Indirect effects of phorid fly parasitoids on the mechanisms of interspecific competition among ants

Indirect effects, which occur when the impact of one species upon another requires the existence of an intermediary species, are apparently very common and may be of greater magnitude than direct effects. Behaviorally mediated indirect effects occur when one species affects the behavior of a second, which in turn affects how that species interacts with a third. I studied behaviorally mediated indirect effects on the mechanisms of competition in two congeneric fire ant species in the presence and absence of parasitoid phorid flies, which parasitized only one ant species. In observational and experimental field studies, the presence of native Texas phorid flies in the genus Pseudacteon decreased food retrieval by their host, Solenopsis geminata (F.), by as much as 50%. In the presence of phorid flies, many S. geminata workers assumed a stationary, curled defensive posture and did not forage. Although the phorid parasitoids had a relatively large effect on exploitative competition, there was no measurable effect on interference competition. Fierce interspecific aggression was observed between S. geminata and S. invicta Buren, and the presence of phorids had no effect on the outcome of these contests. The indirect effects of Pseudacteon parasitoids on Solenopsis fire ant resource retrieval appear to be larger than the direct effect of mortality. Some aspects of the foraging behavior of these Solenopsis species may be, in part, evolutionary adaptations to phorid parasitoid pressure. Because of the relatively large indirect effects, South American Pseudacteon phorids may be promising biocontrol agents of imported fire ants, S. invicta, in the USA. In a laboratory study, a single South American Pseudacteon female was able to significantly decrease food retrieval rates of a North American population of the imported fire ant, S. invicta. Received: 11 May 1998 / Accepted: 18 April 1999     

Ecological interactions among ants in the genus Linepithema, their phorid parasitoids, and ant competitors

Abstract. 1. Many ant species abandon foraging and retreat underground when parasitoids in the dipteran family Phoridae are present. Although the influence of phorids on ant foraging is well documented, their influence on interspecific competition is less studied. This study examined whether phorids influenced the competitive ability of host ants in the genus Linepithema at two sites in Brazil.
2. The phorid Pseudacteon lontrae attacked Linepithema piliferum at one site, while the phorid Pseudacteon pusillus attacked an unknown Linepithema ( Linepithema sp.) at the other site. Phorid parasitoids of Linepithema were far more common than phorids of other ant species.
3. Despite a high abundance of phorids, it was difficult to conclude that they influenced competition. Captures in pitfall traps indicated that host Linepithema were most active during times of day when phorids were inactive.
4. Camponotus rufipes and Brachymyrmex sp., the most common competitors of Linepithema sp. (60% of all interactions), dominated Linepithema sp. during the day regardless of phorids. Remaining ant species could not be evaluated individually because they interacted with Linepithema sp. infrequently.
5. Ectatomma brunneum was the most common competitor of L. piliferum (58% of all interactions). The high abundance of phorids at this site made it impossible to evaluate interactions between E. brunneum and L. piliferum in the absence of phorids.
6. Phorids seldom influenced exploitative competition by causing host Linepithema to abandon the bait when no ant competitors were present.     

Parasite Lost: Chemical and Visual Cues Used by <Emphasis Type="BoldItalic">Pseudacteon</Emphasis> in Search of <Emphasis Type="BoldItalic">Azteca instabilis</Emphasis>

An undescribed species of phorid fly (genus: Pseudacteon) parasitizes the ant Azteca instabilis F Smith, by first locating these ants through the use of both chemical and visual cues. Experiments were performed in Chiapas, Mexico to examine a) the anatomical source of phorid attractants, b) the specific chemicals produced that attract phorids, and c) the nature of the visual cues used by phorids to locate the ants. We determined that phorid-attracting chemicals were present within the dorsal section of the abdomen, the location of the pygidial gland. Further experiments indicate that a pygidial gland compound, 1-acetyl-2-methylcyclopentane, is at least partially responsible for attracting phorid flies to their host. Finally, although visual cues such as movement were important for host location, size and color of objects did not influence the frequency with which phorids attacked moving targets.     

Biogeography and macroecology of phorid flies that attack fire ants in south-eastern Brazil and Argentina

Aim Saevissima group fire ants, Solenopsis richteri and S. invicta, have become serious pests when introduced from Argentina and Brazil to other continents. In South America, Solenopsis are distributed across a great variety of habitats and climates. In North America, S. invicta, introduced free of phorids, now ranges from coast to coast in the south. Success in introducing particular Pseudacteon as agents for the biological control of fire ants has varied across climatic zones. We aimed at assembling all the information about fire ant phorids from Argentina and Brazil, to estimate their richness and geographical ranges, to perform a climatic analysis for these distributions, to define groups and climate‐based communities, and to test and elucidate Rapoport's biogeographical rule. Location Argentina and Brazil (South America). Methods From field and museum collections and historical records, we developed a database of fire ant‐specific phorids throughout their known geographical range. A total of 123 sites with values for 15 climatic variables were mapped between 10° and 38° SL and between 35° and 65° C WL for the presence/absence of phorids. We calculated species richness across all sites combined, and for each phytogeographical region, using rarefaction curves, and ICE and Mmean estimators. We calculated mid‐latitudinal points, geographical ranges and areas for each species. The correlation between mid‐latitudinal point and ranges/areas was tested against a null model generated from the randomization of the raw distributional data. We used several types of multivariate analyses to distinguish groups of phorids by phytogeographical regions, hosts and climate, to find gradients of climate throughout the studied area, to define phorid communities in terms of their relationships with gradients of climate, and to test a mechanism for Rapoport's rule. Results Richness estimations using ICE and Mmean estimators were similar or higher than the observed values depending on the phytogeographical region. Cluster multivariate analyses based on climatic, phytogeographic and host data revealed distinct groupings of Pseudacton. The ‘cerrado’ group was confined to tropical savanna areas. A more ‘widespread’ group included ‘Chaco’ and ‘Maritime’ subgroups defined by their respective association with extreme temperatures or precipitation. Ordination multivariate analyses showed (1) two climatic gradients throughout the study area: one of temperature and the other of precipitation, and (2) that climatic variables significantly explained the observed assemblages of phorids. Positive and negative signs of the eigenvalues from the main axes of a canonical correspondence analysis allowed us to define eight communities whose geographical distribution resembled that of phytogeographical regions. We found a significant and positive correlation between geographical areas and mid latitudinal points, and furthermore, the Mantel test based on climatic variables suggested a mechanism for Rapoport's rule applying in the case of Pseudacteon. Main conclusions Pseudacteon species with greater mid‐latitudinal points occupy broader geographical areas and confront more stressful environmental conditions. Because the composition of Pseudacteon communities is largely determined by climatic variables, the correspondence between climates at sites of origin vs. sites of release should be an important consideration in choosing specific phorids for biocontrol efforts.     

Phorid parasitoids affect foraging activity of Solenopsis richteri under different availability of food in Argentina

1. In Argentina, six species of Pseudacteon parasitoids (Phoridae) attack Solenopsis richteri, one of the two species of South American fire ant that are exotic pests in North America. 2. The presence of these Pseudacteon species significantly reduces the number of ants at food resources in the field, as well as foraging activity generally. 3. Some Pseudacteon not only attack ants walking on trails or at feeding sites, but also at mound entrances, inhibiting workers from leaving to forage. 4. The average size of foraging ants (which prescribes their suitability as hosts) decreased in the presence of phorids. 5. The number of attacking phorids was correlated positively with the number of ants walking towards the food on the trail before the attack. 6. Solenopsis richteri workers responded to manipulations of food size and presence or absence of parasitoids in a risk-adjusting way, i.e. although more foragers were recruited to larger food items, attacking phorids reduced ant foraging activity by the same factor regardless of the size of the food offered. 7. The data suggest that S. richteri colonies juggle the needs to harvest food efficiently, reduce competition, and avoid excess risks from parasitoids in complex ways.     

Ecological dominance of the red imported fire ant, <Emphasis Type="Italic">Solenopsis invicta</Emphasis>, in its native range

Despite the widespread impacts invasive species can have in introduced populations, little is known about competitive mechanisms and dominance hierarchies between invaders and similar taxa in their native range. This study examines interactions between the red imported fire ant, Solenopsis invicta, and other above-ground foraging ants in two habitats in northeastern Argentina. A combination of pitfall traps and baits was used to characterize the ant communities, their dominance relationships, and to evaluate the effect of phorid flies on the interactions. Twenty-eight ant species coexisted with S. invicta in a gallery forest gap, whereas only ten coexisted with S. invicta in a xerophytic forest grassland. S. invicta was the most numerically dominant species in the richest and complex habitat (gallery forest); however it performed better as discoverer and dominator in the simpler habitat. S. invicta was active during day and night. In spite of its poor capacity to discover resources, S. invicta showed the highest ecological dominance and the second-best behavioral dominance after Camponotus blandus. S. invicta won 78% of the interactions with other ants, mostly against its most frequent competitor, Pheidole cf. obscurithorax, dominating baits via mass recruitment and chemical aggression. P. cf. obscurithorax was the best food discoverer. S. invicta won 80% of the scarce interactions with Linepithema humile. Crematogaster quadriformis was one of the fastest foragers and the only ant that won an equal number of contests against S. invicta. The low presence of phorid flies affected the foraging rate of S. invicta, but not the outcome of interspecific interactions. This study revealed that the red imported fire ant ecologically dominated other terrestrial ants in its native range; however, other species were able to be numerically dominant or co-dominant in its presence.     

The introduction of the fire ant parasitoid Pseudacteon nocens in North America: challenges when establishing small populations

Several species of parasitoid phorid flies (Pseudacteon spp., Diptera: Phoridae) have been released into the United States as potential biological control agents for the red imported fire ant, Solenopsis invicta Buren (Hymenoptera: Formicidae). Here we report the first successful introduction and spread of Pseudacteon nocens Borgmeier at a site in Texas, USA. Pseudacteon nocens is an important natural enemy since it is a widespread and often abundant parasitoid of S. invicta in Argentina, where it attacks larger fire ant workers eliciting a strong defensive response. Several years of effort to establish this species previously failed, and here we provide a model to better understand the likelihood of founding new populations when introducing sequential batches of flies in field or laboratory cultures. We also report on a novel method of establishing new populations of phorids in the field using pupae burial boxes to overcome constraints of releasing adult flies or infected worker ants.     

Parasitoid phorid flies of leaf‐cutting ants are negatively affected by loss of forest cover

Habitat fragmentation can have a high impact on parasitoid–ant interactions. Phorid flies are among the most important groups of natural enemies of leaf‐cutting ants. We studied the effects of loss in forest cover upon phorids of the leaf‐cutting ant Acromyrmex nigerSmith (Hymenoptera: Formicidae: Attini) in a fragmented area in the Southeastern Atlantic Forest, Brazil. We sampled 10 forest fragments, five large (>75 ha) and five small (<20 ha), as well as three areas of continuous forest (>1 000 ha). We marked 1–5 colonies of A. niger in the interior of each forest location. At each nest, we collected all of the phorids in interaction with the worker ants for a period of 15 min. We then collected ca. 200 worker ants, which we maintained in the laboratory for rearing phorids from them. We identified three phorid genera – Apocephalus, Myrmosicarius, and Neodohrniphora – which we both observed in the field and reared in the laboratory. The abundance and parasitism percentage were significantly greater in continuous forest sites than in forest fragments, whereas there were no significant differences between fragments of different sizes. These results provide further evidence for the effects of habitat size on the phorid‐Acromyrmex system in a tropical rain forest, based on the abundance of parasitoids both as adults in the field and as reared immature phorids in the laboratory.     

Post-Release Host-Specificity Testing of Pseudacteon tricuspis, a Phorid Parasitoid of Solenopsis Invicta Fire Ants

Inherent in any biological control program is the risk of nontarget effects. Pseudacteon tricuspisBorgmeier, a parasitoid phorid fly, has been introduced to the United States from South America as a potential biocontrol agent of the red imported fire ant, Solenopsis invictaBuren. We conducted tests of host specificity on introduced populations of P. tricuspis, which are attracted to alarm pheromones released by their hosts during events such as mound disturbances and interspecific interactions. We monitored disturbed mounds of S. invicta and its close congener, S. geminata(F.), during the expansion of P. tricuspis across north Florida and after populations had been established for ~3 years. We also tested host acceptance in established populations of P. tricuspis by offering trays containing S. invicta, S. geminata, and 14 additional ant species representing 12 different non-Solenopsis genera. Although P. tricuspiswas commonly observed to hover over and attempt to oviposit on S. invicta, we never observed any parasitization attempts on any other ant species. As predicted by laboratory tests, released populations of P. tricuspis appear to be highly host specific and pose no obvious threat to nontarget species.     

Who is the top dog in ant communities? Resources,parasitoids, and multiple competitive hierarchies

A wide variety of animal communities are organized into interspecific dominance hierarchies associated with the control and harvest of food resources. Interspecific dominance relationships are commonly found to be linear. However, dominance relations within communities can form a continuum ranging from intransitive networks to transitive, linear dominance hierarchies. How interference competition affects community structure depends on the configuration of the dominance interactions among the species. This study explores how resource size and the trait-mediated indirect effect (TMIE) specialist phorid fly parasitoids exert on interference competition, affect the transitive nature of competitive interactions in an assemblage of woodland ants. I quantify the linearity of networks of interactions associated with large and small food resources in the presence and absence of phorid parasitoids. Two distinct, significantly linear dominance hierarchies exist within the ant assemblage depending on the size of the disputed resource. However, the presence of phorid fly parasitoids eliminates the linearity of both dominance hierarchies. The hosts phorid defense behaviors reduce the competitive asymmetries between the host and its subdominant competitors, increasing the indeterminacy in the outcome of competitive interactions. Thus, both resource size variation and phorid-induced TMIEs appear to facilitate coexistence in assemblages of scavenging ants.     

Comparing longevity of Pseudacteon species of different sizes: effect of sugar feeding

Sugar feeding is known to enhance lifespan in many parasitoid species. Several species of phorid flies in the genus Pseudacteon (Diptera: Phoridae) have been introduced in the Southern U.S.A. for the biological control of imported fire ants of Solenopsis species (Hymenoptera: Formicidae). Pseudacteon species are short‐lived flies and little is known about their nutritional ecology. Results from previous studies in our laboratory show that one of the introduced species, Pseudacteon tricuspis Borgmeier, is capable of feeding on sugar sources with a significant increase in lifespan. However, in Pseudacteon phorid flies, the degree to which sugar feeding can enhance the lifespan of other species is not clear, nor is it known whether there is a relationship between body size and longevity. In the present study, the effect of sugar feeding on the survival of three phorid fly species of different body sizes, Pseudacteon cultellatus Borgmeier, Pseudacteon curvatus Borgmeier and Pseudacteon obtusus Borgmeier, is investigated. For all three species, flies continuously provided with 40% or 20% sucrose solution live longer than individuals provided water only or 40% sucrose solution only on the first day of adult life. However, the degree to which sugar feeding enhanced longevity varies by species and is highest for P. obtusus. The data also indicate that longevity in Pseudacteon phorid flies is related to body size: the largest species (P. obtusus) lives significantly longer than the smaller species (P. cultellatus and P. curvatus). These results suggest that adults of all three Pseudacteon species are capable of feeding on sucrose solution and that sugar feeding can enhance their longevity. The significance of these results is discussed in relation to the field performance of Pseudacteon phorid flies as biological control agents of imported fire ants.     

Edge‐mediated reduction of phorid parasitism on leaf‐cutting ants in a Brazilian Atlantic forest

Previous studies have shown that leaf‐cutting ant populations benefit greatly from living in or near the edges of the Brazilian Atlantic forest. One of the mechanisms responsible for this rise in population density is an edge‐mediated increase of pioneer plants, resulting in increased food availability for the ants (i.e., less bottom‐up control). Here, we hypothesized that the release from natural enemies (i.e., less top‐down control) may also contribute to the phenomenon. We investigated whether parasitism of phorid flies on leaf‐cutting ants decreases in colonies located along the forest edge vs. the interior of a large tract of Atlantic forest in northeastern Brazil. For this, we assessed abundance and rates of oviposition attack by phorids in bimonthly intervals over a period of 1 year in 10 adult colonies of Atta cephalotes (L.) (Hymenoptera: Formicidae: Myrmicinae), five at the forest edge and five in the forest interior. The number of phorids attracted by ants at edge colonies was 40% lower than that at interior colonies. The temporal variation in phorid attraction was also significant, with approximately 35% fewer flies in the dry months as compared to the rainy months. As a result of lower phorid abundance, ant workers of edge colonies suffered three times fewer oviposition attacks than those of interior colonies. There was a tendency for fewer attacks during dry months, but the difference in the temporal variation was not significant. Our findings suggest that edge creation contributes to increased leaf‐cutting ant abundance, not only via the attenuation of bottom‐up forces, but also through an environmentally triggered depression of parasitoid abundance/efficiency, possibly because of adverse environmental conditions in edge habitats.     

Dominance in a ground‐dwelling ant community of banana agroecosystem

In tropical ecosystems, ants represent a substantial portion of the animal biomass and contribute to various ecosystem services, including pest regulation and pollination. Dominant ant species are known to determine the structure of ant communities by interfering in the foraging of other ant species. Using bait and pitfall trapping experiments, we performed a pattern analysis at a fine spatial scale of an ant community in a very simplified and homogeneous agroecosystem, that is, a single‐crop banana field in Martinique (French West Indies). We found that the community structure was driven by three dominant species (Solenopsis geminata, Nylanderia guatemalensis, and Monomorium ebeninum) and two subdominant species (Pheidole fallax and Brachymyrmex patagonicus). Our results showed that dominant and subdominant species generally maintained numerical dominance at baits across time, although S. geminata, M. ebeninum, and B. patagonicus displayed better abilities to maintain dominance than P. fallax and N. guatemalensis. Almost all interspecific correlations between species abundances, except those between B. patagonicus and N. guatemalensis, were symmetrically negative, suggesting that interference competition prevails in this ground‐dwelling ant community. However, we observed variations in the diurnal and nocturnal foraging activity and in the daily occurrence at baits, which may mitigate the effect of interference competition through the induction of spatial and temporal niche partitioning. This may explain the coexistence of dominant, subdominant, and subordinate species in this very simplified agroecosystem, limited in habitat structure and diversity.     

Environmental filtering of foraging strategies mediates patterns of coexistence in the fire ants Solenopsis geminata and Solenopsis xyloni,and their interspecific hybrids

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Habitat association and coexistence of endemic and introduced ant species in the Galápagos Islands

1. We investigated ant communities in all main vegetation zones of the model island of Santa Cruz in the Galápagos archipelago (155 collection points, spread over 21 sites; 28 ant species collected), and evaluated the distribution, coexistence, and effect of environmental factors in a community composed of endemic, probably endemic, and introduced ants of the New World and exotic origin. 2. Introduced species were the most frequent, occurring in 98% of the samples, yet endemic and probably endemic species still occurred in 54% of the samples, and constituted one of three most common species. The present study revealed that the habitat type along with altitude and the tree cover are the primary factors shaping ant community composition. Little evidence was found for a competitively structured assemblage of ant species. 3. The present study confirmed the predominance of two dominant invasive species, Solenopsis geminata Fabricius and Wasmannia auropunctata Roger, whose abundances are negatively correlated. The abundance of S. geminata is positively correlated with the overall species richness, and with the proportion of other introduced species. The presence of both invasive ants is associated with a low evenness of ant communities. 4. The present study (i) stresses the dominance of introduced species and the relative resistance of endemic species, (ii) highlights the on‐going processes of species introductions and (iii) points out the need for adequate monitoring and conservation of the pristine and threatened environments that constitute the Galápagos Islands.     

Interactions between ant species increase arrival rates of an ant parasitoid

Behaviours or traits associated with aggression and communication may increase an animal's conspicuousness to predators or parasitoids. Most examples of this come from instances of aggression or communication within a species. We tested whether interspecific encounters between ants enhance the host location success of a parasitoid (Diptera: Phoridae) that attacks ants in the genus Linepithema. At food resources recruited to by Linepithema, parasitoid discovery rates were lower when Linepithema was alone than when other ant species were present. In experimentally controlled encounters, parasitoid discovery rates were elevated when Linepithema confronted an ant species that elicited use of chemicals, but not when it confronted an ant species that primarily elicited physical aggression. These results indicate that phorid parasitoids of Linepithema use the ant's chemicals as host location cues. Because Linepithema is known to abandon food resources in the presence of its phorid parasitoids, its use of chemicals during interspecific encounters may diminish its competitive success when phorids are nearby. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Cascading trait‐mediated interactions induced by ant pheromones

Trait‐mediated indirect interactions (TMII) can be as important as density‐mediated indirect interactions. Here, we provide evidence for a novel trait‐mediated cascade (where one TMII affects another TMII) and demonstrate that the mechanism consists of a predator eavesdropping on chemical signaling. Ants protect scale insects from predation by adult coccinellid beetles – the first TMII. However, parasitic phorid flies reduce ant foraging activity by 50% – the second TMII, providing a window of opportunity for female beetles to oviposit in high‐quality microsites. Beetle larvae are protected from ant predation and benefit from living in patches with high scale densities. We demonstrate that female beetles can detect pheromones released by the ant when attacked by phorids, and that only females, and especially gravid females, are attracted to the ant pheromone. As ants reduce their movement when under attack by phorids, we conclude that phorids facilitate beetle oviposition, thus producing the TMII cascade.     

Resource distribution and soil moisture content can regulate bait control in an ant assemblage in Central Amazonian forest

Resources influence population growth, interspecific interactions, territoriality and, in combination with moisture content, affect terrestrial arthropod distribution and abundance. Ants are usually described as interactive and compete in transitive hierarchies, where the dominants behaviourally exclude subordinate species from food resources. In this study, we evaluated the effects of (i) dominant ants, soil moisture and an artificial resource gradient on the number of ant species attracted to baits; and (ii) how soil moisture and an artificial resource gradient change the number of controlled baits in a Central Amazonian rain forest. We sampled 30 100‐m‐long transects, located at least 200 m apart. The transects were established with six different bait densities varying between six and 41 baits and the soil moisture content was measured at 10 points for each transect. Six ant species were considered dominant, and had negative correlations with the number of species at baits (r² = 0.186; F_1,28 = 6.419; P = 0.017). However, almost half of the transects showed low abundance of dominant species (<30%), and relatively high number of species (mean of 20.1 ± 8.75). Resource availability and soil moisture had negative and positive correlations, with number of controlled baits. These results suggest that, even though the dominance is relatively poorly developed on the floor of this tropical forest, both resource availability and soil moisture affect resource control, and thus, the number of species that use baits.     

Foraging behavior as a determinant of asymmetric competitive interaction between two ant species in a tropical agroecosystem

This work is concerned with elucidating competitive interactions between two neotropical ants, Solenopsis geminata and Pheidole radoszkowskii, focusing on their foraging behavior. When released from competition from P. radoszkowskii, S. geminata increased its foraging activity. On the other hand, when released from competition from S. geminata, P. radoszkowskii did not respond, demonstrating asymmetric competition between the two species. Foraging experiments showed that P. radoszkowskii is more efficient at finding food resources, whereas S. geminata is better at defending the resources once they are encountered. These differences in foraging behavior appear to permit the coexistence of these two species. The practical implications of the results for the management of ant communities in tropical agroecosystems is discussed with respect to the potential use of ants as natural enemies.     

Phoretic relationship between the myceliophagous mite Microdispus lambi (Acari: Microdispidae) and mushroom flies in Spanish crops

We studied the role played by the phorid Megaselia halterata (Wood) and the sciarid Lycoriella auripila (Winnertz) in the phoretic dispersion of the myceliophagous mite Microdispus lambi (Acari: Pygmephoridae). Twenty‐four crops were monitored during 18 months in commercial mushroom farms in Castilla‐La Mancha (Spain). Adults of both species were collected weekly and the mites they carried were counted and identified. Both phorids (19.6%) and sciarids (4.4%) carried the mite M. lambi. The calculated load of each was 3.4 M. lambi mites per phorid and 1.9 per sciarid. The same percentage of male and female phorid was used as vector, but the load was slightly higher for females (1.86 mites per female compared with1.48 mites per male). A mean of 7.2% of the phorids examined in winter were vectors of M. lambi, while in spring and autumn of the first year the average was more than 22%. The mean load did not vary significantly between seasons. Inside the mushroom farms, less than 10% of a small initial population of phorids carried mites (less than two mites per phorid). As the cycle progressed, more than 35% of a larger population of emerging flies did so (average 3.5 mites per phorid vector). At the end of the growth cycle, the flies may fly off to colonise nearby farms, favouring the propagation of M. lambi from infested to uninfested crops. Megaselia halterata is the principal vector of M. lambi in the mushroom farms of Castilla‐La Mancha due to their high numbers, the high percentage carrying mites and the number of M. lambi they carry.