Temnothorax Nylanderi Research Articles

Polydomy enhances foraging performance in ant colonies.

Collective foraging confers benefits in terms of reduced predation risk and access to social information, but it heightens local competition when resources are limited. In social insects, resource limitation has been suggested as a possible cause for the typical decrease in per capita productivity observed with increasing colony size, a phenomenon known as Michener's paradox. Polydomy (distribution of a colony's brood and workers across multiple nests) is believed to help circumvent this paradox through its positive effect on foraging efficiency, but there is still little supporting evidence for this hypothesis. Here, we show experimentally that polydomy enhances the foraging performance of food-deprived Temnothorax nylanderi ant colonies via several mechanisms. First, polydomy influences task allocation within colonies, resulting in faster retrieval of protein resources. Second, communication between sister nests reduces search times for far away resources. Third, colonies move queens, brood and workers across available nest sites in response to spatial heterogeneities in protein and carbohydrate resources. This suggests that polydomy represents a flexible mechanism for space occupancy, helping ant colonies adjust to the environment.

Gene expression patterns underlying parasite-induced alterations in host behaviour and life history.

Many parasites manipulate their hosts' phenotype. In particular, parasites with complex life cycles take control of their intermediate hosts' behaviour and life history to increase transmission to their definitive host. The proximate mechanisms underlying these parasite-induced alterations are poorly understood. The cestode Anomotaenia brevis affects the behaviour, life history and morphology of parasitized Temnothorax nylanderi ants and indirectly of their unparasitized nestmates. To gain insights on how parasites alter host phenotypes, we contrast brain gene expression patterns of T.nylanderi workers parasitized with the cestode, their unparasitized nestmates and unparasitized workers from unparasitized colonies. Over 400 differentially expressed genes between the three groups were identified, with most uniquely expressed genes detected in parasitized workers. Among these are genes that can be linked to the increased lifespan of parasitized workers. Furthermore, many muscle (functionality) genes are downregulated in these workers, potentially causing the observed muscular deformations and their inactive behaviour. Alterations in lifespan and activity could be adaptive for the parasite by increasing the likelihood that infected workers residing in acorns are eaten by their definitive host, a woodpecker. Our transcriptome analysis reveals numerous gene expression changes in parasitized workers and their uninfected nestmates and indicates possible routes of parasite manipulation. Although causality still needs to be established, parasite-induced alterations in lifespan and host behaviour appear to be partly explained by morphological muscle atrophy instead of central nervous system interference, which is often the core of behavioural regulation. Results of this study will shed light upon the molecular basis of antagonistic species interactions.

Taxonomic Synopsis of the Ponto-Mediterranean Ants of Temnothorax nylanderi Species-Group

In the current revisionary work, the Temnothorax nylanderi species-group of myrmicine ants is characterized. Eighteen species belonging to this group in the Ponto-Mediterranean region are described or redefined based on an integrative approach that combines exploratory analyses of morphometric data and of a 658bp fragment of the mitochondrial gene for the cytochrome c oxidase subunit I (CO I). The species group is subdivided into five species complexes: T. angustifrons complex, T. lichtensteini complex, T. nylanderi complex, T. parvulus complex, T. sordidulus complex, and two species, T. angulinodis sp. n. and T. flavicornis (Emery, 1870) form their own lineages. We describe seven new species (T. angulinodis sp. n., T. angustifrons sp. n., T. ariadnae sp. n., T. helenae sp. n., T. lucidus sp. n., T. similis sp. n., T. subtilis sp. n.), raise T. tergestinus (FINZI, 1928) stat.n. to species level, and propose a new junior synonymy for T. saxonicus (SEIFERT, 1995) syn.n. (junior synonym of T. tergestinus). We describe the worker caste and provide high quality images and distributional maps for all eighteen species. Furthermore, we provide a decision tree as an alternative identification key that visually gives an overview of this species-group. We make the first application to Formicidae of the Semantic Phenotype approach that has been used in previous taxonomic revisions.

Morphological variability of intercastes in the ant Temnothorax nylanderi: pattern of trait expression and modularity

Ants have distinct morphological castes (queens and workers), but aberrant queen-worker “intercastes” occasionally occur, both in wild and laboratory conditions. Intercastes are rare, however, such novel phenotypes may have evolutionary significance. Their morphology is highly variable in any given species, providing valuable information about the integration of queen traits (e.g. ocelli, wings, complex segmentation of thorax, large gaster and ovaries, spermatheca). Generally, these traits are all diminished or absent in workers. We used multivariate morphometry to analyze an exceptionally large sample of 101 intercastes of Temnothorax nylanderi. We determined distributions and correlations of traits, and confirmed the mosaic nature of intercastes. Queen-specific traits are not expressed coherently in intercastes, but the possible patterns of trait combination are limited. A large number of small-sized intercastes had disproportionately larger head, ocelli and gaster but smaller thorax. In contrast, queen-like growth of thorax and rudimentary wings only occurred in large-sized intercastes. This is the most comprehensive analysis of intercaste variability, and suggests the existence of constraints on recombination of caste-specific modular traits.

Starvation endurance in the ant Temnothorax nylanderi depends on group size, body size and access to larvae

Social interactions in animal groups can buffer environmental stress and may enhance survival under unfavourable conditions. In the present study, the impact on starvation endurance of social group, access to larvae and cold shock is studied in the ant Temnothorax nylanderi Förster. Resource sharing is expected to lead to grouped workers surviving longer than isolated ones. Access to larvae may increase longevity if larvae serve as food, or may interfere with survival if they induce caring behaviour in workers. Cold shock serves as a stress factor and a negative influence on survival is expected. The results show that isolated workers have a shorter lifespan than grouped workers, which in turn live for a shorter period than grouped workers with larvae. Beneficial ‘group effects’ contribute to group survival and the presence of larvae increases worker survival because the workers presumably feed on the larvae. Thus, improved starvation endurance may reflect an additional benefit of a social lifestyle. Moreover, variance in survival is lower for grouped workers than for isolated workers: group members not only demonstrate improved survival, but also smaller within‐group differences. Although a negative influence on survival is the expected outcome, this type of thermal stress is found to have no direct impact on starvation endurance other than moderating the differences between isolated and grouped workers.

Cold resistance depends on acclimation and behavioral caste in a temperate ant

Adjusting to low temperatures is important for animals living in cold environments. We studied the chill-coma recovery time in temperate ant workers (Temnothorax nylanderi) from colonies collected in autumn and spring in Germany. We experimentally acclimated these ant colonies to cold temperatures followed by warm temperatures. As expected, cold-acclimated workers recovered faster from freezing temperatures, but subsequent heat acclimation did not change the short recovery times observed after cold acclimation. Hence, either heat acclimation improves cold tolerance, possibly as a general response to stress, or at least it does not negate enhanced cold tolerance following cold acclimation. Colonies collected in spring showed similar cold tolerance levels to cold-acclimated colonies in the laboratory. Next, we compared the chill-coma recovery time of different worker castes and found that exterior workers recovered faster than interior workers. This difference may be related to their more frequent exposure to cold, higher activity level, or distinct physiology. Interior workers were also heavier and showed a higher gaster-to-head ratio and thorax ratio compared to exterior workers. An obvious difference between exterior and interior workers is activity level, but we found no link between activity and cold tolerance. This suggests that physiology rather than behavioral differences could cause the increased cold tolerance of exterior workers. Our study reveals the importance of acclimation for cold tolerance under natural and standardized conditions and demonstrates differences in cold tolerance and body dimensions in monomorphic behavioral castes of an ant.

Diaspore Trait Preferences of Dispersing Ants

Elaiosomes of myrmecochorous plant seeds are known to enhance the attraction of diaspore-dispersing ants by serving as a nutritional reward. However, it remained unclear which (nutritional) compounds affect diaspore preferences of ants. We hypothesized that apart from elaiosome/seed-size ratio, volume, and physical surface of diaspores, the quantity and the composition of fatty acids, amino acids, and sugars strongly influence the diaspore preferences of different species. Chemical (nutritional) profiles as well as structural properties of seeds with and without elaiosomes were analyzed and correlated with observed seed choice behavior of ants. Cafeteria experiments in the field confirmed the enhanced attractiveness of elaiosome-bearing seeds for all three ant species tested (Lasius fuliginosus, Myrmica ruginodis, and Temnothorax nylanderi), although seeds lacking elaiosomes also were transported. In multiple-choice cafeteria experiments with simultaneously offered diaspores of 16 plant species with and without elaiosome and with highly varying structural and chemical properties, all three ant species showed distinct preferences for certain diaspore species. Correlation analyses confirmed that the presence of an elaiosome represents the crucial factor that favors ant diaspore dispersal. In addition, the composition and the content of free amino acids, and to varying degrees fatty acids, were found to significantly affect preferences of each ant species, whereas the effect of single fatty acids acting as chemical triggers for diaspore transport by ants, as supposed by several studies, was not confirmed. In conclusion, although at least some diaspore species lacking elaiosomes attract ants for diaspore removal services by presenting nutritional seed coats, the production of elaiosomes seems to provide a worthwhile investment. Elaiosomes ensure rapid diaspore detection and removal due to chemical cue compounds and by offering a highly nutritional food supply, probably fitting the nutritional demands of ants.

Characterizing the Collective Personality of Ant Societies: Aggressive Colonies Do Not Abandon Their Home

Animal groups can show consistent behaviors or personalities just like solitary animals. We studied the collective behavior of Temnothorax nylanderi ant colonies, including consistency in behavior and correlations between different behavioral traits. We focused on four collective behaviors (aggression against intruders, nest relocation, removal of infected corpses and nest reconstruction) and also tested for links to the immune defense level of a colony and a fitness component (per-capita productivity). Behaviors leading to an increased exposure of ants to micro-parasites were expected to be positively associated with immune defense measures and indeed colonies that often relocated to other nest sites showed increased immune defense levels. Besides, colonies that responded with low aggression to intruders or failed to remove infected corpses, showed a higher likelihood to move to a new nest site. This resembles the trade-off between aggression and relocation often observed in solitary animals. Finally, one of the behaviors, nest reconstruction, was positively linked to per-capita productivity, whereas other colony-level behaviors, such as aggression against intruders, showed no association, albeit all behaviors were expected to be important for fitness under field conditions. In summary, our study shows that ant societies exhibit complex personalities that can be associated to the physiology and fitness of the colony. Some of these behaviors are linked in suites of correlated behaviors, similar to personalities of solitary animals.

Cuticular hydrocarbons in two parapatric species of ants and their hybrid

Discrimination between nestmates and non-nestmates in social insects is thought to rely on the pattern of cuticular hydrocarbons. We investigated the cuticular hydrocarbon profiles of 2 parapatric sibling ant species, Temnothorax nylanderi (Förster, 1850) and Temnothorax crassispinus (Karavaiev, 1926), and their hybrid. We found that although the profiles show considerable similarities, a discriminant analysis based on the relative peak areas of cuticular hydrocarbons separates the 3 taxa. The profiles of hybrids were not consistently intermediate between those of the paternal species, suggesting either non-additive interactions among the parental biosynthetic pathways or systematic differences in environment-derived odor cues.

How ants find each other; temporal and spatial patterns in nuptial flights

Reproduction is a key factor in understanding population ecology and therefore species occurrence. However, patterns in reproductive behaviour for distinct ant species remain insufficiently known. In this paper strategies in mate finding are studied for six ant species (Lasius niger, Lasius umbratus, Temnothorax nylanderi, Myrmica rubra, Myrmica ruginodis, Stenamma debile) in a forest – forest edge – agricultural field gradient. Using window traps, we studied whether these species had a restricted nuptial flight season, displayed swarming behaviour, and whether the alates aggregated at the forest edge. The flight season was limited to one month or less for L. niger, T. nylanderi, M. rubra, M. ruginodis and S. debile. Swarming behaviour occurred in all but one (L. umbratus) species. Although none of the six species seemed to have highest nest density at the forest edge, three of them, M. rubra, M. ruginodis and S. debile, showed male aggregations there, indicating this to be the main reproduction site. This last finding could be due to a more suitable micro-climate, but most likely, edges are conspicuous land marks which are used by ants to meet mates. The behavioural patterns of ant sexuals at the forest edge can influence dispersal possibilities in fragmented landscapes, reproductive success and nest densities.

Hierarchy length in orphaned colonies of the ant Temnothorax nylanderi

Workers of the ant Temnothorax nylanderi form dominance orders in orphaned colonies in which only one or a few top-ranking workers begin to produce males from unfertilized eggs. Between one and 11 individuals initiated 80% of all aggression in 14 queenless colonies. As predicted from inclusive fitness models (Molet M, van Baalen M, Monnin T, Insectes Soc 52:247-256, 2005), hierarchy length was found to first increase with colony size and then to level off at larger worker numbers. The frequency and skew of aggression decreased with increasing size, indicating that rank orders are less pronounced in larger colonies.

Nestmate recognition and intraspecific chemical and genetic variation in Temnothorax ants

The evolutionary stability of social cooperation requires altruistic acts to be directed mainly towards related individuals. To maintain significant intragroup relatedness, social insects have evolved sophisticated recognition systems. We compared nestmate recognition abilities of three native European species of the ant genus Temnothorax in relation to variation in cuticular chemicals and intraspecific genetic diversity. Colony take-over and fusions are common in two sibling species T. nylanderi and T. crassispinus, but rare in T. unifasciatus, suggesting relatively inefficient nestmate recognition in the former species. Temnothorax nylanderi and T. crassispinus discriminated less effectively between nestmates and non-nestmates than T. unifasciatus. While chemical profiles of different colonies strongly overlapped in T. crassispinus and T. nylanderi, 97% of all T. unifasciatus samples were, in a discriminant analysis, correctly assigned to their respective colony. Chemical distances between colonies correlated with aggression levels in T. unifasciatus, but not in the other two species. Finally, preliminary genetic results indicated that haplotype divergence within T. nylanderi and T. crassispinus is remarkably low, whereas T. unifasciatus and other congeners are genetically more variable. In theory, loss of genetic variation, including depletion at loci involved in the production of cuticular hydrocarbons, could decrease colony recognition efficiency. We suggest that this might be the case in T. nylanderi and T. crassispinus, leading to colony fusions in the field. Despite their different life histories, these native species show similarities to invasive ants where genetic bottlenecks arising from founder effects are thought to result in a loss of colony boundaries.

Distribution and genetic divergence of two parapatric sibling ant species in Central Europe

The two sibling ant species Temnothorax nylanderi and Temnothorax crassispinus are widely distributed throughout deciduous forests in Europe. Their resemblance in morphology and similar ecological requirements suggest that they evolved from the same ancestral species in different glacial refugia and re-immigrated into Central Europe after the last ice age. Here, we show that the two species are parapatrically distributed in south-eastern Germany and hybridize along a narrow contact zone close to the continental divide. Phylogeographical data based on the mitochondrial genes cytochrome oxidase subunit I and cytochrome b suggest that the dominant haplotypes for T. nylanderi and T. crassispinus might have diverged already 1.5–2 Mya. Intraspecific variability is extremely low in both species, which might be explained by severe bottlenecks during rapid postglacial expansion into Central Europe.

Ecology, life history and resource allocation in the ant, Leptothorax nylanderi.

We aimed at identifying the causal basis of previously shown interrelations between demographic and genetic colony structure, ecological factors and split sex ratios in the ant, Leptothorax nylanderi. Colony-level variation in sex allocation was only partly explained by annual fluctuations during eight study years and by resource availability as indicated by sexual production of colonies. Allocation ratios were highly male-biased in dense populations with ephemeral nest sites and high frequencies of colonies containing several unrelated matrilines. Field observations and experimental manipulations showed that nest site limitation leads to such heterogeneous colonies. Laboratory experiments demonstrated that genetic heterogeneity directly causes male-biased investment, although relatedness asymmetry is not influenced by invasion of unrelated queens. The influence of genetic composition on allocation strategies might either be explained by negative feedback mechanisms connected with habitat saturation or by a lower efficiency of heterogeneous colonies. Our results thus demonstrate which factors other than variation in relatedness asymmetry can explain split sex ratios in ants. An empirical test of a model on reproductive allocation revealed on-going queen-worker conflict over colony growth and sexual reproduction. Workers controlled reproductive allocation, but queen-worker conflict ceased in large colonies with a high survival rate.

Behavior of Usurping Queens in Colonies of the Ant species Leptothorax nylanderi (Hymenoptera: Formicidae)

Animals can reduce their costs of parental care by exploiting the behavior of unrelated conspecifics. Such 'intraspecific parasitism' has been observed in the small ant species, Leptothorax nylanderi. This study aims to identify how parasite queens of Leptothorax nylanderi are integrated into established colonies. Both in natural multi-queen colonies and in manipulated laboratory colonies, workers were temporarily aggressive towards the usurper queen, which herself remained initially passive and inconspicuous but later attacked the resident queen, the workers and the brood. Queen-queen aggression led to the elimination of one of the two queens and the restoration of monogyny. Usurpation is not associated with size polymorphism in L nylanderi.

Intraspecific parasitism and split sex ratios in a monogynous and monandrous ant ( Leptothorax nylanderi )

Sex ratios were bimodally distributed in a population of the monogynous and monandrous ant Leptothorax nylanderi during each of 3 study years. The population-wide investment ratios suggested worker control of sex allocation. Nest-level variation in the proportional investment in virgin queens was not affected by the presence or absence of a queen and only slightly by collecting year, but was correlated with nest size, total sexual investment and, unexpectedly, with differences in nestmate relatedness: small, low-investment nests and nests with several worker lineages produced male-biased sex ratios. Colonies containing several worker lineages arise from usurpation of mature colonies by unrelated founding queens and the fusion of unrelated colonies under strong nest site limitation. In contrast to facultatively polygynous and polyandrous species of social insects, where workers can maximize their inclusive fitness by adjusting sex ratios according to the degree of relatedness asymmetry, workers in mixed colonies of L. nylanderi do not benefit from manipulating sex allocation, as here relatedness asymmetries appear to be the same as in homogeneous colonies.

Modification of morphological characters and cuticular compounds in worker ants Leptothorax nylanderi induced by endoparasites Anomotaenia brevis

Anomotaenia brevis (Cestoda, Cyclophyllidea) induces major changes in the morphological characters of the host ant, Leptothorax nylanderi (Hymenoptera, Formicidae): alteration of pigmentation, lowering of adult mean size, reduction of legs, eyes and head, enlargement of petiole. The presence of parasites in adult ants also modifies the quantity of cuticular compounds but not their quality. The parasite induces some changes in the synthesis/release of 13 cuticular hydrocarbons. The higher the number of parasites within a worker, the larger the quantitative changes in four cuticular hydrocarbons in comparison with normal ants. Such modifications (morphology, chemistry, behaviour) may explain the partial intolerance exerted by normal workers against the parasitized ants.

Interspecific hybridisations in natural populations of ants by example of a regional fauna (Hymenoptera, Formicidae)

The identification of hybrids by high-precision stereomicroscopy and chorological data is described in detail by example of the ant species Formica bruni and pressilabris. In a synopsis of heterogenous data, the overall hybrid frequency in the ant fauna of Central Europe is estimated. 17 of the 164 ant species of this region are demonstrated and further 2 species strongly suspected to hybridise. The low-frequency hybridisers, showing overall hybrid ratios < 3%, were native elements of the Central European fauna before the onset of human cultivation and experienced direct interspecific contact for longer periods of their natural history. They developed more effective mechanisms of reproductive isolation beginning at the prezygotic level. Extremely high (12-31%) local hybridisation ratios occurred in species that invaded the area after anthropogenic changes in landscape structure. The segregated distribution of invaders and autochthonous species in the precultural period apparently did not impose the need to evolve more effective mechanisms for reproductive isolation. Prezygotic mechanisms in particular are deficient. In local situations, 19% of Lasius jensi matings and 44% of Leptothorax albipennis matings leading to successful nest foundations were matings with heterospecific partners. Signs for a dissolution of interspecific phenotypic differences are not detectable in most of the species. The dispersal of hybrid genotypes is apparently inhibited. Factors that probably stabilise the genomic integrity of hybridizing parent species are: (a) inability to produce hybrid queens (in L. jensi×umbratus), (b) aneuploidy of F1 females with inability to perform a balanced meiosis (in hybrids of L. albipennis with 3 other species), and (c) selection against hybrids in the epigenetic environment of alleles (Leptothorax nylanderi×slavonicus). Extreme ratios of heterospecific matings would mean a dangerous drain of genetic material. A mechanism to reduce these losses is postulated, consisting of a 'cleptogamy' (theft of heterospecific sperm) by queens that missed a conspecific male, an establishment of a functioning colony of F1 hybrid workers, a depression of the rearing or fertility of hybrid queens, and a maximum production of own sons by the queen. This mechanism could explain the persistence of rare species (Lasius jensi or Leptothorax albipennis) in an environment of more populous heterospecific hybridisation partners.

Effects of Ant Species Composition on Seed Removal in Deciduous Forest in Eastern Europe

Ant species contribute differently to the removal of elaiosome-bearing seeds. Microhabitats on the deciduous forest floor differ in the composition of ant species; these differences may influence the dispersal of myrmecochorous plants. This study examined removal rates of Viola odorata seeds which were dispersed by foragers of four ant species Formica polyctena (FP), Lasius fuliginosus (LF), Myrmica rubra (MR). and Leptothorax nylanderi (LN). We observed (I) the visit rates of ants and removal rates of seeds at three microhabitats with different ant species composition: FP territory with MR and LN. LF territory with AIR and LN, AIR territory with LN; (2) the contribution of different ant species to the seed removal rates; (3) effects of microsite type on the duration of seed manipulation by ant workers; (4) number of seeds handled by workers of different ant species: (5) effects of learning and recruitment on seed removal rates in three ant species FP. LF, and MR. At each territory the main contribution to visit rates and to seed removal rates was made by the most dominant ant species. The duration of seed manipulation by AIR ants at both FP and LF territories was significantly shorter than that by FP and LF ants. Under the pressure of large LF and FP, the AIR workers left the seed depot twice as quickly as in the absence of other ant species. Thus, duration of seed manipulation by MR was significantly shorter at FP, and especially, at LF territories than at MR territory. In all territories the duration of seed manipulation by LN was significantly longer than by other ant species. Learning (MR) or recruitment (FP) took place during seed removal. There were no differences in the duration of seed manipulation between experienced and inexperienced ants of both LF and FP. Inexperienced individuals of MR manipulated seeds twice as long as experienced ones. Experienced ants removed seeds with higher frequency than inexperienced ones regardless of ant species. It is concluded that ant species composition strongly influences the removal rates of seeds and potentially the fate of seeds of a myrmecochore. FP territory, providing the fastest removal of the fruit content, is presumably the most advantageous for the myrmecochore.

Mating frequency of Leptothorax nylanderi ant queens determined by microsatellite analysis

The queen mating frequency of the ant Leptothorax nylanderi was investigated by analyzing three (GA)n repeat microsatellite loci in queens and workers in 12 colonies from a single German population. Microsatellite genotypes indicate that all queens were inseminated by a single male. The genotypes of the males produced in queenright colonies suggest that they were offspring of the queen.