Impact of Long-Term Floods on Spatial Dynamics of Myrmica scabrinodis, a Host Ant of a Highly Threatened Scarce Large Blue (Phengaris teleius).

Extensively used wet meadows with high species diversity are under threat in Europe by anthropogenic pressure. The increasing frequency of prolonged flooding is emerging as an additional threat to this fragile environment. In our study, we investigated how prolonged flooding affects the spatial distribution and temporal dynamics (through mortality and recolonization process) of the host ant species Myrmica scabrinodis, which is essential for the survival of the endangered Scarce large blue (Phengaris teleius). The study was conducted in the flood-prone Ljubljansko barje plain situated on the southern edge of the species' global range. Prolonged flooding in the study area, possibly affecting the past and current distribution of the host ant M. scabrinodis, was recorded in 2010, 2013, and 2017. In 2020, we set 160 ant traps to estimate the distribution of host ants in a system of meadows covering the entire gradient of flood history. Results indicate that M. scabrinodis survives the flooding for up to three days, starting to disappear if flooding persists longer. After the flooding recedes, ants gradually recolonize empty habitats from the surrounding upland refugia. Our spatial analyses predict that the average recolonization speed was about 29 m per year and that in a year, ants compensate for the mortality effects of 1.8 days of flooding by recolonization in a year. These results show that flooding should be considered as an additional (in some areas, a major) threat to the endangered P. teleius through its deleterious effects on the host ant species.

Extrafloral Nectary-Bearing Plants Recover Ant Association Benefits Faster and More Effectively after Frost-Fire Events Than Frost.

The Cerrado confronts threats such as fire and frost due to natural or human-induced factors. These disturbances trigger attribute changes that impact biodiversity. Given escalating climate extremes, understanding the effects of these phenomena on ecological relationships is crucial for biodiversity conservation. To understand how fire and frost affect interactions and influence biological communities in the Cerrado, our study aimed to comprehend the effects of these two disturbances on extrafloral nectar (EFN)-bearing plants (Ouratea spectabilis, Ochnaceae) and their interactions. Our main hypothesis was that plants affected by fire would grow again more quickly than those affected only by frost due to the better adaptation of Cerrado flora to fire. The results showed that fire accelerated the regrowth of O. spectabilis. Regrowth in plants with EFNs attracted ants that proved to be efficient in removing herbivores, significantly reducing foliar herbivory rates in this species, when compared to the species without EFNs, or when ant access was prevented through experimental manipulation. Post-disturbance ant and herbivore populations were low, with frost leading to greater reductions. Ant richness and diversity are higher where frost precedes fire, suggesting that fire restores Cerrado ecological interactions better than frost, with less impact on plants, ants, and herbivores.

The Pattern of Social Parasitism in Maculinea teleius Butterfly Is Driven by the Size and Spatial Distribution of the Host Ant Nests.

The parasitic relationship between Maculinea butterflies and Myrmica ants has been extensively studied but little information is available on the spatial occurrence of Maculinea larvae. We searched for the presence of Maculinea teleius in 211 ant nests at two sites in two crucial phases of its life cycle, i.e., in autumn, during the initial larval development, and in the following late spring, before pupation. We assessed variations in the proportion of infested nests and factors correlated with spatial distributions of parasites in Myrmica colonies. The parasitism rate in autumn was very high (∼50% of infestation rate) but decreased in the following spring. The most important factor explaining parasite occurrence in both seasons was the nest size. Further factors, such as the presence of other parasites, the Myrmica species or the site, concurred to explain the differential survival of Ma. teleius until the final development. Irrespective of the host nest distribution, the parasite distribution changed from even in autumn to clumped in late spring. Our work showed that the survival of Ma. teleius is correlated with colony features but also with the nest spatial distribution, which therefore should be taken into consideration in conservation strategies aiming at preserving these endangered species.

The Intranidal Myrmecophiles of the Maltese Islands with Notes on Messor Nests as Repositories of Biodiversity.

The intranidal myrmecophilous arthropod fauna of the Maltese Islands is reviewed. Thirty species from nine orders are found to be obligate myrmecophiles, of which four species are recorded from the Maltese archipelago for the first time: Phrurolithus sp. (Araneae: Phrurolithidae), Pogonolaelaps canestrinii (Berlese, 1904), Gymnolaelaps messor Joharchi, Halliday, Saboori & Kamali, 2011 and G. myrmecophilus (Berlese, 1892) (Mesostigmata: Laelapidae). Phrurolithus also represents the first record of the family Phrurolithidae in Malta. Notes on the biology and local distribution of each species are provided, including ant-myrmecophile associations, of which two appear to be previously unknown: the occurrence of Smynthurodes betae Westwood, 1849 (Hemiptera: Aphididae) in the nest of Plagiolepis pygmaea (Latreille, 1798) and Phrurolithus in the nest of Pheidole pallidula (Nylander, 1849). Fourteen additional species are found to be either only occasionally myrmecophilic, accidental ant-guests or potentially myrmecophilous, the latter remaining ambiguous due to a lack of knowledge of their biology. Of these, the family Caeculidae (Arachnida: Trombidiformes) represents a new record for the Maltese Islands, on the basis of Microcaeculus sp. occurring in a nest of Camponotus barbaricus Emery, 1905. Preliminary results indicate that Messor nests may be repositories of considerable myrmecophile diversity, with the most unique symbionts.

Quantifying the impacts of symbiotic interactions between two invasive species: the tawny crazy ant (Nylanderia fulva) tending the sorghum aphid (Melanaphis sorghi).

The establishment of new symbiotic interactions between introduced species may facilitate invasion success. For instance, tawny crazy ant (Nylanderia fulva Mayr) is known to be an opportunistic tender of honeydew producing insects and this ants' symbiotic interactions have exacerbated agricultural damage in some invaded regions of the world. The invasive sorghum aphid (Melanaphis sorghi Theobald) was first reported as a pest in the continental United States-in Texas and Louisiana-as recent as 2013, and tawny crazy ant (TCA) was reported in Texas in the early 2000s. Although these introductions are relatively recent, TCA workers tend sorghum aphids in field and greenhouse settings. This study quantified the tending duration of TCA workers to sorghum aphids and the impact of TCA tending on aphid biomass. For this study aphids were collected from three different host plant species (i.e., sugarcane, Johnson grass, and sorghum) and clone colonies were established. Sorghum is the main economic crop in which these aphids occur, hence we focused our study on the potential impacts of interactions on sorghum. Quantification of invasive ant-aphid interactions, on either stems or leaves of sorghum plants, were conducted in greenhouse conditions. Our results show that although these two invasive insect species do not have a long coevolutionary history, TCA developed a tending interaction with sorghum aphid, and aphids were observed excreting honeydew after being antennated by TCA workers. Interestingly, this relatively recent symbiotic interaction significantly increased overall aphid biomass for aphids that were positioned on stems and collected from Johnson grass. It is recommended to continue monitoring the interaction between TCA and sorghum aphid in field conditions due to its potential to increase aphid populations and sorghum plant damage.

Protective Benefits of Tending Ants to a Critically Endangered Butterfly.

Ants provide protection to various organisms via myrmecophilous relationships. Most notably, ants and several butterfly species are involved in mainly mutualistic interactions. Previous field studies have shown that butterfly larval survival is increased in the presence of tending ants, suggesting that ants are providing protection against insect predation or parasitism. Here, we conducted a series of timed observational trials under laboratory conditions to assess larval survival and ant protection from insect predators for a myrmecophilous lycaenid butterfly. We focused on a critically endangered butterfly, the Miami blue (Cyclargus thomasi bethunebakeri) (Comstock and Huntington) (Lepidoptera: Lycaenidae), and its most common ant associate, the Florida carpenter ant (Camponotus floridanus) (Buckley) (Hymenoptera: Formicidae), to test this assumption of ant protection. We found that ants provide significant protection to Miami blue larvae, with later instar larvae receiving a higher level of protection due to differences in tending frequencies. These results will aid in informing conservation management and future organism reintroductions for this endangered butterfly.

Temporal variation in the effect of ants on the fitness of myrmecophilic plants: seasonal effect surpasses periodic benefits.

Plants provide extrafloral nectar, which is a food resource taken by ants, especially aggressive species that may act as plant guards. To our knowledge, no study has been conducted to concurrently investigate the fluctuation of plant fitness over its whole reproductive season, recording and comparing both short periods (different samplings during the plant's reproductive season) and the season/pooled data (all fruits produced during the reproductive season). Here, by assigning plants to either ant-present or absent treatments, we investigated the influence of the protective foliage-dwelling ant, Camponotus crassus, on the flower bud and fruit production of four extrafloral nectaried plants (Ancistrotropis firmula, Bionia coriacea, Cochlospermum regium, and Peixotoa tomentosa) throughout their annual reproductive season. Periodic samples in the field revealed a large variation in plant reproduction throughout the season; the increases in buds and fruits were not constantly higher in plants with ants, and in fact, plants without ants had more reproductive structures sometimes. Nonetheless, the examination of the pooled data, i.e., cumulative number of flower buds and fruits produced during the reproductive season, revealed the plants with ants produced more flower buds and fruits (e.g., up to two-fold greater in A. firmula) compared to ant-absent treatments. Our results indicate the effects of ants on plant reproduction are not constant over time, but the net benefits to plants with ants are reflected in increased fruit production. Therefore, the investigations of the benefit of ants on plants should consider the whole plant's reproductive season rather than single samplings within plant reproduction period.

The Ecological Significance of Aphid Cornicles and Their Secretions.

Aphid cornicles are abdominal appendages that secrete an array of volatile and nonvolatile compounds with diverse ecological functions. The emission of alarm pheromones yields altruistic benefits for clone-mates in the aphid colony, which is essentially a superorganism with a collective fate. Secreted droplets also contain unsaturated triglycerides, fast-drying adhesives that can be lethal when smeared on natural enemies but more often impede their foraging efficiency. The longest cornicles have evolved in aphids that feed in exposed locations and are likely used to scent-mark colony intruders. Reduced cornicles are associated with reliance on alternative defenses, such as the secretion of protective waxes or myrmecophily. Root-feeding and gall-forming lifestyles provide protected feeding sites and are associated with an absence of cornicles. In some eusocial gall-formers, soldier morphs become repositories of cornicle secretion used to defend the gall, either as menopausal apterae that defend dispersing alatae or as sterile first instars that dispatch predators with their stylets and use cornicle secretions as a construction material for gall repair. Collectively, the evidence is consistent with an adaptive radiation of derived cornicle functions molded by the ecological lifestyle of the aphid lineage.

Mixed pollination system and floral signals of Paepalanthus (Eriocaulaceae): insects and geitonogamy ensure high reproductive success.

BACKGROUND AND AIMS: Eriocaulaceae exhibit a great variety of floral traits associated with insect (e.g. nectariferous structures) and wind pollination (unisexual flowers, exposed sexual organs and small pollen grains), as well as the 'selfing syndrome' (small flowers, short distance between stigma and anthers, and temporal overlap of male and female phases). Paepalanthus bifidus, P. subtilis and P. tortilis are related species that differ in form, size and colour of floral structures. We aimed to investigate the pollination and reproductive biology of these three species. METHODS: We analysed the floral biology, floral visitors, pollinator behaviour, and the contribution of insects, wind and spontaneous geitonogamy to fruit set. We also evaluated the floral colour and scent of the species. Colour reflectance of capitula of each species was measured and plotted in models of insect vision. Floral scent samples were extracted and the compounds were compared to vegetative scent samples. KEY RESULTS: In all species, the staminate and pistillate flowers are arranged in alternating cycles with a temporal overlap between these phases. Ants were the most frequent floral visitors and were effective pollinators in P. bifidus and P. tortilis, while flies were occasional pollinators in P. tortilis. Floral visitors were not observed in P. subtilis. In all species, fruits were produced by spontaneous geitonogamy, with no evidence of wind pollination. According to the models of insect vision, the colours of the capitula of P. bifidus and P. subtilis are the most inconspicuous for ants and flies. We found no difference between the emission of volatiles of inflorescences and vegetative structures. CONCLUSIONS: This study suggests that ant pollination might be more widespread in Eriocaulaceae than currently assumed. Furthermore, for small monocarpic plants, mixed mating strategies are most favourable, by ensuring reproduction either by outcrossing when pollinators are abundant or by spontaneous geitonogamy when pollinations are scarce/absent.

Chemical and behavioural strategies along the spectrum of host specificity in ant-associated silverfish.

BACKGROUND: Host range is a fundamental trait to understand the ecological and evolutionary dynamics of symbionts. Increasing host specificity is expected to be accompanied with specialization in different symbiont traits. We tested this specificity-specialization association in a large group of 16 ant-associated silverfish species by linking their level of host specificity to their degree of behavioural integration into the colony and to their accuracy of chemically imitating the host's recognition system, i.e. the cuticular hydrocarbon (CHC) profile. RESULTS: As expected, facultative associates and host generalists (targeting multiple unrelated ants) tend to avoid the host, whereas host-specialists (typically restricted to Messor ants) were bolder, approached the host and allowed inspection. Generalists and host specialists regularly followed a host worker, unlike the other silverfish. Host aggression was extremely high toward non-ant-associated silverfish and modest to low in ant-associated groups. Surprisingly, the degree of chemical deception was not linked to host specificity as most silverfish, including facultative ant associates, imitated the host's CHC profile. Messor specialists retained the same CHC profile as the host after moulting, in contrast to a host generalist, suggesting an active production of the cues (chemical mimicry). Host generalist and facultative associates flexibly copied the highly different CHC profiles of alternative host species, pointing at passive acquisition (chemical camouflage) of the host's odour. CONCLUSIONS: Overall, we found that behaviour that seems to facilitate the integration in the host colony was more pronounced in host specialist silverfish. Chemical deception, however, was employed by all ant-associated species, irrespective of their degree of host specificity.

Nectaries in ferns: their taxonomic distribution, structure, function, and sugar composition.

PREMISE: Extrafloral nectaries have mainly been studied in angiosperms, but have also been reported in 39 fern species. Here we provide a global review of nectaries in ferns and examined their structure, function, and nectar sugar composition in two genera. METHODS: We searched in the literature and living plant collections of botanical gardens for indications of fern nectaries, observed nectar-feeding animals, studied the morphoanatomy in the two genera Aglaomorpha and Campyloneurum, and analyzed the total sugar concentrations and ratios of 16 species. Diurnal nectar release was observed with time-lapse photography. RESULTS: We found evidence for nectaries in 101 species of ferns from 11 genera and 6 families. Most of the nectary-bearing species were tree ferns (Cyatheaceae) and epiphytic ferns of the family Polypodiaceae. Nectaries consisted of cytoplasm-rich parenchyma with large nuclei and an epidermis with or without stomata, were attached to amphiphloic vascular bundles, and released nectar on the lower leaf surface mainly on expanding leaves during the night. Sugar concentrations varied between species (3.8-15.3%) but not between genera, and were sucrose-dominant (3 spp.), sucrose-rich (7), or hexose-rich (3). In the greenhouse, introduced ants, scale insects, and snails fed on the nectar. CONCLUSIONS: The wide taxonomic distribution, variable morphology, locations, and sugar compositions point to multiple evolutionary origins of fern nectaries. Nectar release in young leaves might attract mutualistic ants to protect leaves against herbivores only during this most vulnerable developmental stage. Even ex-situ, fern nectar is a valuable food source because it attracted several opportunistic animal species.

Taxonomy of Homoeusa Kraatz, 1856 (Coleoptera, Staphylinidae) from the East Palearctic: I. Homoeusarufescens (Sharp, 1874) and a new allied species.

There is insufficient information to identify most species of the myrmecophilous rove beetle genus Homoeusa. In this paper, after examining the type material, Homoeusarufescens (Sharp, 1874) is redescribed in detail and its new allied species Homoeusaovata sp. nov. is described. We also observed the behavior of these two species in the field; the behavior was similar to that reported for H.acuminata (Märkel, 1842). A checklist of Homoeusa from the Palearctic and Nearctic is also provided.

Two new species and a new record of emGaeolaelaps/em Evans amp; Till (Acari: Laelapidae) from Altai Mountains, Russia.

This paper reports on three species of mites of the genus Gaeolaelaps from Altai Mountains, Russia. Gaeolaelaps altaiensis sp. nov., G. lenis Vatankhah Nemati were collected from nests of Lasius sp. and Formica sp. (Hymenoptera: Formicidae), respectively, and G. euparadactylifer sp. nov. from moss on the shore of the Teletskoye Lake. The new species have the unusual character state of short peritremes, ending near the middle of coxae II. In G. euparadactylifer sp. nov., the pretarsal paradactyli on legs II-IV are strongly elongated, apically pointed and with their tips projecting well beyond the claws. Both new species are described based on morphological characters of the adults. In addition, the male of Gaeolaelaps lenis is described for the first time.

Characterization of the complete mitochondrial genome of Neoasterolepisma foreli (Insecta: Zygentoma: Lepismatidae) and the phylogeny of basal Ectognatha.

The silverfish Neoasterolepisma foreli belongs to the family Lepismatidae within Zygentoma and is well known for the peculiar habit of living in strict association with ant nests (myrmecophily). In this study, we describe its mitochondrial genome, a circular molecule of 15,398 bp including the canonical 13 PCGs, 22 tRNAs, 2 rRNAs, as well as a 403 bp AT-rich region. A phylomitogenomic analysis of the new sequence, alongside basal hexapod mtDNAs, confirmed the monophyly of all orders, with some uncertainty over the position of the enigmatic Tricholepidion gertschi that would make Zygentoma paraphyletic. Neoasterolepisma foreli is recovered in a basal position within family Lepismatidae, at odd with our current understanding of the group that would, in turn, suggest a closer relationship with the genus Lepisma (Mendes, 1991).

Recognition and revision of the Phelister blairi group (Histeridae, Histerinae, Exosternini).

Forty-nine new species of Neotropical Exosternini are described in this work, representing the newly recognized Phelister blairi species group, within the large, heterogeneous taxon Phelister. Eight previously described species are also assigned to this group. Relationships within are indicated with several informal subgroups: P. blairi subgroup: (P. blairi Hinton, 1935, P. erwini sp. nov., P. fimbriatus sp. nov., P. stellans sp. nov., P. sparsus sp. nov., P. pretiosus sp. nov., P. trigonisternus Marseul, 1889, P. globosus sp. nov., P. serratus sp. nov., P. geminus sp. nov., P. parana sp. nov., P. asperatus sp. nov., P. uniformis sp. nov., P. miscellus sp. nov., P. inbio sp. nov., P. sculpturatus Schmidt, 1893, P. tunki sp. nov., P. praedatoris Reichensperger, 1939, P. ifficus sp. nov., P. genieri sp. nov., P. marginatus sp. nov., P. vazdemelloi sp. nov., P. dilatatus sp. nov., P. spectabilis sp. nov., P. pervagatus sp. nov.); P. amazoniae subgroup: (P. morbidus sp. nov., P. annulatus sp. nov., P. sphaericus sp. nov., P. geijskesi Kanaar, 1997, P. fraternus sp. nov., P. conjunctus sp. nov., P. chabooae sp. nov., P. striatinotum Wenzel & Dybas, 1941, P. notandus Schmidt, 1893, P. amazoniae (Lewis, 1898) comb. nov., P. arcuatus sp. nov.); P. gregarius subgroup: (P. gregarius sp. nov., P. praecisus sp. nov., P. rudis sp. nov., P. incongruens sp. nov., P. congruens sp. nov., P. praesignis sp. nov.); P. umens subgroup: (P. umens sp. nov., P. almeidae sp. nov., P. chicomendesi sp. nov., P. microdens sp. nov., P. matatlantica sp. nov.); P. curvipes subgroup: (P. curvipes sp. nov., P. vilavelha sp. nov.); P. rio subgroup: (P. rio sp. nov., P. semotus sp. nov., P. uncinatus sp. nov., P. inscriptus sp. nov.); incertae sedis - unplaced to subgroup: (P. incertus sp. nov., P. okeefei sp. nov., P. blairoides sp. nov., P. pirana sp. nov.). Lectotypes are designated for the following species: P. trigonisternus Marseul, P. sculpturatus Schmidt, P. praedatoris Reichensperger, P. notandus Schmidt, and Discoscelis amazoniae Lewis. Preliminary phylogenetic analyses of the broader Neotropical Exosternini do not support the monophyly of the P. blairi group, nor of all of these subgroups, but the majority do fall within one large clade (which is potentially paraphyletic with respect to some other Neotropical exosternine genera). More work on the phylogeny and taxonomy of this diverse fauna is needed.

Rhadinoscelidia lixa sp. nov. (Hymenoptera, Chrysididae, Loboscelidiinae) found on an ant nest in Thailand.

Rhadinoscelidia lixa sp. nov. is described from Thailand. It is the fifth species of the genus and second species from Thailand. A biological note on the species with its associated ants is provided.

A new species of Aricoris Westwood, 1851 (Lepidoptera: Riodinidae) from the Neotropical Cerrado savanna.

A new species of Aricoris Westwood, 1851, until now treated erroneously as Aricoris tutana (Godart, [1824]), is described and illustrated from the Neotropical Cerrado savanna. Aricoris emeryi Callaghan, Lemes Kaminski, sp. nov. can be identified by a set of characters on wings and male genitalia that differentiates it from other species belonging to the constantius group.

On systematic status of Spindasis syama Horsfield, [1829] in Taiwan and the Philippines (Lepidoptera: Lycaenidae: Aphnaeini).

Based upon both morphological and gene barcode evidence, the population of Spindasis syama in Taiwan is described as Spindasis syama lamuae, ssp. nov., and the taxon negrita Felder, 1862 is revised as a species endemic to the Philippines (stat. rev.).

Oviposition of Minstrellus grandis (Lepidoptera: Riodinidae) in a harmful ant-plant symbiosi

The oviposition behavior of the rare butterfly Minstrellus grandis (Callaghan, 1999) (Lepidoptera: Riodinidae) is recorded for the first time. Two females laid eggs on the old leaves of an unidentified Triplaris sp. (Polygonaceae), a myrmecophytic plant typically known as 'Triplaria' or 'novice' tree, inhabited by aggressive 'taxi' ants of the genus Pseudomyrmex sp. (Hymenoptera: Formicidae). These observations suggest that M. grandis caterpillars live associated with one of the most harmful types of Amazon ant-plant symbiosis. (AU)

Profound head modifications in Claviger testaceus (Pselaphinae, Staphylinidae, Coleoptera) facilitate integration into communities of ants.

Clavigeritae is a group of obligate myrmecophiles of the rove beetle subfamily Pselaphinae (Staphylinidae). Some are blind and wingless, and all are believed to depend on ant hosts through feeding by trophallaxis. Phylogenetic hypotheses suggest that their ancestors, as are most pselaphines today, were free-living predators. Morphological alterations required to transform such beetles into extreme myrmecophiles were poorly understood. By studying the cephalic morphology of Claviger testaceus, we demonstrate that profound changes in all mouthpart components took place during this process, with a highly unusual connection of the maxillae to the hypopharynx, and formation of a uniquely transformed labium with a vestigial prementum. The primary sensory function of the modified maxillary and labial palps is reduced, and the ventral mouthparts transformed into a licking/'sponging' device. Many muscles have been reduced, in relation to the coleopteran groundplan or other staphylinoids. The head capsule contains voluminous glands whose appeasement secretions are crucial for the beetle survival in ant colonies. The brain, in turn, has been shifted into the neck region. The prepharyngeal dilator is composed of an entire series of bundles. However, the pharynx does not show any peculiar adaptations to taking up liquid food. We demonstrate that far-reaching cephalic modifications characterize C. testaceus, and that the development of appeasement glands and adaptation of the mouthparts to trophallaxis determine the head architecture of this extreme myrmecophile.