Is two company or a crowd: How does conspecific density affect the small-scale dispersal of a weed biocontrol agent?

To predict the growth and spread of an insect population introduced for the biological control of weeds, one must first understand the factors affecting the movement of individuals in the population. The purpose of this study was to determine how the dispersal rate of Aphthona lacertosa (Rosenhauer) (Chrysomelidae) was affected by conspecific density and by the characteristics of leafy spurge (Euphorbia esula L.: Euphorbiaceae) in patches where these beetles feed. In 2002 in Manitoba and in 2003 in Alberta, Canada, between 200 and 2500 insects were released in small patches (<10 m²) of spurge. The number and location of beetles within patches was monitored over subsequent days. In 1 m² plots within patches, spurge ramet density, the proportions of vegetative and reproductive ramets, and ramet height were measured. In both years, beetle movement within patches and emigration from patches, was not affected by conspecific density. In Manitoba in 2002, beetles aggregated non-randomly on either vegetative or reproductive ramets within plots, but plot characteristics were not related to the formation of aggregations. In Alberta in 2003, plots in which beetles aggregated had significantly higher spurge density but did not differ in either the proportion of vegetative ramets or in the amount of non-spurge vegetation. These results suggest that density-dependent dispersal does not limit the population's ability to reach densities up to 2500 beetles/m².     

Phenology of first and peak emergence of Aphthona lacertosa and A. nigriscutis: Two flea beetles introduced for biological control of leafy spurge, Euphorbia esula L.

Nonlinear models were used to estimate first emergence and peak abundance dates for Aphthona lacertosa Rosenhauer and A. nigriscutis Foudras, two flea beetles introduced to control leafy spurge, Euphorbia esula L., in North America. For model development, 26 field sites were sampled for flea beetle abundance at weekly intervals for eight weeks in three western Minnesota counties in 2000, 2001, and 2002. A three-parameter Weibull function, fit to observed cumulative probability distributions, were used to predict accumulated degree-days (ADD) to first emergence. Bias testing indicated the Weibull function provided a useful estimate of first emergence for A. lacertosa (304 ADD, lower developmental threshold 7.5 °C), but failed to produce a useful estimate for A. nigriscutis. A third-order polynomial was used to approximate seasonal abundance and predict peak abundance for each species. Estimated ADD to peak abundance of A. lacertosa was 594 ± 24 (DD > 7.5 °C) and 670 ± 15 (DD > 9.3 °C) for A. nigriscutis. Models were validated with additional data sets from Minnesota, Montana, and North Dakota. Estimated date of peak emergence provided useful predictions of peak emergence for Minnesota and North Dakota, but failed to predict peak emergence in Montana. We speculate that variation in climate and environmental conditions between Midwestern states and Montana were responsible for differing emergence patterns. We conclude that phenology models should be developed regionally to provide useful predictions of peak emergence for land managers. Maps were developed for Minnesota to spatially display predicted dates of peak abundance for A. lacertosa and A. nigriscutis.     

Evidence for the influence of conspecific chemical cues on <Emphasis Type="Italic">Aphthona nigriscutis</Emphasis> (Coleoptera: Chrysomelidae) behaviour and distribution

Although the distribution of biological control agents may have a significant effect upon their impacts, the mechanisms regulating these distributions are often unknown. Such is the case with Aphthona nigriscutis, a classical biological control agent of leafy spurge in North America. These beetles assume aggregated distributions at some sites but disperse rapidly at others. The potential influence of plant and insect-factors upon aggregation and dispersal was investigated to try to explain these observations. Male beetles produce a putative aggregation pheromone. Responses of conspecifics to male-associated cues are greater when beetles are feeding on host plants. Densities of beetle groups greatly impact their attractiveness. Males are more sensitive to dispersal cues and females are more sensitive to congregation cues.     

Temporal dynamics of leafy spurge (Euphorbia esula) and two species of flea beetles (Aphthona spp.) used as biological control agents

The goal of this study was to evaluate the biological control program of leafy spurge (Euphorbia esula) in a large natural area, Theodore Roosevelt National Park, western North Dakota, USA. Aphthona lacertosa and Aphthona nigriscutis have been released at more than 1800 points in the 18,600-ha South Unit of the park beginning in 1989; most releases have occurred since 1994. We established permanent vegetation plots throughout the infested area of the park and determined stem counts and biomass of leafy spurge and abundance of the two flea beetle species at these plots each year from 1999 to 2001. Both biomass and stem counts declined over the 3 years of the study. Both species of flea beetle are well established within the park and have expanded into areas where they were not released. A. nigriscutis was more abundant than A. lacertosa in the grassland areas we surveyed, but in all other habitats abundances were similar. Using structural equation models, only A. lacertosa could be shown to have a significant effect on counts of mature stems of leafy spurge. A. nigriscutis numbers were positively correlated with stem counts of mature stems. Previous year’s stem counts had the greatest influence on change in stem counts over each 2-year time step examined with structural equation models.     

Effect of Leafy Spurge (Euphorbia esula) Genotype on Feeding Damage and Reproduction of Aphthona spp.: Implications for Biological Weed Control

North American leafy spurge (Euphorbia esula L.) is genetically diverse and composed of multiple genotypes introduced from several areas of Europe and Asia. Five species of leafy spurge flea beetle (Aphthona spp.) have been introduced as biological control agents for leafy spurge, but were collected in a relatively small region of Europe. Greenhouse and field experiments were conducted to determine if observed variation in feeding preference and reproduction of Aphthona spp. on North American leafy spurge may be due in part to leafy spurge genotype. Leafy spurge genotypes were collected from Austria; Manitoba in Canada; and the states of Montana, Nebraska, North Dakota, South Dakota, and Wyoming in the United States. Leafy spurge genotype affected feeding but not egg laying by Aphthona spp. adults. Aphthona czwalinae/lacertosa fed slightly less in a free-choice test on a genotype from Manitoba (7%) compared to genotypes from Nebraska and North Dakota (14%). Aphthona flava tended to feed less on the Nebraska genotype than any other genotype evaluated. Reproduction of Aphthona spp. was greatly affected by leafy spurge genotype. For instance, A. czwalinae/lacertosa produced 72 adults per plant from a Nebraska genotype compared to 11 to 32 adults per plant from all other genotypes evaluated. Thus, some observed variation in establishment and reproduction of Aphthona spp. can be attributed to leafy spurge genotype.     

First Quantitative Study of Rove Beetles in Oklahoma Winter Wheat Fields

Adult rove beetles (Staphylinidae) were sampled every 7–14 days from one winter wheat field located in each of the four major wheat growing regions of Oklahoma during the 1999–2000 and 2000–2001 growing seasons. The number of cereal aphids per tiller, wheat plant growth stage, and wheat tiller density also were estimated. A total of 12 genera representing 13 species of beetles were collected from the field. The density of rove beetles was generally low, ranging from 0.003 beetles per m² in fall to 0.106 beetles per m² in spring. Rove beetle communities differed among seasons. After accounting for the effect of season, there was no statistically significant association between rove beetle community structure and field location, aphid density, wheat plant growth stage, or wheat plant density. Most rove beetle species showed no association with a particular season, however, Aleochara notula Erichson, Lathrobium sp., and Oxypoda sp. were present predominantly in fall, while Bisnius inquitus Erichson was associated with winter. Oxypoda sp. was the most abundant rove beetle in winter wheat fields in spring and was relatively abundant in winter, but was not collected from wheat fields in fall. Tachyporus jocosus Say was present in wheat fields during all seasons. T. jocosus was the most abundant rove beetle species in the winter wheat fields in fall and winter and was the second most abundant species during spring.     

Confinement Behavior of Cape Honey Bees (Apis mellifera capensis Esch.) in Relation to Population Densities of Small Hive Beetles (Aethina tumida Murray)

We quantified the effects of increasing small hive beetle (Aethina tumida Murray) populations on guarding behavior of Cape honey bees (Apis mellifera capensis, an African subspecies). We found more confinement sites (prisons) at the higher (50 beetles per colony) rather than lower (25 beetles per colony) beetle density. The number of beetles per prison did not change with beetle density. There were more guard bees per beetle during evening than morning. Neither guard bee nor beetle behavior varied with beetle density or over time. Forty-six percent of all beetles were found among the combs at the low beetle density and this increased to 58% at the higher one. In neither instance were beetles causing depredation to host colonies. Within the limits of the experiment, guarding behavior of Cape honey bees is relatively unaffected by increasing beetle density (even if significant proportions of beetles reach the combs).     

Integration of biological control agents with other weed management technologies: Successes from the leafy spurge (Euphorbia esula) IPM program

An invasive weed can occupy a variety of environments and ecological niches and generally no single control method can be used across all areas the weed is found. Biological control agents integrated with other methods can increase and/or improve site-specific weed control, but such combinatorial approaches have not been widely utilized. The successful leafy spurge (Euphorbia esula L.) control program provides examples for future integrated weed programs that utilize biological control agents with traditional methods. Weed control methods can be used separately, such as when the leafy spurge gall midge (Spurgia esulae Gagné) reduced seed production in wooded areas while herbicides prevented further spread outside the tree line. Traditional methods also can be used directly with biological control agents. Incorporation of Aphthona spp. with herbicides has resulted in more rapid and complete leafy spurge control than either method used alone. Also, the insect population often increased rapidly following herbicide treatment, especially in areas where Aphthona spp. were established for several years but had been ineffective. Incorporation of Aphthona spp. with sheep or goat grazing has resulted in a larger decline in leafy spurge production than insects alone and in weed density than grazing alone. Controlled burns can aid establishment of biological control agents in marginally suitable environments, but timing of the fire must be coordinated to the insect’s life-cycle to ensure survival. Integration of biological control agents with revegetation programs required the agent to be the last method introduced because the cultivation and herbicide treatments necessary to establish desirable grasses and forbs were destructive to the insect. In a practical application, herbicides were combined with Aphthona spp. to help the insect establish and control leafy spurge in the habitat of the western prairie fringed orchid (Platanthera praeclara Sheviak and Bowles), an endangered species. Several experimental designs can be used to evaluate biological control agents with cultural, mechanical, and chemical control methods or with additional biological agents.     

Effectiveness of honey bees in delivering the biocontrol agent Bacillus subtilis to blueberry flowers to suppress mummy berry disease

Honey bees are important pollinators of commercial blueberries in the southeastern United States, and blueberry producers often use supplemental bees to achieve adequate fruit set. However, honey bees also vector the plant pathogenic fungus Monilinia vaccinii-corymbosi which infects open blueberry flowers through the gynoecial pathway causing mummy berry disease. Here, we report the results of a 3-year field study to test the hypothesis that using bee hives equipped with dispensers containing the biocontrol product Serenade, a commercial formulation of the bacterium Bacillus subtilis which has shown activity against flower infection by M. vaccinii-corymbosi in laboratory experiments, can reduce mummy berry disease incidence when honey bees are used as pollinators in blueberries. Individual honey bees carried 5.1–6.4 × 10⁵ colony-forming units (CFU) of B. subtilis when exiting hive-mounted dispensers with Serenade. On caged rabbiteye blueberry bushes in the field, population densities of B. subtilis vectored by honey bees reached a carrying capacity of <10³ CFU per flower stigma within 2 days of exposure, and there was a highly significant non-linear relationship between B. subtilis populations per stigma and bee activity, expressed as number of legitimate flower visits per time interval per cage (R = 0.6928, P < 0.0001, n = 32). Honey bee density (1600 or 6400 individuals per 5.8-m³ cage) and Serenade treatment (presence or absence of the product in hive-mounted dispensers) significantly (P < 0.05) affected the incidence of fruit mummification on caged bushes, whereby increasing bee density increased disease incidence and application of Serenade reduced disease levels. Taken together, results of this study suggest that use of a hive-dispersed biocontrol product such as Serenade as a supplement during pollination can reduce the risk of mummy berry disease. This may be a prudent practice that optimizes the benefits to pollination of high bee densities while reducing the associated disease-vectoring risk.     

Attack behavior and host utilization ofAphthona chinchihi (Col.: Chrysomelidae), a potential biological control agent ofEuphorbia esula (Euphorbiaceae, leafy spurge) in North America

Aphthona chinchihi Chen was collected in China feeding on leafy spurge (Euphorbia esula L.). Studies were conducted on its host specificity in the laboratory, using field collected adults and their progeny.Aphthona chinchihi can effectively complement the impact of the other natural enemies of leafy spurge established from Europe in the U.S.A. and Canada. The adults feed on leaves and shoots and the larvae, which cause the main damage to the plant, feed on the hypogeous portion of the plant, seriously stressing the plant and preventing its vegetative spread. The host range ofA. chinchihi was studied with tests on adult feeding and oviposition, larval survival and host suitability, using 40 plant species or varieties distributed in 12 families. The experiments demonstrated that it has a high level of specificity. This flea beetle completed its life cycle only on leafy spurge. Also, because of its ecological valence,A. chinchihi has a very good potential as a biocontrol agent in North America.     

Horizontal transmission of a microsporidium from the convergent lady beetle, Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae), to three coccinellid species of Nova Scotia

Convergent lady beetles, Hippodamia convergens Guérin-Méneville, are a popular choice for aphid control in North America. An unidentified microsporidium was found in H. convergens adults that were purchased from a commercial insectary in 2004. This study examined egg cannibalism and egg predation as a means of horizontal transmission of the unidentified microsporidium among H. convergens larvae and three coccinellid species found in Nova Scotia: Coccinella septempunctata (seven-spotted lady beetle), C. trifasciata perplexa (three-banded lady beetle), and Harmonia axyridis (multicolored Asian lady beetle). The microsporidium was transmitted with 100% efficiency when first instars fed on microsporidia-infected eggs. Mean spore count data from smear preparations of infected beetles suggest that the infection was as heavy in C. trifasciata perplexa (a native coccinellid) (11.2 ± 0.96 spores/100 μm²) as it was in H. convergens (the natural host) (12.8 ± 1.16) but lighter in the introduced species C. septempunctata (7.5 ± 0.65) and H. axyridis (0.8 ± 0.11). For all of the beetle species examined, larval development was significantly longer for microsporidia-infected individuals than for their uninfected cohorts. The microsporidium had no effect on larval mortality. Based on the results of this study, field-collected H. convergens should be examined for microsporidia and uninfected individuals should be used to rear individuals for release in biological control programs. However, this is unlikely to happen because H. convergens are relatively easy and inexpensive to collect from their overwintering sites for redistribution.     

Social encapsulation of the small hive beetle (<Emphasis Type="Italic">Aethina tumida</Emphasis> Murray) by European honeybees (<Emphasis Type="Italic">Apis mellifera</Emphasis> L.)

Summary European and African subspecies of honeybees (Apis mellifera L.) utilize social encapsulation to contain the small hive beetle (Aethina tumida Murray), a honeybee colony scavenger. Using social encapsulation, African honeybees successfully limit beetle reproduction that can devastate host colonies. In sharp contrast, European honeybees often fail to contain beetles, possibly because their social encapsulation skills may be less developed than those of African honeybees. In this study, we quantify beetle and European honeybee behaviours associated with social encapsulation, describe colony and time (morning and evening) differences in these behaviours (to identify possible circadian rhythms), and detail intra-colonial, encapsulated beetle distributions. The data help explain the susceptibility of European honeybees to depredation by small hive beetles. There were significant colony differences in a number of social encapsulation behaviours (the number of beetle prisons and beetles per prison, and the proportion of prison guard bees biting at encapsulated beetles) suggesting that successful encapsulation of beetles by European bees varies between colonies. We also found evidence for the existence of circadian rhythms in small hive beetles, as they were more active in the evening rather than morning. In response to increased beetle activity during the evening, there was an increase in the number of prison guard bees during evening. Additionally, the bees successfully kept most (~93%) beetles out of the combs at all times, suggesting that social encapsulation by European honeybees is sufficient to control small populations of beetles (as seen in this study) but may ultimately fail if beetle populations are high.Received 20 January 2003; revised 21 April 2003; accepted 29 April 2003.     

Biological control ofSenecio jacobaea in Northern California,an enduring success

Senecio jacobaea, a poisonous weed from Eurasia, was brought under successful biological control in the Ft. Bragg, California area by 1976, through the combined action of the defoliating cinnabar moth (Tyria jacobaeae) and a root feeding flea beetle (Longitarsus jacobaeae). In 1987, 4 previously infested Ft. Bragg sites (3 sites where control had been documented and another unstudied site) were examined.Senecio jacobaea densities at these sites were 0.0, 0.0, 0.01 and 0.18 plants/m², indicating both continued and improved control of the weed. The flea beetle and the cinnabar moth both persist at the sites, despite very low numbers ofS. jacobaea plants. The control ofS. jacobaea has resulted in the return of near natural vegetation at the 2 coastal prairie sites and regained productivity at the 2 pasture sites. c/o American Embassy APO San Francisco 96301     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Supercolonies of the invasive yellow crazy ant, Anoplolepis gracilipes, on an oceanic island: Forager activity patterns, density and biomass

Summary. Ants have the capacity to reach unusually high densities, mostly in their introduced ranges. Numerical dominance is often cited as key to the ability of exotic ants to displace native ant species, reduce the abundance of invertebrates and negatively impact upon bird, land crab and other vertebrate populations. On Christmas Island, Indian Ocean, the yellow crazy ant, Anoplolepis gracilipes (Jerdon), forms supercolonies, where extremely high densities of foraging ants have contributed to ‘invasional meltdown’ in rainforest areas. Densities of up to 2254 foraging ants per m² and a biomass of 1.85 g per m² were recorded, and nest densities reached 10.5 nest entrances per m². Populations of A. gracilipes can overcome and kill red endemic land crabs (Gecarcoidea natalis) over 100 times their own biomass. This is the highest recorded density of foraging ants, and adds another element to the definition of ‘supercolony’ of unicolonial ants. This paper documents one extreme in a continuum of densities of unicolonial, invasive ant species and highlights the need to incorporate forager densities into invasive ant research.Received 17 November 2004; revised 14 February 2005, accepted 21 February 2005.     

塔克拉玛干沙漠北缘柽柳灌丛沙堆对甲虫的庇护作用

以塔克拉玛干沙漠北缘典型荒漠区柽柳灌丛沙堆-柽柳包为研究对象,在2005—2006年期间调查了柽柳包及相邻沙漠裸地地表甲虫多样性。结果表明:2005年和2006年柳包上地表甲虫个体数量和物种数均高于沙漠裸地,柽柳包上的地表甲虫物种数分别为21种和16种,沙漠裸地中分别为10种和12种,柽柳包上物种丰富度明显高于沙漠裸地。选取优势种髋胫小土甲(Penthicicus koltzei Reitter)进行相对种群密度的估算,髋胫小土甲在柽柳包上分布数量较多,密度可达到每平方米在13头以上;但沙漠裸地仅有6头左右。柽柳包对甲虫多样性有明显的庇护所的作用,有利于维持甲虫物种多样性。2种不同生境条件下地表甲虫的数量存在明显的季节变动,其数量高峰分别出现在2005年6月和2006年7月,可能与不同季节环境的极端性(如气温峰值和洪水早晚)导致的食物丰富程度有关。     

Open field host selection and behavior by tamarisk beetles (Diorhabda spp.) (Coleoptera: Chrysomelidae) in biological control of exotic saltcedars (Tamarix spp.) and risks to non-target athel (T. aphylla) and native Frankenia spp.

Biological control of invasive saltcedars (Tamarix spp.) in the western U.S. by exotic tamarisk leaf beetles, Diorhabda spp., first released in 2001 after 15 years of development, has been successful. In Texas, beetles from Crete, Greece were first released in 2004 and are providing control. However, adults alight, feed and oviposit on athel (Tamarix aphylla), an evergreen tree used for shade and as a windbreak in the southwestern U.S. and México, and occasionally feed on native Frankenia spp. plants. The ability of tamarisk beetles to establish on these potential field hosts was investigated in the field. In no-choice tests in bagged branches, beetle species from Crete and Sfax, Tunisia produced 30–45% as many egg masses and 40–60% as many larvae on athel as on saltcedar. In uncaged choice tests in south Texas, adult, egg mass and larval densities were 10-fold higher on saltcedar than on adjacent athel trees after 2 weeks, and damage by the beetles was 2- to 10-fold greater on saltcedar. At a site near Big Spring, in west-central Texas, adults, egg masses and 1st and 2nd instar larvae were 2- to 8-fold more abundant on saltcedar than on athel planted within a mature saltcedar stand being defoliated by Crete beetles, and beetles were 200-fold or less abundant or not found at all on Frankenia. At a site near Lovelock, Nevada, damage by beetles of a species collected from Fukang, China was 12–78% higher on saltcedar than on athel planted among mature saltcedar trees undergoing defoliation. The results demonstrate that 50–90% reduced oviposition on athel and beetle dispersal patterns within resident saltcedar limit the ability of Diorhabda spp. to establish populations and have impact on athel in the field.     

Ecological and genetic data indicate recovery of the endangered coral Acropora palmata in Los Roques, Southern Caribbean

The rapid decline of Acropora cervicornis and Acropora palmata has often been linked with coral reef deterioration in the Caribbean; yet, it remains controversial whether these species are currently recovering or still declining. In this study, the status of ten populations of A. palmata in Los Roques National Park (LRNP), Venezuela is presented. Six of these populations showed signs of recovery. Ten 80 m² belt-transects were surveyed at each of the ten reef sites. Within belt-transects, each colony was measured (maximum diameter and height) and its status (healthy, diseased or injured) was recorded. Populations in recovery were defined by a dominance of small to medium-sized colonies in densities >1 colony per 10 m², together with 75% undamaged colonies, a low prevalence of diseases (<10%), and a low density of predators (0.25 snails per colony). Based on allozyme analysis of seven polymorphic loci in four populations (N = 30), a moderate to high-genetic connectivity among these populations (F _ST = 0.048) was found with a predominance of sexual over asexual reproduction (N* : N = 1; N _go : N = 0.93–1). Both ecological and molecular data support a good prognosis for the recovery of this species in Los Roques.     

Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia)

Little is known about how tropical land-use systems contribute to the conservation of functionally important insect groups, including dung beetles. In a study at the margin of Lore Lindu National Park (a biodiversity hotspot in Central Sulawesi, Indonesia) dung-beetle communities were sampled in natural forest, young secondary forest, agroforestry systems (cacao plantations with shade trees) and annual cultures (maize fields), each with four replicates (n = 16 sites). At each site we used 10 pitfall traps, baited with cattle dung, along a 100 m transect for six 3-day periods. The number of trapped specimens and species richness at the natural forest sites was higher than in all land-use systems, which did not significantly differ. Each land-use system contained, on average, 75% of the species richness of the natural forest, thereby indicating their importance for conservation. However, a two-dimensional scaling plot based on NESS indices (m = 6) indicated distinct dung beetle communities for both forest types, while agroforestry systems and annual cultures exhibited a pronounced overlap. Mean body size of dung beetles was not significantly influenced by land-use intensity. Five of the six most abundant dung beetle species were recorded in all habitats, whereas the abundance of five other species was significantly related to habitat type. Mean local abundance and number of occupied sites were closely correlated, further indicating little habitat specialisation. The low dung beetle diversity (total of 18 recorded species) may be due to the absence of larger mammals in Sulawesi during historical times, even though Sulawesi is the largest island of Wallacea. In conclusion, the dung beetle fauna of the lower montane forest zone in Central Sulawesi appears to be relatively robust to man-made habitat changes and the majority of species did not exhibit strong habitat preferences.     

Bigger ant colonies reduce herbivory and herbivore residence time on leaves of an ant-plant: Azteca muelleri vs. Coelomera ruficornis on Cecropia pachystachya

Summary The effect of defence force size in colonies of the ant Azteca muelleri on the time spent to localize, attack and expel the specialized herbivorous beetle Coelomera ruficornis from Cecropia pachystachya bushes was studied in an area of Atlantic forest in northeastern Brazil. Our results show that Azteca muelleri expel Coelomera ruficornis from Cecropia pachystachya and that the number of ants leaving a colony (defence force size) is negatively correlated with the residence time of an adult beetle on the plant. Colonies with larger defence forces recruited larger numbers of ants, resulting in faster herbivore discovery (r ²=0.80; n=17; P<0.001) and reduced herbivore residence time on a leaf (r ²=0.79 n=23; P<0.001) before being driven off by the ants. We also found a negative and significant relationship between herbivore damage on leaves and ant colony size (r ²=0.28; n=17; P<0.05). We conclude that larger colonies have more individuals available to patrol a plant and recruit defenders toward herbivores. This reduces the time spent to locate and expel susceptible herbivores from the plant. Since the plant probably benefits from reduced herbivory and the plant provides food for the ants, the association between Azteca muelleri and Cecropia pachystachya appears mutualistic.