Geometrical multiscale tortuosity of desert ant walking trajectories.

Desert ants stand out as some of the most intriguing insect navigators, having captured the attention of scientists for decades. This includes the structure of walking trajectories during goal approach and search behaviour for the nest and familiar feeding sites. In the present study, we analysed such trajectories with regard to changes in walking direction. The directional change of the ants was quantified, i.e. an angle θ between trajectory increments of a given arclength λ was computed. This was done for different length scales λ, according to our goal of analysing desert ant path characteristics with respect to length scale. First, varying λ through more than two orders of magnitude demonstrated Brownian motion characteristics typical of the random walk component of search behaviour. Unexpectedly, this random walk component was also present in - supposedly rather linear - approach trajectories. Second, there were small but notable deviations from a uniform angle distribution that is characteristic of random walks. This was true for specific search situations, mostly close to the (virtual) goal position. And third, experience with a feeder position resulted in straighter approaches and more focused searches, which was also true for nest searches, albeit to a lesser extent. Taken together, these results both verify and extend previous studies on desert ant path characteristics. Of particular interest are the ubiquitous Brownian motion signatures and specific deviations thereof close to the goal position, indicative of unexpectedly structured search behaviour.

Geometrical multiscale tortuosity of desert ant walking trajectories.

Desert ants stand out as some of the most intriguing insect navigators, having captured the attention of scientists for decades. This includes the structure of walking trajectories during goal approach and search behaviour for the nest and familiar feeding sites. In the present study, we analysed such trajectories with regard to changes in walking direction. The directional change of the ants was quantified, i.e. an angle θ between trajectory increments of a given arclength λ was computed. This was done for different length scales λ, according to our goal of analysing desert ant path characteristics with respect to length scale. First, varying λ through more than two orders of magnitude demonstrated Brownian motion characteristics typical of the random walk component of search behaviour. Unexpectedly, this random walk component was also present in - supposedly rather linear - approach trajectories. Second, there were small but notable deviations from a uniform angle distribution that is characteristic of random walks. This was true for specific search situations, mostly close to the (virtual) goal position. And third, experience with a feeder position resulted in straighter approaches and more focused searches, which was also true for nest searches, albeit to a lesser extent. Taken together, these results both verify and extend previous studies on desert ant path characteristics. Of particular interest are the ubiquitous Brownian motion signatures and specific deviations thereof close to the goal position, indicative of unexpectedly structured search behaviour.

Flupyradifurone negatively affects survival, physical condition and mobility in the two-spotted lady beetle (Adalia bipunctata).

Lady beetles play a crucial role in natural ecosystems and agricultural settings. Unfortunately, these insects and more specifically the two-spotted lady beetle (Adalia bipunctata) are currently facing a severe decline in populations due to various stressors, with pesticide exposure being a significant threat. Flupyradifurone is a relatively newly introduced insecticide and as existing research is mainly elucidating its effects on bees there remains a limited understanding of its effects on non-hymenopteran insects, including lady beetles. In this study we investigated the impact of acute orally applied flupyradifurone doses on survival and sublethal parameters such as physical condition and mobility on A. bipunctata. Our findings revealed a significant increase in mortality among individuals subjected to flupyradifurone doses of 19 ng/individual (corresponding to >1.5-2.0 ng active substance (a.s.)/mg body weight (bw). The calculated LD50 of flupyradifurone at 48 h was 2.11 ng a.s./mg bw corresponding to an amount of 26.38 ng/individual. Sublethal consequences were observable immediately after pesticide application. Even at doses as low as 2 ng/individual (corresponding to >0.0-0.5 ng a.s./mg bw), flupyradifurone induced trembling and temporary immobility in treated animals. Furthermore, pesticide intoxication led to hypoactivity, with less distance covered and a decline in straightness of locomotion. In conclusion, our study underscores the harmful effects of flupyradifurone on the two-spotted lady beetle at doses notably lower than those affecting bees. These findings stress the importance of additional research to attain a more holistic understanding of pesticide impacts not only on a broader range of non-target arthropods species, but also on various exposure routes as well as lethal and sublethal effects.

Influence of the pesticide flupyradifurone on mobility and physical condition of larval green lacewings.

Global pesticide use in agriculture is one reason for the rapid insect decline in recent years. The relatively new pesticide flupyradifurone is neurotoxic to pest insects but considered harmless to bees according to previous risk assessments. With this study, we aim to investigate lethal and sublethal effects of flupyradifurone on larvae of the beneficial arthropod Chrysoperla carnea. We treated the animals orally with field-realistic concentrations of flupyradifurone and examined lethality as well as effects on condition, mobility and locomotion. For the lethal dose 50, we determined a value of > 120-200 ng/mg (corresponding to a mean amount of 219 ng/larva) after 168 h. Abnormal behaviors such as trembling and comatose larvae were observed even at the lowest concentration applied (> 0-20 ng/mg, 59 ng/larva). Mobility analysis showed impaired activity patterns, resulting in acute hypoactivity at all pesticide concentrations and time-delayed hyperactivity in larvae treated with > 40-60 ng/mg (100 ng/larva) and > 80-100 ng/mg (120 ng/larva), respectively. Even locomotion as a fundamental behavioral task was negatively influenced throughout larval development. In conclusion, our results demonstrate that flupyradifurone impacts life and survival of lacewing larvae and may pose-despite its status as bee-friendly-a major threat to insect fauna and environment.

Mechanosensory pathways of scorpion pecten hair sensillae-Adjustment of body height and pecten position.

Scorpions' sensory abilities are intriguing, especially the rather enigmatic ventral comb-like chemo- and mechanosensory organs, the so-called pectines. Attached ventrally to the second mesosomal segment just posterior to the coxae of the fourth walking leg pair, the pectines consist of the lamellae, the fulcra, and a variable number of pecten teeth. The latter contain the bimodal peg sensillae, used for probing the substrate with regard to chemo- and mechanosensory cues simultaneously. In addition, the lamellae, the fulcra and the pecten teeth are equipped with pecten hair sensillae (PHS) to gather mechanosensory information. Previously, we have analyzed the neuronal pathway associated with the peg sensillae unraveling their somatotopic projection pattern in dedicated pecten neuropils. Little is known, however, regarding the projections of PHS within the scorpion nervous system. Behavioral and electrophysiological assays showed involvement of PHS in reflexive responses but how the information is integrated remains unresolved. Here, we unravel the innervation pattern of the mechanosensory pecten hair afferents in Mesobuthus eupeus and Euscorpius italicus. By using immunofluorescent labeling and injection of Neurobiotin tracer, we identify extensive arborizations of afferents, including (i) ventral neuropils, (ii) somatotopically organized multisegmental sensory tracts, (iii) contralateral branches via commissures, and (iv) direct ipsilateral innervation of walking leg neuromeres 3 and 4. Our results suggest that PHS function as sensors to elicit reflexive adjustment of body height and obstacle avoidance, mediating accurate pecten teeth alignment to guarantee functionality of pectines, which are involved in fundamental capacities like mating or navigation.

Influence of caste and subcaste characteristics on locomotion in the ant Camponotus fellah.

Worker polymorphism in ants has evolved repeatedly, with considerable differences in the morphometry of worker subcastes. Such body size differences and especially caste- and subcaste-specific characteristics might significantly influence locomotion. Therefore, we performed a comprehensive locomotion analysis along gradients in both body size and walking speed of Camponotus fellah worker subcastes, and of males, which have rarely been studied to date because of their short life spans associated with mating flights. We provide a detailed description of the morphometry and size differences of C. fellah castes and subcastes and analyse locomotion in the different polymorphic groups in terms of absolute and relative walking speeds (mesosoma lengths per second). Our results reveal that body size and shape affect locomotion behaviour to different extents in the worker subcastes (minor workers, medias, major workers) and in males. Nevertheless, C. fellah ants use the same overall locomotion strategy, with males and major workers reaching considerably lower walking speeds than minors and medias. Body size thus mainly affects walking speed. Minor workers reach the highest relative velocities by high relative stride lengths in combination with large vertical and lateral centre of mass oscillations and clearly higher stride frequencies of up to 25 Hz. Locomotion of males was characterised by clearly lower walking speeds, wider footprint positions, significant phase shifts and a notable dragging of the shorter hind legs. However, general walking parameters of males differed less from those of the female workers than expected as a result of division of labour in the colony.

Locomotion in the pseudoscorpion Chelifer cancroides: forward, backward and upside-down walking in an eight-legged arthropod.

While insect locomotion has been intensively studied, there are comparably few studies investigating octopedal walking behaviour, and very little is known about pseudoscorpions in particular. Therefore, we performed an extensive locomotion analysis during forward, backward and upside-down walking in the cosmopolitan pseudoscorpion Chelifer cancroides. During forward locomotion, we observed C. cancroides to freeze locomotion frequently for short time periods. These microstops were barely visible to the naked eye with a duration of 100-200 ms. Our locomotion analysis revealed that C. cancroides performs a statically stable and highly coordinated alternating tetrapod gait during forward and backward walking, with almost complete inversion of the tetrapod schemes, but no rigidly fixed leg coordination during upside-down walks with low walking speeds up to 4 body lengths per second. Highest speeds (up to 17 body lengths per second), mainly achieved by consistent leg coordination and strong phase shifts, were observed during backward locomotion (escape behaviour), whereas forward walking was characterised by lower speeds and phase shifts of ∼10% between two loosely coupled leg groups within one tetrapod. That is, during the movement of one tetrapod group, the last and the third leg are almost synchronous in their swing phases, as are the second and the first leg. A special role of the second leg pair was demonstrated, probably mainly for stability reasons and related to the large pedipalps.

New records of soldier flies from Iran with a current checklist and taxonomic notes to some species (Diptera: Stratiomyidae).

Soldier fly records of Iran belonging to six species are presented, which includes the first record of Odontomyia annulata (Meigen, 1822) and another record of Adoxomyia cinerascens (Loew, 1873). Odontomyia annulata shows some morphological differences from the description found in the literature. With the addition of O. annulata, the Iranian soldier fly fauna comprises now 48 species. The eastern border of the known range of this species extended to the eastern Black Sea coast and the north-western Caucasus region. Now it has been shifted about 1200 km to the east.

Allometry in desert ant locomotion (Cataglyphis albicans and Cataglyphis bicolor) - does body size matter?

Desert ants show a large range of adaptations to their habitats. They can reach extremely high running speeds, for example, to shorten heat stress during foraging trips. It has recently been examined how fast walking speeds are achieved in different desert ant species. It is intriguing in this context that some species exhibit distinct intraspecific size differences. We therefore performed a complete locomotion analysis over the entire size spectrum of the species Cataglyphis bicolor, and we compared this intraspecific dataset with that of the allometrically similar species Cataglyphis albicans. Emphasis was on the allometry of locomotion: we considered the body size of each animal and analysed the data in terms of relative walking speed. Body size was observed to affect walking parameters, gait patterns and phase relationships in terms of absolute walking speed. Unexpectedly, on a relative scale, all ants tended to show the same overall locomotion strategy at low walking speeds, and significant differences occurred only between C. albicans and C. bicolor at high walking speeds. Our analysis revealed that C. bicolor ants use the same overall strategy across all body sizes, with small ants reaching the highest walking speeds (up to 80 body lengths s-1) by increasing their stride length and incorporating aerial phases. By comparison, C. albicans reached high walking speeds mainly by a high synchrony of leg movement, lower swing phase duration and higher stride frequency ranging up to 40 Hz.

Structure of the pecten neuropil pathway and its innervation by bimodal peg afferents in two scorpion species.

The pectines of scorpions are comb-like structures, located ventrally behind the fourth walking legs and consisting of variable numbers of teeth, or pegs, which contain thousands of bimodal peg sensillae. The associated neuropils are situated ventrally in the synganglion, extending between the second and fourth walking leg neuromeres. While the general morphology is consistent among scorpions, taxon-specific differences in pecten and neuropil structure remain elusive but are crucial for a better understanding of chemosensory processing. We analysed two scorpion species (Mesobuthus eupeus and Heterometrus petersii) regarding their pecten neuropil anatomy and the respective peg afferent innervation with anterograde and lipophilic tracing experiments, combined with immunohistochemistry and confocal laser-scanning microscopy. The pecten neuropils consisted of three subcompartments: a posterior pecten neuropil, an anterior pecten neuropil and a hitherto unknown accessory pecten neuropil. These subregions exhibited taxon-specific variations with regard to compartmentalisation and structure. Most notable were structural differences in the anterior pecten neuropils that ranged from ovoid shape and strong fragmentation in Heterometrus petersii to elongated shape with little compartmentalisation in Mesobuthus eupeus. Labelling the afferents of distinct pegs revealed a topographic organisation of the bimodal projections along a medio-lateral axis. At the same time, all subregions along the posterior-anterior axis were innervated by a single peg's afferents. The somatotopic projection pattern of bimodal sensillae appears to be common among arachnids, including scorpions. This includes the structure and organisation of the respective neuropils and the somatotopic projection patterns of chemosensory afferents. Nonetheless, the scorpion pecten pathway exhibits unique features, e.g. glomerular compartmentalisation superimposed on somatotopy, that are assumed to allow high resolution of substrate-borne chemical gradients.

Arthropod spatial cognition.

The feats of arthropods, and of the well-studied insects and crustaceans in particular, have fascinated scientists and laymen alike for centuries. Arthropods show a diverse repertoire of cognitive feats, of often unexpected sophistication. Despite their smaller brains and resulting lower neuronal capacity, the cognitive abilities of arthropods are comparable to, or may even exceed, those of vertebrates, depending on the species compared. Miniature brains often provide parsimonious but smart solutions for complex behaviours or ecologically relevant problems. This makes arthropods inspiring subjects for basic research, bionics, and robotics. Investigations of arthropod spatial cognition have originally concentrated on the honeybee, an animal domesticated for several thousand years. Bees are easy to keep and handle, making this species amenable to experimental study. However, there are an estimated 5-10 million arthropod species worldwide, with a broad diversity of lifestyles, ecology, and cognitive abilities. This high diversity provides ample opportunity for comparative analyses. Comparative study, rather than focusing on single model species, is well suited to scrutinise the link between ecological niche, lifestyle, and cognitive competence. It also allows the discovery of general concepts that are transferable between distantly related groups of organisms. With species diversity and a comparative approach in mind, this special issue compiles four review articles and ten original research reports from a spectrum of arthropod species. These contributions range from the well-studied hymenopterans, and ants in particular, to chelicerates and crustaceans. They thus present a broad spectrum of glimpses into current research on arthropod spatial cognition, and together they cogently emphasise the merits of research into arthropod cognitive achievements.

Accuracy and spread of nest search behaviour in the Saharan silver ant, Cataglyphis bombycina, and in the salt pan species, Cataglyphis fortis.

Desert ants of the genus Cataglyphis are renowned for their navigation abilities, especially for their beeline homing after meandering foraging excursions reaching several hundreds of meters in length. A spiralling nest search is performed when an ant misses the nest entrance upon completing its homebound travel. We examined the nest search behaviours of two desert ant species dwelling in different habitats-Cataglyphis bombycina living in the dunes of the Sahara and Cataglyphis fortis found in the salt pans of North Africa. The two species show distinct differences in walking behaviour. C. bombycina performs a strict tripod gait with pronounced aerial phases, high stride frequencies, and extremely brief ground contact times. In view of these peculiarities and the yielding sand dune substrate, we hypothesised that homing accuracy, and namely distance measurement by stride integration, should be lower in C. bombycina, compared to the well-studied C. fortis with less specialised walking behaviour. We tested this hypothesis in ants' homebound runs from a feeding site in a linear channel setup. Surprisingly, the accuracies of nest searches were similar in the two ant species, and search accuracy was also independent of the walking substrate, soft dune sand or a hard floor. The spread of the nest search, by contrast, differed significantly between the two species, C. bombycina exhibiting a larger search spread. This may be interpreted as an increased path integration uncertainty due to the above locomotor specialisations, or as a compensation strategy accounting for the silver ants' particular environmental and behavioural situation.

The impact of implant material and patient age on the long-term outcome of secondary cranioplasty following decompressive craniectomy for severe traumatic brain injury.

BACKGROUND: Secondary cranioplasty (CP) is considered to support the neurological recovery of patients after decompressive craniectomy (DC), but the treatment success might be limited by complications associated to confounders, which are not yet fully characterized. The aim of this study was to identify the most relevant factors based on the necessity to perform revision surgeries. METHODS: Data from 156 patients who received secondary CP following DC for severe traumatic brain injury (TBI) between 1984 and 2015 have been retrospectively analyzed and arranged into cohorts according to the occurrence of complications requiring surgical intervention. RESULTS: Cox regression analysis revealed a lower revision rate in patients with polymethylmethacrylate (PMMA) implants than in patients with autologous calvarial bone (ACB) implants (HR 0.2, 95% CI 0.1 to 1.0, p = 0.04). A similar effect could be observed in the population of patients aged between 18 and 65 years, who had a lower risk to suffer complications requiring surgical treatment than individuals aged under 18 or over 65 years (HR 0.4, 95% CI 0.2 to 0.9, p = 0.02). Revision rates were not influenced by the gender (p = 0.88), timing of the CP (p = 0.53), the severity of the TBI (p = 0.86), or the size of the cranial defect (p = 0.16). CONCLUSIONS: In this study, the implant material and patient age were identified as the most relevant parameters independently predicting the long-term outcome of secondary CP. The use of PMMA was associated with lower revision rates than ACB and might provide a therapeutic benefit for selected patients with traumatic cranial defects.

High-speed locomotion in the Saharan silver ant, Cataglyphis bombycina.

The diurnal thermophilic Saharan silver ant, Cataglyphis bombycina, is the fastest of the North African Cataglyphis desert ant species. These highly mobile ants endure the extreme temperatures of their sand dune environment with outstanding behavioural, physiological and morphological adaptations. Surprisingly, C. bombycina has comparatively shorter legs than its well-studied sister species Cataglyphis fortis from salt pan habitats. This holds despite the somewhat hotter surface temperatures and the more yielding sand substrate. Here, we report that C. bombycina employs a different strategy in reaching high running speeds, outperforming the fastest known runs of the longer-legged C. fortis ants. Video analysis across a broad range of locomotor speeds revealed several differences to C. fortis Shorter leg lengths are compensated for by high stride frequencies, ranging beyond 40 Hz. This is mainly achieved by a combination of short stance phases (down to 7 ms) and fast leg swing movements (up to 1400 mm s-1). The legs of one tripod group exhibit almost perfect synchrony in the timings of their lift-offs and touch-downs, and good tripod coordination is present over the entire walking speed range (tripod coordination strength values around 0.8). This near synchrony in leg movement may facilitate locomotion across the yielding sand dune substrate.

Primary processing neuropils associated with the malleoli of camel spiders (Arachnida, Solifugae): a re-evaluation of axonal pathways.

BACKGROUND: Arachnids possess highly specialized and unorthodox sense organs, such as the unique pectines of Scorpiones and the malleoli of Solifugae. While the external morphology, numbers, and shapes of sensory organs are widely used in taxonomic studies, little is known about the internal anatomy of these organs and their associated processing neuropils in the central nervous system. Camel spiders (Solifugae) possess pedipalps and first walking legs heavily endowed with sensory structures, as well as conspicuous malleoli located ventrally on the proximal fourth walking legs. Malleoli are fan-shaped organs that contain tens of thousands of presumptive chemoreceptor neurons, but mechanoreceptive structures are absent. RESULTS: Here, we examine the organization of the synganglion based on microCT analysis, 3D reconstruction of serial paraffin sections, and backfill preparations to trace the malleolar pathway. The projection area of malleolar afferents is intriguingly located in the most anterior ventral nerve cord, located in between the pedipalpal neuromere hemispheres. However, malleolar axon bundles are separated by a thin soma layer that points to an anteriad projection of the fourth walking leg neuromere. A conspicuous projection neuron tract that may receive additional input from pedipalpal sensory organs connects the malleolar neuropil with the mushroom bodies in the protocerebrum. CONCLUSION: Arthropod chemosensory appendages or organs and primary processing neuropils are typically located in the same segment, which also holds true in Solifugae, although the malleolar neuropil is partially shifted towards the pedipalpal neuromere. A comparison of the malleoli in Solifugae and the pectines in Scorpiones, and of their primary processing neuropils, reveals certain similarities, while striking differences are also evident. Similarities include the ventral arrangement of peg-shaped sensory structures on the respective segmental appendage, exposing dense arrays of chemoreceptive sensilla, and projections to a primary processing neuropil with glomerular subdivision. Differences are, e.g., the lack of mechanoreceptive afferents and an associated processing neuropil.

Two distance memories in desert ants-Modes of interaction.

Navigation plays an essential role for many animals leading a mobile mode of life, and for central place foragers in particular. One important prerequisite for navigation is the ability to estimate distances covered during locomotion. It has been shown that Cataglyphis desert ants, well-established model organisms in insect navigation, use two odometer mechanisms, namely, stride and optic flow integration. Although both mechanisms are well established, their mode of interaction to build one odometer output remains enigmatic. We tackle this problem by selectively covering the ventral eye parts in Cataglyphis fortis foragers, the eye regions responsible for optic flow input in odometry. Exclusion of optic flow cues was implemented during different sections of outbound and inbound travel. This demonstrated that the two odometers have separate distance memories that interact in determining homing distance. Possible interpretations posit that the two odometer memories (i) take on different relative weights according to context or (ii) compete in a winner-take-all mode. Explanatory values and implications of such interpretations are discussed. We are able to provide a rough quantitative assessment of odometer cue interaction. An understanding of the interaction of different odometer mechanisms appears valuable not only for animal navigation research but may inform discussions on sensor fusion in both behavioural contexts and potential technical applications.

Scorpions pectines - Idiosyncratic chemo- and mechanosensory organs.

Scorpions possess specialised chemosensory appendages, the pectines. These comb-shaped limbs are located ventrally behind the walking legs. Like the antennae of mandibulate arthropods, they also serve a mechanosensory function. However, more than 90% of the sometimes well above 100,000 sensory neurons projecting from a pectine to the central nervous system are chemosensory. There are two primary projection neuropils. The posterior one, immediately adjacent to the pectine nerve entrance, has an intriguing substructure reminiscent of the olfactory glomeruli observed in the primary chemosensory neuropils of many arthropods and indeed of most bilaterian animals. There are further similarities, particularly to the antennal lobes of mandibulate arthropods, including dense innervation by a relatively small number of putative serotonergic interneurons and the presence of GABA immunoreactivity, indicative of inhibitory interactions. Scorpion idiosyncrasies include the flattened shape and broad size range of the glomerulus-like neuropil compartments. Further, these compartments are often not clearly delimited and form layers in the neuropil that are arranged like onion peels. In summary, the pectine appendages of scorpions and their central nervous projections appear as promising study subjects, particularly regarding comparative examination of chemosensory representation and processing strategies. The possibility of combined, rather than discrete, representations of chemo- and mechanosensory inputs should merit further study.

Clinical outcome and osteoarthritic changes after surgical treatment of isolated capitulum humeri fractures with a minimum follow-up of five years.

PURPOSE: The aim of this study was to evaluate the functional outcome of patients treated for a fracture of the capitulum humeri and to analyze the grade of osteoarthritic changes. METHODS: Patients undergoing surgical reconstruction of isolated capitulum humeri fractures were included in a retrospective comparative analysis. Fractures were classified according to the Arbeitsgemeinschaft für Osteosynthesefragen (AO) and Dubberley and were functionally evaluated by the American Shoulder and Elbow Surgeons Score (ASES) and the Mayo Elbow Performance Index (MEPI). Levels of arthritic changes were measured with the Broberg and Morrey Score. Frequency of complications was evaluated whereby all results were compared to the ipsilateral, unaffected side. RESULTS: Thirteen patients, ten females (76.9 %) and three males (23.1 %), with a mean age of 48.7 ± 13.3 years were included in the study. Long-term follow-up range of motion (ROM) in the sagittal plane was significantly influenced by time of surgery (p < 0.001), and long-term follow-up with respect to pronation and supination by ROM of the healthy control (p < 0.05). The average ASES score was 37.8, and the Mayo Elbow Performance Index (MEPI) was 92.7. The mean level of degenerative arthritic changes was 1.9 ± 0.6 on the fractured side and significantly less (0.8 ± 0.8) on the healthy side (pF-test < 0.001). At a mean follow-up of 118.5 ± 52.4 months neither nonunion nor avascular necrosis were observed in any case. However, six cases of heterotopic ossification were identified. CONCLUSIONS: Satisfying functional outcomes and a low rate of osteoarthritic changes can be expected after the presented open reduction and internal fixation of capitulum humeri fractures. LEVEL OF EVIDENCE: Level IV - Case series; therapeutic study.

Nest and food search behaviour in desert ants, Cataglyphis: a critical comparison.

North African desert ants, Cataglyphis, use path integration to calculate a home vector during their foraging trips, constantly informing them about their position relative to the nest. This home vector is also used to find the way back to a productive feeding site the ant has encountered and thus memorized. When the animal fails to arrive at its goal after having run off the home or food vector, a systematic search is initiated. The basic search strategies are identical for nest and food searches, consisting of a search spiral superimposed by a random walk. While nest searches have been investigated in much detail, food site searches have received comparatively little attention. Here, we quantify and compare nest and food site searches recorded under similar conditions, particularly constant nest-feeder distance, and we observe notable differences in nest and food search performances. The parameters of nest searches are relatively constant and improve little with experience, although those small improvements had not been recognized previously. Food searches, by contrast, are more flexible and cover smaller or larger areas, mainly depending on the reliability of food encounter over several visits. Intriguingly, food site searches may be significantly more focussed than nest searches, although the nest should be the most important goal in an ant's life. These results demonstrate both adaptability and high accuracy of the ants' search programme.

Food searches and guiding structures in North African desert ants, Cataglyphis.

North African desert ants, Cataglyphis fortis, use path integration as their primary means of navigation. The ants also use landmarks when these are available to improve navigation accuracy. Extended landmarks, such as walls and channels, may serve further functions, for example, local guidance or triggering of local vectors. The roles of such structures were usually examined in homing animals but not during food searches. When searching for familiar feeding sites, Cataglyphis may show intriguing deviations from expected search performances. These may result from the presence of extended landmarks, namely experimental channels. Here we scrutinise this hypothesis of landmark guidance in food searches. We prevented the ants from seeing the channel walls by covering their eyes, except the dorsal rim area. This experiment was repeated in the open test field with an alley of black cylinders to extend our findings to a more normal foraging environment. Ants with covered eyes did not deviate from expected search performances, whereas ants with normal eyes extended their searches along the axis of the leading structures by 15-20%, in both channels and landmark alleys. This demonstrates that Cataglyphis orients along extended landmarks when searching for familiar food sources and alters its search pattern accordingly.