3 % from 1985 to 1986 and by 49.5 % from 1986 to 1987. Natality (recruitment), expressed as number of newly established saplings, was 13 ind. / ha / yr. Recruitment is apparently discontinuous and bound to years with a relatively high precipitation.     

Hatching date influences age at first reproduction in the black-headed gull

In long-lived colonial birds, age at recruitment is an important life-history character. Variation in this parameter may reflect differences in several factors, including competitive ability and breeding strategies. Further, these differences may be due to timing of hatching (for instance through differences in competitive ability). We investigated the age of first-time breeders in relation to hatching date in a black-headed gull Larus ridibundus colony situated in central France, from 1979 to 1993. Age at first breeding was estimated for four groups of individuals (total n=550) according to their hatching date, using a recent capture-recapture methodology which allowed us to estimate recruitment rate without the limiting assumptions of methods relying on simple return rates. The age at first breeding was negatively correlated with the hatching date of individuals: individuals hatched earlier in the season started breeding at a younger age than individuals born later. Proportionally more 2-year-old late-hatched individuals were seen breeding on small peripheral colonies than young early-hatched individuals. This difference disappeared after age 3 years. These results strongly suggest that individuals hatched late in the season start to breed on peripheral colonies before recruiting to their natal colony. A difference of few weeks in hatching date has consequences which can last for several years.     

Variability of seedling recruitment under dominant, moss, and litter removal over four years

In a four-year removal experiment we investigated the effect of removing a dominant species (Nardus stricta), the litter layer and moss layer on species composition of established vegetation and on numbers and species composition of seedlings. The experiment was conducted in an oligotrophic wet meadow (Molinion with some features ofViolion caninae according to phytosociological classification). After four years of the experiment, the redundancy analysis (RDA) did not reveal any changes in the composition of the established plant community. Seedling numbers and their species composition, however, varied significantly both among treatments and over years. Whereas the differences between treatments were mainly in total seedling numbers, the years differed in both seedling numbers and relative representation of individual species. In particular,Myosotis nemorosa seedlings highly increased their abundance in a favourable year 1995 constituting one third of all the seedlings found, whereas in other years their abundance was between 4% and 10%. The effects of year and treatment were not additive (significant year × treatment interaction). This means that plots under different treatments are differently affected by climatic conditions of individual years. The greatest increase of seedling recruitment was observed in the plots with the moss layer removed. Microsites with high seedling recruitment persisted in the plots for no more than two years. Why the established vegetation composition does not differ among treatments even after four years, despite the relatively pronounced and immediate response of seedlings? The number of species with regular seedling recruitment is limited. They are mostly perennials, and consequently, it might take even longer than four years before recruitment limitation affects the population size of established plants.     

The relationship between the distribution of worker sizes and new worker production in the ant Formica neorufibarbis

While it is commonly assumed that variation in worker sizes within a single ant colony increases colony efficiency, there is little causal evidence of a link between worker size variation and colony performance. I tested whether the range of worker sizes within colonies of the ant species Formica neorufibarbis affected new worker production. Removing large workers from colonies lowered the rate of new worker production. A study of unmanipulated colonies indicated that colonies did not maintain a full range of worker sizes; mean worker head widths varied from 0.89-1.24 mm. Colonies naturally missing large workers did not have lower rates of worker production, suggesting that the relative size, not the absolute size, of workers within colonies was important. These are the first results to directly link the range of worker sizes to a component of colony fitness in a natural setting.     

Spatial distribution, size and density of nests of Myrmicaria opaciventris Emery (Formicidae, Myrmicinae)

Summary: We recorded over the course of one year the location of nests of Myrmicaria opaciventris on five sites in the region surrounding Yaoundé (Cameroon). On these sites, the colonies are polydomous and polygynous. The size of the largest colony, which was made up of 27 nests, was evaluated at 216,575 - 24,750 individuals of which 147,230 - 15,336 were workers. The nests are interconnected by trails that the workers dig during the rainy season. They consist of trenches that are slowly closed to form tunnels. Thanks to these tunnels, the workers are sheltered from their predators when going from one nest to another. They also have easy access to food sources far from the nests. We noted a seasonal variation in the location of the nests. It consists of a kind of budding. Depending on the case, the initial nest might survive or end up by being abandoned during the main rainy season or early in the dry season. The spatial distribution of the nests in the field is, in general, random and the density is lower during the dry season (from 6.2 to 26.4 nests/ha) than during the rainy season (from 12.1 to 50 nests/ha).     

Dispersal and recruitment during population growth in a colonial bird, the great cormorant Phalacrocorax carbo sinensis

While the factors influencing reproduction and survival in colonial populations are relatively well studied, factors involved in dispersal and settlement decisions are not well understood. The present study investigated exchanges of great cormorants Phalacrocorax carbo sinensis among six breeding colonies over a 13‐year period when the breeding population in Denmark increased from 2800 to 36 400 nests. We used a multistate capture‐recapture model that combined multisite resightings and recoveries to examine simultaneously recruitment, natal dispersal, breeding dispersal and annual survival of first‐year, immature and breeding great cormorants. Mean survival of first‐year birds (0.50±0.09, range=0.42–0.66 among colonies) was lower than survival of breeders (0.90±0.06, range=0.81–0.97). Mean survival of immature birds over the study period was 0.87±0.08. Dispersal from a colony increased with decreasing mean brood size in the colony in both first‐time and experienced breeders. The choice of the settlement colony in first‐time breeders was affected by conditions in the natal colony and in the colonies prospected during the pre‐breeding years. In particular, first‐time breeders recruited to colonies where they could expect better breeding success. Experienced breeders relied mainly on cues present early in the season and on their own breeding experience to choose a new breeding colony. Newly established colonies resulted mainly from the immigration of first‐time breeders originating from denser colonies. Dispersal was distance‐dependent and first‐time breeders dispersed longer distances than breeders. We suggest that the prospecting behaviour allows first‐time breeders to recruit in nearby as well as more distant potential breeding colonies. Dispersing breeders preferred to settle in neighbouring colonies likely to benefit from their experience with foraging areas. We discuss the importance of these movements for growth and expansion of the breeding population.     

Invasive alien plants affect grassland ant communities, colony size and foraging behaviour

Ants are dominant members of many terrestrial ecosystems and are regarded as indicators of environmental changes. However, little is known about the effects of invasive alien plants on ant populations, particularly as regards the density, spatial distribution and size of ant colonies, as well as their foraging behaviour. We addressed these questions in a study of grassland ant communities on five grasslands invaded by alien goldenrods (Solidago sp.) and on five non-invaded grasslands without this plant. In each grassland, seven 100 m² plots were selected and the ant colonies counted. Ant species richness and colony density was lower in the plots on the invaded grasslands. Moreover, both of these traits were higher in the plots near the grassland edge and with a higher number of plant species in the grasslands invaded by goldenrods but not in the non-invaded ones. On average, ant colony size was lower on the invaded grasslands than the non-invaded ones. Also, ant workers travelled for longer distances to collect food items in the invaded areas than they did in the non-invaded ones, even after the experimental removal of some ant colonies in order to exclude the effect of higher colony density in the latter. Our results indicate that invasive alien goldenrods have a profound negative effect on grassland ant communities which may lead to a cascade effect on the whole grassland ecosystem through modification of the interactions among species. The invasion diminishes a major index of the fitness of ants, which is a colony’s size, and probably leads to increased foraging effort of workers. This, in turn, may have important consequences for the division of labour and reproductive strategies within ant colonies.