Functional response and mutual interference in the parasitoid Coptera haywardi (Oglobin) (Hymenoptera: Diapriidae) attacking Anastrepha ludens (Loew) (Diptera: Tephritidae) pupae.

Functional response and mutual interference are important attributes of natural enemies that should be analysed in species with the potential to be used as biological control agents in order to increase the predictive power of the possible benefits and/or consequences of their release in the field. Our main objective was to determine the functional response and mutual interference of Coptera haywardi (Oglobin), a pupal parasitoid of economically important fruit flies (Diptera: Tephritidae). The functional response of C. haywardi on A. ludens pupae corresponded to a type II model, with an attack rate of 0.0134 host pupa/h and a handling time of 1.843 h, which reveals a meticulous selection process of pupal hosts. The effect of mutual interference among foraging females was negatively correlated with increased parasitoid density in the experimental arena, showing a gradual decline in attack rate per individual female. The increase in the number of foraging females also had an impact on the number of oviposition scars per pupa and the number of immature parasitoids per dissected pupa, but not on the percentage of adult emergence or the sex ratio. Our results suggest that C. haywardi could act as a complementary parasitoid in the control of fruit fly pupae, since the random distribution of these pupae in the soil would decrease the possibility of aggregation and mutual interference between foraging females.

Superparasitism and fitness parameters in three native wasp parasitoids (Braconidae: Opiinae) of the Mexican fruit fly, Anastrepha ludens (Diptera: Tephritidae).

Knowledge on reproductive strategies and host use in insect parasitoids is fundamental for biological control purposes. Superparasitism and multiparasitism, oviposition in a previously parasitized host by a female of the same or different species, respectively, may impact pest management decisions. Our objective was to determine the occurrence of superparasitism and multiparasitism in three species of native larval-pupal solitary endoparasitoids that attack Anastrepha Shiner species (Diptera: Tephritidae) in the Neotropical region, and the possible effect on offspring fitness parameters. Doryctobracon crawfordi (Viereck), Utetes anastrephae (Viereck), and Opius hirtus (Fischer) occur in sympatry in Mexico, and are currently under consideration for use as biocontrol agents. Experiments were conducted under laboratory conditions with females acting alone (self-superparasitism), females in groups of the same species (conspecific superparasitism), and females in mixed groups (multiparasitism). Our results showed that self-superparasitism is an uncommon strategy in the three native species and is rare under conditions of intraspecific competition. In the case of multiparasitism, a higher number of immature stages of U. anastrephae was observed, compared to those of D. crawfordi and O. hirtus. However, it is not clear yet if this was due to some adult female trait or to the competitive ability of the larvae. We conclude that most females of the native species studied appeared to avoid superparasitism, specifically when acting alone, suggesting a high discrimination ability, which is probably a result of a close relationship and evolutionary history with Anastrepha hosts.

A Neoponera villosa (Fabricius, 1804) Worker Pupa with too Many Legs (Hymenoptera: Formicidae).

Several somatic and sexual anomalies such as gynandromorphism and other morphological malformations have been described in ants, but cases of supernumerary legs in natural populations are rare, with only six cases reported to date. Moreover, few cases of abnormalities have been reported for the Ponerinae subfamily, with only three cases of genetic chimeras and five teratological cases worldwide. We report here a new case of teratology in this subfamily, with a seven-legged worker pupa found in a Neoponera villosa colony nesting in an epiphytic tank bromeliad. This is the second case for Mexico of a morphological anomaly in a ponerine ant, but the first report of a teratological case in the subfamily for this country.

Myrmozercon mites are highly host specific: two new species of Myrmozercon Berlese associated with sympatric Camponotus ants in southern Quintana Roo, Mexico.

Two new species of Myrmozercon, M. serratus sp. nov. and M. spatulatus sp. nov., are described based on adults and deutonymphs collected in association with ants in Mexico. They represent the first records of this genus from the Neotropic s.s. faunal region. Both new species are associated with hosts in the genus Camponotus from the same small area of a coastal lagoon, which share the same nesting habit preferences, providing strong evidence for host specificity. All but one colony of C. atriceps hosted mites, whereas they occurred in only half of the colonies of C. rectangularis. There was a significant positive correlation between the abundance of C. atriceps sexual ants and the abundance of mites. We summarize the known host associations for the genus Myrmozercon and discuss host specificity. Larvae of both mite species were collected on the wings of males and gynes suggesting that egg laying occurs on the hosts reproductive caste. Two hypotheses explaining this observation are discussed, larvae may be phoretic on winged sexuals, favoring mite co-dispersal with hosts, or larvae reside on the alates as a refuge from predation.

First Record of the Elusive Ant Parasitoid Horismenus floridensis (Hymenoptera: Eulophidae) in Mexico and New Association with an Ant Host.

Most eulophid wasps are primary parasitoids, mainly of endophytic insect larvae (Diptera, Coleoptera, Lepidoptera, Hymenoptera, Hemiptera, and Thysanoptera), but can also attack spider eggs, gall-forming mites, or nematodes. A few species are known to parasitize ants. Here we report on the occurrence of Horismenus floridensis (Schauff and Boucek) attacking Camponotus atriceps (Smith) in southern Mexico (Campeche), expanding the distribution for this eulophid species and the range of its potential hosts. We also provide an updated list of the Horismenus Walker species found in Mexico, which currently includes 21 identified species. This is the second host ant ever recorded for H. floridensis and the first reliable record of C. atriceps as a host for this eulophid wasp. The first host ant reported from Florida 35 years ago was the closely related valid species, C. floridanus (Buckley), erroneously synonymized at that time with C. atriceps (formerly, C. abdominalis (Fabricius)).

Cohabitation with aggressive hosts: description of a new microhisterid species in nests of a ponerine ant with ecological notes.

A new clown beetle species, Bacanius neoponerae, is described from Mexican nests of the arboreal ponerine ant Neoponera villosa found in the tank bromeliad Aechmea bracteata. Adult beetles were found in brood chambers or inner refuse piles, but also outside the ant nests, in decaying organic matter between the bromeliad leaves. No direct interactions between ants and microhisterid beetles could be observed. Several lines of evidence suggest a close relationship either with the ants, specific microhabitats within the ant nests or the bromeliads. Sample site elevation, colony size, monthly rainfall and collecting site were the main variables predicting the association. Almost half of the N. villosa colonies were associated with the microhisterids, and larger colonies favored their presence, especially during the driest months of the year. Two specimens were found in a nest of another ant species, Camponotus atriceps, also inhabiting A. bracteata. The new species is the seventh of the genus Bacanius reported from Mexico. This is the second time a species of this genus is associated with ants, and the fourth record of a histerid beetle cohabiting with ponerine ants. The small size of these beetles and their very protective body structure may facilitate their cohabitation with such aggressive hosts.

Vocalizations of wild West Indian manatee vary across subspecies and geographic location.

Geographic variation in the vocal behavior of manatees has been reported but is largely unexplored. Vocalizations of wild West Indian manatees (Trichechus manatus) were recorded with hydrophones in Florida from Florida manatees (Trichechus manatus latirostris), and in Belize and Panama from Antillean manatees (Trichechus manatus manatus) to determine if calls varied between subspecies and geographic regions. Calls were visually classified into five categories: squeaks, high squeaks, squeals, squeak-squeals, and chirps. From these five categories, only three call types (squeaks, high squeaks and squeals) were observed in all three populations. Six parameters from the temporal and frequency domains were measured from the fundamental frequency of 2878 manatee vocalizations. A repeated measures PERMANOVA found significant differences for squeaks and high squeaks between each geographic location and for squeals between Belize and Florida. Almost all measured frequency and temporal parameters of manatee vocalizations differed between and within subspecies. Variables that may have influenced the variation observed may be related to sex, body size, habitat and/or other factors. Our findings provide critical information of manatee calls for wildlife monitoring and highlight the need for further study of the vocal behavior of manatees throughout their range.

Co-occurrence in ant primary parasitoids: a Camponotus rectangularis colony as host of two eucharitid wasp genera.

Different assemblages of parasitoids may attack a given host species and non-random distribution patterns in parasitoid species assemblages have been reported on various occasions, resulting in co-occurrence at the population, colony, or even individual host levels. This is the case for different closely related species of eucharitid wasps (a family of specialized ant parasitoids) sharing similar niches and co-occurring on the same host at different levels. Here we reviewed all known associations between eucharitid wasps and the ant host genus Camponotus Mayr, 1861 and reported new ant-parasitoid associations. In addition, we report a new case of co-occurrence in eucharitid wasps, at the host colony level, involving a new undescribed species of Pseudochalcura Ashmead, 1904 and an unidentified species of Obeza Heraty, 1985, which attack the common but very poorly known neotropical arboreal ant Camponotus rectangularis Emery, 1890. Most attacks were solitary, but various cocoons were parasitized by two (16%) or three (8%) parasitoids. Globally, parasitism prevalence was very low (3.7%) but showed an important variability among samples. Low parasitism prevalence along with host exposure to parasitoid attack on host plants and overlapping reproductive periods of both parasitoid species may have allowed the evolution of co-occurrence. We also provided some additional data regarding the host ant nesting habits, the colony composition and new symbiotic associations with membracids and pseudococcids. The seemingly polydomous nesting habits of C. rectangularis could play a part in the reduction of parasitism pressure at the population level and, combined with occasionally important local parasitism rates, could also contribute to some parts of the colonies escaping from parasites, polydomy possibly representing an effective parasitism avoidance trait.

A new ant-butterfly symbiosis in the forest canopy fills an evolutionary gap.

Myrmecophilous butterflies can establish complex symbiotic relationships with ants. A caterpillar wandering among the brood of the aggressive ponerine ant Neoponera villosa was found inside the core of a nest built in the myrmecophytic bromeliad Aechmea bracteata. This is the first caterpillar found living inside a ponerine ant nest. Its DNA barcode was sequenced, and an integrative approach was used to identify it as Pseudonymphidia agave, a poorly known member of the subtribe Pachythonina in the riodinid tribe Nymphidiini. The cuticle of the tank-like caterpillar lacks projections or tubercles and is covered dorsally by specialized flat setae that form an armor of small plates. Ant-organs potentially related to caterpillar-ant signaling, such as perforated cupola organs and tentacle nectary organs, are present. These morphological traits, together with evidence of social integration (direct contact with host brood, protective morphology, slow movement, no host aggressiveness), suggest that P. agave is a symbiotic, social parasite of N. villosa, preying on its host brood. However, several knowledge gaps remain, including oviposition site, dependence on bromeliad association, steps to colony integration, and larval diet through development. Carnivory has been reported in all known members of the subtribe Pachythonina (caterpillars prey on honeydew-producing hemipterans) suggesting a shift to myrmecophagy inside the ant nests as a possible evolutionary transition.

Nest Site Selection during Colony Relocation in Yucatan Peninsula Populations of the Ponerine Ants Neoponera villosa (Hymenoptera: Formicidae).

In the Yucatan Peninsula, the ponerine ant Neoponera villosa nests almost exclusively in tank bromeliads, Aechmea bracteata. In this study, we aimed to determine the factors influencing nest site selection during nest relocation which is regularly promoted by hurricanes in this area. Using ants with and without previous experience of Ae. bracteata, we tested their preference for refuges consisting of Ae. bracteata leaves over two other bromeliads, Ae. bromeliifolia and Ananas comosus. We further evaluated bromeliad-associated traits that could influence nest site selection (form and size). Workers with and without previous contact with Ae. bracteata significantly preferred this species over others, suggesting the existence of an innate attraction to this bromeliad. However, preference was not influenced by previous contact with Ae. bracteata. Workers easily discriminated between shelters of Ae. bracteata and A. comosus, but not those of the closely related Ae. bromeliifolia. In marked contrast, ants discriminated between similar sized Ae. bracteata and Ae. bromeliifolia plants, suggesting that chemical cues and plant structure play an important role. Size was also significant as they selected the largest plant when provided two dissimilar Ae. bracteata plants. Nest site selection by N. villosa workers seems to depend on innate preferences but familiarization with plant stimuli is not excluded.

Mexico ants: incidence and abundance along the Nearctic-Neotropical interface.

Mexico is one of the most biodiverse countries in the world, with an important proportion of endemism mainly because of the convergence of the Nearctic and Neotropical biogeographic regions, which generate great diversity and species turnover at different spatial scales. However, most of our knowledge of the Mexican ant biota is limited to a few well-studied taxa, and we lack a comprehensive synthesis of ant biodiversity information. For instance, most of the knowledge available in the literature on Mexican ant fauna refers only to species lists by states, or is focused on only a few regions of the country, which prevents the study of several basic and applied aspects of ants, from diversity and distribution to conservation. Our aims in this data paper are therefore (1) to compile all the information available regarding ants across the Mexican territory, and (2) to identify major patterns in the gathered data set and geographic gaps in order to direct future sampling efforts. All records were obtained from raw data, including both unpublished and published information. After exhaustive filtering and updating information and synonyms, we compiled a total of 21,731 records for 887 ant species distributed throughout Mexico from 1894 to 2018. These records were concentrated mainly in the states of Chiapas (n = 6,902, 32.76%) and Veracruz de Ignacio de la Llave (n = 4,329, 19.92%), which together comprise half the records. The subfamily with the highest number of records was Myrmicinae (n = 10,458 records, 48.12%), followed by Formicinae (n = 3,284, 15.11%) and Ponerinae (n = 1,914, 8.8%). Most ant records were collected in the Neotropical region of the country (n = 12,646, 58.19%), followed by the Mexican transition zone (n = 5,237, 24.09%) and the Nearctic region (n = 3,848, 17.72%). Native species comprised 95.46% of the records (n = 20,745). To the best of our knowledge, this is the most complete data set available to date in the literature for the country. We hope that this compilation will encourage researchers to explore different aspects of the population and community research of ants at different spatial scales, and to aid in the establishment of conservation policies and actions. There are no copyright restrictions. Please cite this data paper when using its data for publications or teaching events.

Fine-tuned intruder discrimination favors ant parasitoidism.

A diversity of arthropods (myrmecophiles) thrives within ant nests, many of them unmolested though some, such as the specialized Eucharitidae parasitoids, may cause direct damage to their hosts. Ants are known to discriminate between nestmates and non-nestmates, but whether they recognize the strength of a threat and their capacity to adjust their behavior accordingly have not been fully explored. We aimed to determine whether Ectatomma tuberculatum ants exhibited specific behavioral responses to potential or actual intruders posing different threats to the host colony and to contribute to an understanding of complex ant-eucharitid interactions. Behavioral responses differed significantly according to intruder type. Ants evicted intruders that represented a threat to the colony's health (dead ants) or were not suitable as prey items (filter paper, eucharitid parasitoid wasps, non myrmecophilous adult weevils), but killed potential prey (weevil larvae, termites). The timing of detection was in accordance with the nature and size of the intruder: corpses (a potential source of contamination) were detected faster than any other intruder and transported to the refuse piles within 15 min. The structure and complexity of behavioral sequences differed among those intruders that were discarded. Workers not only recognized and discriminated between several distinct intruders but also adjusted their behavior to the type of intruder encountered. Our results confirm the previously documented recognition capabilities of E. tuberculatum workers and reveal a very fine-tuned intruder discrimination response. Colony-level prophylactic and hygienic behavioral responses through effective removal of inedible intruders appears to be the most general and flexible form of defense in ants against a diverse array of intruders. However, this generalized response to both potentially lethal and harmless intruders might have driven the evolution of ant-eucharitid interactions, opening a window for parasitoid attack and allowing adult parasitoid wasps to quickly leave the natal nest unharmed.

Context dependent life-history shift in Macrodinychus sellnicki mites attacking a native ant host in Colombia.

Ant parasitoidism has been reported in seven of the 26 recognized species of the mite genus Macrodinychus (Machrodynichidae). Macrodynichus sellnicki, previously reported as a parasitoid of the invasive ant Nylanderia fulva in Colombia, is now reported, in the same region, as attacking a native host, Ectatomma sp. 2 (E. ruidum complex). The mite develops within the protective silk cocoon of an Ectatomma pupa and waits for the emergence of the young ant before leaving the cocoon, unmolested. Overall nest prevalence was relatively high (34.6% of the 52 nests containing cocoons) but pupae prevalence was low (4.0%, n = 1401 cocoons). Mite life-history (parasite or parasitoid) was context dependent, shifting according to the intensity of the attack on a same host. Contrary to the strictly parasitoidic association of M. sellnicki with N. fulva, single mite attacks against E. ruidum did not result in host killing and solitary M. sellnicki (78.6% of the cases) behaved as parasites. However, in 21.4% of the attacks (0.9% of all available host pupae) more than one mite was involved and behaved as parasitoids, draining the host of its internal fluids and killing it. This is the first association of a macrodinychid mite with a species of the subfamily Ectatomminae, and the first ant associated mite for which such a context dependent life-style shift is described.

Mandible strike: the lethal weapon of Odontomachus opaciventris against small prey.

In order to study both the hunting efficiency and the flexibility of their predatory behavior, solitary hunters of the trap-jaw ant Odontomachus opaciventris were offered small prey (termites, fruit flies and tenebrionid larvae), presenting different morphological or defensive characteristics. The monomorphic hunters showed a moderately flexible predatory behavior characterized by short capture sequences and a noteworthy efficiency of their mandible strike (76.7-100% of prey retrievals), even when presented with Nasutitermes soldiers. Contrary to most poneromorph ants, antennal palpation of the prey before the attack was always missing, no particular targeted region of the prey's body was preferred, and no 'prudent' posture was ever exhibited. Moreover, stinging was regularly performed on bulky, fast moving fruit flies, very scarcely with sclerotized tenebrionid larvae, but never occurred with Nasutitermes workers or soldiers despite their noxious chemical defense. These results suggest that, whatever the risk linked to potentially dangerous prey, O. opaciventris predatory strategy optimizes venom use giving top priority to the swiftness and strength of the lethal trap-jaw system used by hunters as first strike weapon to subdue rapidly a variety of small prey, ranging from 0.3 to 2 times their own body size and from 0.1 to 2 times their weight. Such risk-prone predatory behavior is likely to be related to the large size of O. opaciventris colonies where the death of a forager might be of lesser vital outcome than in small colony-size species.

Hidden biodiversity in entomological collections: The overlooked co-occurrence of dipteran and hymenopteran ant parasitoids in stored biological material.

Biological collections around the world are the repository of biodiversity on Earth; they also hold a large quantity of unsorted, unidentified, or misidentified material and can house behavioral information on species that are difficult to access or no longer available to science. Among the unsorted, alcohol-preserved material stored in the Formicidae Collection of the 'El Colegio de la Frontera Sur' Research Center (Chetumal, Mexico), we found nine colonies of the ponerine ant Neoponera villosa, that had been collected in bromeliads at Calakmul (Campeche, Mexico) in 1999. Ants and their brood were revised for the presence of any sign of parasitism. Cocoons were dissected and their content examined under a stereomicroscope. Six N. villosa prepupae had been attacked by the ectoparasitoid syrphid fly Hypselosyrphus trigonus Hull (Syrphidae: Microdontinae), to date the only known dipteran species of the Microdontinae with a parasitoid lifestyle. In addition, six male pupae from three colonies contained gregarious endoparasitoid wasps. These were specialized in parasitizing this specific host caste as no gyne or worker pupae displayed signs of having been attacked. Only immature stages (larvae and pupae) of the wasp could be obtained. Due to the long storage period, DNA amplification failed; however, based on biological and morphological data, pupae were placed in the Encyrtidae family. This is the first record of an encyrtid wasp parasitizing N. villosa, and the second example of an encyrtid as a primary parasitoid of ants. Furthermore, it is also the first record of co-occurrence of a dipteran ectoparasitoid and a hymenopteran endoparasitoid living in sympatry within the same population of host ants. Our findings highlight the importance of biological collections as reservoirs of hidden biodiversity, not only at the taxonomic level, but also at the behavioral level, revealing complex living networks. They also highlight the need for funding in order to carry out biodiversity inventories and manage existing collections.

Rediscovery and reclassification of the dipteran taxon Nothomicrodon Wheeler, an exclusive endoparasitoid of gyne ant larvae.

The myrmecophile larva of the dipteran taxon Nothomicrodon Wheeler is rediscovered, almost a century after its original description and unique report. The systematic position of this dipteran has remained enigmatic due to the absence of reared imagos to confirm indentity. We also failed to rear imagos, but we scrutinized entire nests of the Brazilian arboreal dolichoderine ant Azteca chartifex which, combined with morphological and molecular studies, enabled us to establish beyond doubt that Nothomicrodon belongs to the Phoridae (Insecta: Diptera), not the Syrphidae where it was first placed, and that the species we studied is an endoparasitoid of the larvae of A. chartifex, exclusively attacking sexual female (gyne) larvae. Northomicrodon parasitism can exert high fitness costs to a host colony. Our discovery adds one more case to the growing number of phorid taxa known to parasitize ant larvae and suggests that many others remain to be discovered. Our findings and literature review confirm that the Phoridae is the only taxon known that parasitizes both adults and the immature stages of different castes of ants, thus threatening ants on all fronts.

Macrodinychus mites as parasitoids of invasive ants: an overlooked parasitic association.

Mites are frequent ant symbionts, yet the exact nature of their interactions with their hosts is poorly known. Generally, myrmecophilous mites show adaptations for dispersal through phoresis, but species that lack such an adaptation may have evolved unusual specialized relationships with their hosts. The immature stages of Macrodinychus multispinosus develop as ectoparasitoids of pupae of the invasive ant Paratrechina longicornis. Feeding stages show regressed locomotor appendages. These mites complete their development on a single host, sucking all of its body content and therefore killing it. Locally high proportions of parasitized host pupae suggest that M. multispinosus could serve as a biological control agent. This is the ninth species of Macrodinychus reported as ant parasite, and the third known as parasitoid of invasive ants, confirming a unique habit in the evolution of mite feeding strategies and suggesting that the entire genus might be parasitic on ants. Several mites' characteristics, such as their protective morphology, possible viviparity, lack of a specialized stage for phoretic dispersal, and low host specificity, combined with both the general low aggressiveness of invasive P. longicornis towards other ants and its possible susceptibility to generalist ectoparasites would account for the host shift in native macrodinychid mites.

Local and Landscape Drivers of Ant Parasitism in a Coffee Landscape.

Parasitism of ants that nest in rotting wood by eucharitid wasps was studied in order to examine whether habitat and season influence ant parasitism, vegetation complexity and agrochemical use correlate with ant parasitism, and whether specific local and landscape features of agricultural landscapes correlate with changes in ant parasitism. In a coffee landscape, 30 coffee and 10 forest sites were selected in which local management (e.g., vegetation, agrochemical use) and landscape features (e.g., distance to forest, percent of rustic coffee nearby) were characterized. Rotten logs were sampled and ant cocoons were collected from logs and cocoons were monitored for parasitoid emergence. Sixteen ant morphospecies in three ant subfamilies (Ectatomminae, Ponerinae, and Formicinae) were found. Seven ant species parasitized by two genera of Eucharitidae parasitoids (Kapala and Obeza) were reported and some ant-eucharitid associations were new. According to evaluated metrics, parasitism did not differ with habitat (forest, high-shade coffee, low-shade coffee), but did increase in the dry season for Gnamptogenys ants. Parasitism increased with vegetation complexity for Gnamptogenys and Pachycondyla and was high in sites with both high and low agrochemical use. Two landscape variables and two local factors positively correlated with parasitism for some ant genera and species. Thus, differences in vegetation complexity at the local and landscape scale and agrochemical use in coffee landscapes alter ecological interactions between parasitoids and their ant hosts.

How to escape from the host nest: imperfect chemical mimicry in eucharitid parasitoids and exploitation of the ants' hygienic behavior.

Communication in ants is based to a great extent on chemical compounds. Recognition of intruders is primarily based on cuticular hydrocarbon (CHC) profile matching but is prone to being cheated. Eucharitid wasps are specific parasitoids of the brood of ants; the immature stages are either well integrated within the colony or are protected within the host cocoons, whereas adult wasps at emergence must leave their host nest to reproduce and need to circumvent the ant recognition system to escape unscathed. The behavioral interactions between eucharitid wasps and workers of their host, the Neotropical ant Ectatomma tuberculatum, are characterized. In experimental bioassays, newly emerged parasitoids were not violently aggressed. They remained still and were grabbed by ants upon contact and transported outside the nest; host workers were even observed struggling to reject them. Parasitoids were removed from the nest within five minutes, and most were unharmed, although two wasps (out of 30) were killed during the interaction with the ants. We analyzed the CHCs of the ant and its two parasitoids, Dilocantha lachaudii and Isomerala coronata, and found that although wasps shared all of their compounds with the ants, each wasp species had typical blends and hydrocarbon abundance was also species specific. Furthermore, the wasps had relatively few CHCs compared to E. tuberculatum (22-44% of the host components), and these were present in low amounts. Wasps, only partially mimicking the host CHC profile, were immediately recognized as alien and actively removed from the nest by the ants. Hexane-washed wasps were also transported to the refuse piles, but only after being thoroughly inspected and after most of the workers had initially ignored them. Being recognized as intruder may be to the parasitoids' advantage, allowing them to quickly leave the natal nest, and therefore enhancing the fitness of these very short lived parasitoids. We suggest that eucharitids take advantage of the hygienic behavior of ants to quickly escape from their host nests.

Queen dimorphism and reproductive capacity in the ponerine ant, Ectatomma ruidum Roger.

We report here the first case of queen dimorphism in a ponerine ant species. A total of 550 colonies of the Neotropical ponerine ant Ectatomma ruidum, from two natural populations in southeastern Mexico, were investigated for polymorphism and reproductive function within the queen caste. The distribution of different morphological traits (head and thorax widths, and scutum and alitrunk lengths) was shown to be bimodal. Thorax width and head width were significantly isometric for both macro- and microgynes and thorax proportions did not change with the category of the queen. Microgynes appear as an isometric reduction of the normal queens. On average, macrogynes were found to be approximately 20% larger in size and twice as heavy as microgynes (and up to three times heavier in terms of dry weight). Their wing surface was 72% greater. There was no difference between the two morphs in terms of their capacity to be inseminated and all dealate females present in natural colonies could be considered as true potentially reproductive queens whatever the class to which they belonged. Ovary size, number of ovarioles per ovary, and number of mature oocytes produced were significantly higher for macrogynes than for microgynes, but small queens were able to function as active egg-layers. However, the number of egg-laying individuals was significantly lower for this form, although egg-laying activity appeared to be independent of queen weight. Despite its markedly reduced reproductive capacity compared to macrogynes, the distribution of the microgyne form of E. ruidum in both studied populations was very broad, reaching one-third of all alate and dealate females and extending to one-third of all colonies. In 72.2% of the colonies where alate and/or dealate microgynes were present, both forms co-occurred. Fat content analysis results supported the idea that semi-claustral independent colony founding, typical for this species, would essentially be achieved by macrogynes, while microgynes would be adopted by established colonies. We argue that queen dimorphism in E. ruidum could represent an attractive alternative dispersal strategy for female sexuals. The small females may constitute an important reserve of potential reproductives at a very low energetic cost to the colony, the production of a macrogyne being about 7.5 times more costly than a microgyne. The significantly greater `wing surface/body weight' ratio of microgynes also suggests their greater capacity for dispersion. The combination of such a capacity for microgynes, along with their ability to contribute in the production of both female morphs, including the reproductively efficient macrogynes, would contribute to insuring genetic reassortment at the population level and could explain, in part, the ecological success of E. ruidum in Neotropical zones.