Spider dung beetles: coordinated cooperative transport without a predefined destination.

Cooperative transport allows for the transportation of items too large for the capacity of a single individual. Beyond humans, it is regularly employed by ants and social spiders where two or more individuals, with more or less coordinated movements, transport food to a known destination. In contrast to this, pairs of male and female dung beetles successfully transport brood balls to a location unknown to either party at the start of their common journey. We found that, when forced to overcome a series of obstacles in their path, transport efficiency of pairs of beetles was higher than of solo males. To climb tall obstacles with their common ball of dung, the female assisted the leading male in lifting the ball by steadying and pushing it upwards in a 'headstand' position during the climb initiation. Finally, we show that pairs were faster than single beetles in climbing obstacles of different heights. Our results suggest that pairs of Sisyphus beetles cooperate in the transportation of brood balls with coordinated movements, where the male steers and the female primarily assists in lifting the ball. Taken together, this is to our knowledge, the first quantitative study of cooperative food transport without a known goal to aim for.

The balbyter ant Camponotus fulvopilosus combines several navigational strategies to support homing when foraging in the close vicinity of its nest.

Many insects rely on path integration to define direct routes back to their nests. When shuttling hundreds of meters back and forth between a profitable foraging site and a nest, navigational errors accumulate unavoidably in this compass- and odometer-based system. In familiar terrain, terrestrial landmarks can be used to compensate for these errors and safely guide the insect back to its nest with pin-point precision. In this study, we investigated the homing strategies employed by Camponotus fulvopilosus ants when repeatedly foraging no more than 1.25 m away from their nest. Our results reveal that the return journeys of the ants, even when setting out from a feeder from which the ants could easily get home using landmark information alone, are initially guided by path integration. After a short run in the direction given by the home vector, the ants then switched strategies and started to steer according to the landmarks surrounding their nest. We conclude that even when foraging in the close vicinity of its nest, an ant still benefits from its path-integrated vector to direct the start of its return journey.

Mechanisms of spectral orientation in a diurnal dung beetle.

Ball rolling dung beetles use a wide range of cues to steer themselves along a fixed bearing, including the spectral gradient of scattered skylight that spans the sky. Here, we define the spectral sensitivity of the diurnal dung beetle Kheper lamarcki and use the information to explore the orientation performance under a range of spectral light combinations. We find that, when presented with spectrally diverse stimuli, the beetles primarily orient to the apparent brightness differences as perceived by their green photoreceptors. Under certain wavelength combinations, they also rely on spectral information to guide their movements, but the brightness and spectral directional information is never fully disentangled. Overall, our results suggest the use of a dichromatic, primitive colour vision system for the extraction of directional information from the celestial spectral gradient to support straight-line orientation. This article is part of the theme issue 'Understanding colour vision: molecular, physiological, neuronal and behavioural studies in arthropods'.

Mapping the distribution and tree canopy cover of Jacaranda mimosifolia and Platanus × acerifolia in Johannesburg's urban forest.

This study investigated the distribution and the tree canopy cover (TCC) of the two most prominent street trees (Jacaranda mimosifolia and Platanus × acerifolia) in Johannesburg, using the multispectral SPOT 6 satellite data and field survey GPS points. The importance of the spectral bands (Blue, Green, Red and NIR) and the NDVI index in discriminating between the tree species was quantified using five separability indices (Divergence, Bhattacharyya, Transformed Divergence, Jeffries-Matusita and M-statistic). The visual comparison of the Blue band and the NDVI histograms between the two species and other vegetation type showed the lowest feature overlap, suggesting the highest separability between paired classes. This was further supported by the highest Divergence value for the Blue band (3.68) and NDVI index (2.48) followed by the M-statistic (0.8 and 0.73, respectively) indicating good to moderate separability between the two species, respectively. The results were also consistent with the RF classification where the Blue band and NDVI index were the most important variables for the discrimination between the two species with an overall accuracy of 88% (kappa = 8). The TCC of J. mimosifolia and P. × acerifolia constituted 38% of the total vegetation cover in the city. These findings not only would help prioritize the increase of targeted vegetation cover in low cover areas, but will also provide a valuable information for assessment and protection of vulnerable species such as P. × acerifolia from the threat of the polyphagous shot hole borer, Euwallacea fornicatus in Johannesburg.

The interplay of directional information provided by unpolarised and polarised light in the heading direction network of the diurnal dung beetle Kheper lamarcki.

The sun is the most prominent source of directional information in the heading direction network of the diurnal, ball-rolling dung beetle Kheper lamarcki. If this celestial body is occluded from the beetle's field of view, the distribution of the relative weight between the directional cues that remain shifts in favour of the celestial pattern of polarised light. In this study, we continue to explore the interplay of the sun and polarisation pattern as directional cues in the heading direction network of K. lamarcki. By systematically altering the intensity and degree of the two cues, we effectively change the relative reliability as they appear to the dung beetle. The response of the beetle to these modifications allows us to closely examine how the weighting relationship of these two sources of directional information is influenced and altered in the heading direction network of the beetle. We conclude that the process by which K. lamarcki relies on directional information is very likely done based on Bayesian reasoning, where directional information conveying the highest certainty at a particular moment is afforded the greatest weight.

Cold-induced anesthesia impairs path integration memory in dung beetles.

Path integration is a general mechanism used by many animals to maintain an updated record of their position in relation to a set reference point.1-11 To do this, they continually integrate direction and distance information into a memorized home vector. What remains unclear is how this vector is stored, maintained, and utilized for successful navigation. A recent computational model based on the neuronal circuitry of the insect central complex suggests that home vector memories are encoded across a set of putative memory neurons and maintained through ongoing recurrent neural activity.12 To better understand the nature of the home vector memory and experimentally assess underlying mechanisms for maintaining it, we performed a series of experiments on the path integrating dung beetle Scarabaeus galenus.13 We found that, while the directional component of the home vector was maintained for up to 1 h, the distance component of the vector memory decreased gradually over time. Using cold-induced anesthesia, we disrupted the neural activity of beetles that had stored a home vector of known length and direction. This treatment diminished both components of the home vector memory, but by different amounts-the homing beetles lost their distance memory before their directional memory. Together, these findings present new insights into the functional properties of home vector memories and provide the first empirical evidence that a biological process that can be disrupted by cold-induced anesthesia is essential to support homing by path integration.

The finely defined shift work schedule of dung beetles and their eye morphology.

In nature, nothing is wasted, not even waste. Dung, composed of metabolic trash and leftovers of food, is a high-quality resource and the object of fierce competition. Over 800 dung beetle species (Scarabaeinae) compete in the South African dung habitat and more than 100 species can colonize a single dung pat. To coexist in the same space, using the same food, beetles divide the day between them. However, detailed diel activity periods and associated morphological adaptations have been largely overlooked in these dung-loving insects. To address this, we used a high-frequency trapping design to establish the diel activity period of 44 dung beetle species in their South Africa communities. This allowed us to conclude that the dung beetles show a highly refined temporal partitioning strategy, with differences in peak of activity even within the diurnal, crepuscular, and nocturnal guilds, independent of nesting behavior and taxonomic classification. We further analyzed differences in eye and body size of our 44 model species and describe their variability in external eye morphology. In general, nocturnal species are bigger than crepuscular and diurnal species, and as expected, the absolute and relative eye size is greatest in nocturnal species, followed by crepuscular and then diurnal species. A more surprising finding was that corneal structure (smooth or facetted) is influenced by the activity period of the species, appearing flat in the nocturnal species and highly curved in the diurnal species. The role of the canthus-a cuticular structure that partially or completely divides the dung beetle eye into dorsal and ventral parts-remains a mystery, but the large number of species investigated in this study nevertheless allowed us to reject any correlation between its presence and the nesting behavior or time of activity of the beetles.

Dataset, including a photo-guide, of alien plants sold in traditional medicine markets and healthcare outlets in three South African cities, specifically by traders of Indian, West African, East African, and Chinese origin.

This dataset is a an inventory of 475 alien plant taxa (447 identified to species), including a photo-guide to 96 plants, mostly sold as traditional medicines in three South African cities by traders of South African, West African, East African, Indian and Chinese origin (Williams et al., 2021). The dataset also incorporates species documented in a literature survey of alien plants used for traditional medicines in South Africa. The species inventory is a consolidation of the data from two separate investigations of 106 medicinal plant traders: firstly, a study conducted in 2010/2011 of 77 traders in markets and shops in Johannesburg, Pretoria and Durban (Williams et al., 2021); and secondly, a study conducted in 2017/2018 of plants sold by 29 (im)migrant traders of West African, East African, and Indian origin in Johannesburg, and of alien species listed in a TCM (Traditional Chinese Medicine) catalogue (Burness, 2019). Accompanying each plant photograph in the photo-guide is the following information: species name; common name(s) provided by the survey respondents; invasive alien plant category; introduction status; voucher specimen number(s); nationality of the medicine traders; and, notes on source localities (e.g. imported or collected in southern Africa). Overall, most of the taxa were from the Asteraceae (12%), Fabaceae (9%) and Poaceae (5%). The species are mostly unlisted (76%) with respect to their legal status in South Africa in terms of the National Environmental Management: Biodiversity Act (NEM:BA), 2004 Alien & Invasive Species (A&IS) regulations. The most frequently recorded species in the various surveys were Glycyrrhiza glabra, Acorus calamus, Angelica sinensis and Zingiber officinale.

Compass Cue Integration and Its Relation to the Visual Ecology of Three Tribes of Ball-Rolling Dung Beetles.

To guide their characteristic straight-line orientation away from the dung pile, ball-rolling dung beetles steer according to directional information provided by celestial cues, which, among the most relevant are the sun and polarised skylight. Most studies regarding the use of celestial cues and their influence on the orientation system of the diurnal ball-rolling beetle have been performed on beetles of the tribe Scarabaeini living in open habitats. These beetles steer primarily according to the directional information provided by the sun. In contrast, Sisyphus fasciculatus, a species from a different dung-beetle tribe (the Sisyphini) that lives in habitats with closely spaced trees and tall grass, relies predominantly on directional information from the celestial pattern of polarised light. To investigate the influence of visual ecology on the relative weight of these cues, we studied the orientation strategy of three different tribes of dung beetles (Scarabaeini, Sisyphini and Gymnopleurini) living within the same biome, but in different habitat types. We found that species within a tribe share the same orientation strategy, but that this strategy differs across the tribes; Scarabaeini, living in open habitats, attribute the greatest relative weight to the directional information from the sun; Sisyphini, living in closed habitats, mainly relies on directional information from polarised skylight; and Gymnopleurini, also living in open habitats, appear to weight both cues equally. We conclude that, despite exhibiting different body size, eye size and morphology, dung beetles nevertheless manage to solve the challenge of straight-line orientation by weighting visual cues that are particular to the habitat in which they are found. This system is however dynamic, allowing them to operate equally well even in the absence of the cue given the greatest relative weight by the particular species.

Elevated atmospheric CO2 adversely affects a dung beetle's development: Another potential driver of decline in insect numbers?

Insect declines have been attributed to several drivers such as habitat loss, climate change, invasive alien species and insecticides. However, in the global context, these effects remain patchy, whereas insect losses appear to be consistent worldwide. Increases in atmospheric CO2 concentrations are known to have indirect effects on herbivorous insects, but the effects on other insects are largely unexplored. We wondered if elevated atmospheric CO2 (eCO2 ) could influence the growth and survival of insects, not via rising temperature, nor through their changes in food quality, but by other means. Rearing tunnelling dung beetle Euoniticellus intermedius (Reiche, 1848) at pre-industrial (250 parts per million [ppm]), current (400 ppm) and eCO2 levels (600 and 800 ppm), we found that exposure to eCO2 resulted in longer developmental times and increased mortality. Elevated CO2 also caused reduction of adult size and mass which is detrimental to dung beetle fitness. Additional results showed associated increases in CO2 levels inside dung brood balls, dung pH and respiration rates of the soil surrounding the developing dung beetles (CO2 flux). We thus hypothesize that elevated CO2 increases competition for O2 and nutrients between soil microbiota and subterranean insects. Given that many insect orders spend at least part of their life underground, our findings indicate the possibility of a negative ubiquitous effect of eCO2 on a large portion of the earth's insect biota. These findings therefore suggest an important area for future research on the soil community in the context of atmospheric change.

Light pollution forces a change in dung beetle orientation behavior.

Increasing global light pollution1,2 threatens the night-time darkness to which most animals are adapted. Light pollution can have detrimental effects on behavior,3-5 including by disrupting the journeys of migratory birds,5,6 sand hoppers,7-9 and moths.10 This is particularly concerning, since many night-active species rely on compass information in the sky, including the moon,11,12 the skylight polarization pattern,13,14 and the stars,15 to hold their course. Even animals not directly exposed to streetlights and illuminated buildings may still experience indirect light pollution in the form of skyglow,3,4 which can extend far beyond urban areas.1,2 While some recent research used simulated light pollution to estimate how skyglow may affect orientation behavior,7-9 the consequences of authentic light pollution for celestial orientation have so far been neglected. Here, we present the results of behavioral experiments at light-polluted and dark-sky sites paired with photographic measurements of each environment. We find that light pollution obscures natural celestial cues and induces dramatic changes in dung beetle orientation behavior, forcing them to rely on bright earthbound beacons in place of their celestial compass. This change in behavior results in attraction toward artificial lights, thereby increasing inter-individual competition and reducing dispersal efficiency. For the many other species of insect, bird, and mammal that rely on the night sky for orientation and migration, these effects could dramatically hinder their vital night-time journeys.

How Dung Beetles Steer Straight.

Distant and predictable features in the environment make ideal compass cues to allow movement along a straight path. Ball-rolling dung beetles use a wide range of different signals in the day or night sky to steer themselves along a fixed bearing. These include the sun, the Milky Way, and the polarization pattern generated by the moon. Almost two decades of research into these remarkable creatures have shown that the dung beetle's compass is flexible and readily adapts to the cues available in its current surroundings. In the morning and afternoon, dung beetles use the sun to orient, but at midday, they prefer to use the wind, and at night or in a forest, they rely primarily on polarized skylight to maintain straight paths. We are just starting to understand the neuronal substrate underlying the dung beetle's compass and the mystery of why these beetles start each journey with a dance.

Feasible or foolish: Attempting restoration of a Parthenium hysterophorus invaded savanna using perennial grass seed.

The annual herb Parthenium hysterophorus L. (Asteraceae), remains one of Southern Africa's most significant invasive weeds, commonly invading savannas, and their rangelands, causing severe losses to agriculture, livestock production and native biodiversity. Previous studies have suggested that perennial grasses may act as useful competitive species, capable of suppressing the growth and invasion of P. hysterophorus. To explore this, a total of 48 plots were established within an invaded savanna, using a randomised block design, and included treatments with and without the clearing of P. hysterophorus, as well as with and without the sowing of native perennial grass seed (Anthephora pubescens, Chloris gayana, Cynodon dactylon, Digitaria eriantha, Eragrostis curvula, Panicum maximum and Themeda triandra). Plots were assessed yearly in terms of P. hysterophorus density and growth as well as grass species composition, basal cover, and biomass over a three-year period. Clearing alone was found to exacerbate invasion, increasing P. hysterophorus density by 40%. Whereas the sowing of grass seed, in both the cleared and uncleared plots, increased the abundance of perennial grass species by 28%, subsequently reducing the size, reproductive output and density of P. hysterophorus over the three years. In addition, these sowing efforts contributed towards partial restoration of the plots, enhancing grass basal cover by ~15% and biomass production by 17%. Overall, this research suggests that sowing of native grass species, with or without clearing, may be a useful supplementary control or restoration tool towards the long-term management of P. hysterophorus invasions in managed savannas and rangelands in Southern Africa.

Effects of cadmium toxicity on the physiology and growth of a halophytic plant, Tamarix usneoides (E. Mey. ex Bunge).

Heavy metal polluted soils can be remediated using plants, a process called phytoremediation. However, high concentrations of heavy metals can negatively affect plant physiology and growth. We experimentally evaluated the effects of cadmium (Cd) on the growth, (i.e. height, shoot and biomass) and physiology (i.e. leaf chlorophyll and relative water contents) of Tamarix usneoides. In a greenhouse experiment, T. usneoides clones were subjected to a once off treatment of 100 mmol/l NaCl with three different Cd concentrations (6, 12, and 18 mg/kg) applied 3 times/week for eight weeks. We predicted that plant health would decrease with an increase in Cd concentration. Results revealed a 35.9% reduction in chlorophyll content between the 18 mg/kg Cd treated plants and the control, suggesting that T. usneoides experienced a reduction in photosynthetic rate, which in turn influenced the growth and relative water content (RWC) of the plant. Although T. usneoides' growth and physiology were significantly decreased at 12 and 18 mg/kg Cd concentrations, the plants tolerated up to 6 mg/kg Cd concentration, a level found in most anthropogenic Cd-contaminated soils. Tamarix usneoides should thus be confirmed as a good phytoremediation candidate once its ability to extract, translocate and concentrate Cd has been determined.

A dung beetle that path integrates without the use of landmarks.

Unusual amongst dung beetles, Scarabaeus galenus digs a burrow that it provisions by making repeated trips to a nearby dung pile. Even more remarkable is that these beetles return home moving backwards, with a pellet of dung between their hind legs. Here, we explore the strategy that S. galenus uses to find its way home. We find that, like many other insects, they use path integration to calculate the direction and distance to their home. If they fail to locate their burrow, the beetles initiate a distinct looping search behaviour that starts with a characteristic sharp turn, we have called a 'turning point'. When homing beetles are passively displaced or transferred to an unfamiliar environment, they initiate a search at a point very close to the location of their fictive burrow-that is, a spot at the same relative distance and direction from the pick-up point as the original burrow. Unlike other insects, S. galenus do not appear to supplement estimates of the burrow location with landmark information. Thus, S. galenus represents a rare case of a consistently backward-homing animal that does not use landmarks to augment its path integration strategy.

Sexual Dimorphism in Anthonomus santacruzi (Coleoptera: Curculionidae): a Biological Control Agent of Solanum mauritianum Scopoli (Solanaceae).

There is evident variation in body size amongst Anthonomus santacruzi Hustache, 1924, weevils. The aims of this study were to assess if the variation in body size in A. santacruzi weevils is a result of sexual dimorphism and what features can be used to distinguish males from females. The weevils were collected from field sites in Mpumalanga, South Africa, where they were introduced as biocontrol agents of Solanum mauritianum Scopoli. Body structures and the presence/absence of the tergal notch was examined under an optical stereomicroscope and SEM to assess differences between sexes. The morphometric analysis of the body structures included rostrum length (base-apex and antennal insertion-apex), elytral length and width, pronotum length and width, first tarsus length, first tibia length, funiculus length and total body length. Rostrum length, elytra length and width and total body length were significantly larger in females than in males. A tergal notch in the 8th abdominal tergite was present in males and absent in females. The body structure; of rostrum length, elytra length and width and total body length overlapped between sexes in some specimens. The abdominal tergal notch was found to be the most useful body structure to distinguish males from females in A. santacruzi.

Implications of elevated carbon dioxide on the susceptibility of the globally invasive weed, Parthenium hysterophorus, to glyphosate herbicide.

BACKGROUND: The noxious annual herb, Parthenium hysterophorus L. (Asteraceae), is an invasive weed of global significance, threatening food security, biodiversity and human health. In South Africa, chemical control is frequently used to manage P. hysterophorus, however, concern surrounds increasing atmospheric CO2 levels, which may reduce the efficacy of glyphosate against the weed. Therefore, this study aimed to determine the susceptibility of P. hysterophorus to glyphosate (1L/ha: recommended) after being grown for five generations in Convirons under ambient (400 ppm) and elevated (600 and 800 ppm) CO2 . RESULTS: Glyphosate efficacy decreased with increasing CO2 , with mortalities of 100, 83 and 75% recorded at 400, 600 and 800 ppm, respectively. Parthenium hysterophorus experienced enhanced growth and reproduction under elevated CO2, however, glyphosate application was highly damaging, reducing the growth and flowering of plants across all CO2 treatments. Physiologically, glyphosate-treated plants, in all CO2 treatments, suffered severe declines of >90% in chlorophyll content, maximum quantum efficiency (F v /Fm ), photon absorption (ABS/RC), electron transport (ET 0 /RC) and performance index (PI ABS ), albeit at slower rates for plants grown under elevated CO2 . Low levels of recovery from glyphosate were documented only for plants grown under elevated CO2 and was attributed to their increased biomass. CONCLUSION: These results suggest that increasing CO2 levels may hinder chemical control efforts used against P. hysterophorus in the future, advocating for further investigation using multigenerational CO2 studies and the maintenance of effective spraying programs at present. © 2020 Society of Chemical Industry.

Tamarix efficiency in salt excretion and physiological tolerance to salt-induced stress in South Africa.

This study, investigated the salt excretion efficiency and the level of the physiological response to salt-induced stresses between the native and exotic Tamarix species as well as their hybrids (Tamarix chinensis × Tamarix ramosissima and Tamarix chinensis × Tamarix usneoides). Ten potted plants from each of the five taxa were exposed to salt at a concentration of 3% (w/w) (180 mM) for 3 weeks. Measurements of electro-conductivity (EC), physiological parameters such as stomatal conductance, chlorophyll fluorescence, and water pressure and plant growth were taken from salt-treated and control plants. The EC in the exotic T. chinensis significantly increased by >30% compared with all other Tamarix taxa, suggesting that it is the most effective taxon for phytoremediation. Although there was no significant difference in plant growth between T. chinensis and T. usneoides, they both showed a significantly greater plant growth than the other taxa. However, the plant physiological parameters indicated that T. usneoides was less stressed by the salt exposure than the T. chinensis and the others. Thus, considering the T. usneoides greater tolerance to salt-induced and/water stresses and the strict environmental regulations of planting exotic Tamarix, the native Tamarix remains the preferred plant of choice for phytoremediation in South Africa.

Straight-line orientation in the woodland-living beetle Sisyphus fasciculatus.

To transport their balls of dung along a constant bearing, diurnal savannah-living dung beetles rely primarily on the sun for compass information. However, in more cluttered environments, such as woodlands, this solitary compass cue is frequently hidden from view by surrounding vegetation. In these types of habitats, insects can, instead, rely on surrounding landmarks, the canopy pattern, or wide-field celestial cues, such as polarised skylight, for directional information. Here, we investigate the compass orientation strategy behind straight-line orientation in the diurnal woodland-living beetle Sisyphus fasciculatus. We found that, when manipulating the direction of polarised skylight, Si. fasciculatus responded to this change with a similar change in bearing. However, when the apparent position of the sun was moved, the woodland-living beetle did not change its direction of travel. In contrast, the savannah-living beetle Scarabaeus lamarcki responded to the manipulation of the solar position with a corresponding change in bearing. These results suggest that the dominant compass cue used for straight-line orientation in dung beetles may be determined by the celestial cue that is most prominent in their preferred habitat.

The effect of step size on straight-line orientation.

Moving along a straight path is a surprisingly difficult task. This is because, with each ensuing step, noise is generated in the motor and sensory systems, causing the animal to deviate from its intended route. When relying solely on internal sensory information to correct for this noise, the directional error generated with each stride accumulates, ultimately leading to a curved path. In contrast, external compass cues effectively allow the animal to correct for errors in its bearing. Here, we studied straight-line orientation in two different sized dung beetles. This allowed us to characterize and model the size of the directional error generated with each step, in the absence of external visual compass cues (motor error) as well as in the presence of these cues (compass and motor errors). In addition, we model how dung beetles balance the influence of internal and external orientation cues as they orient along straight paths under the open sky. We conclude that the directional error that unavoidably accumulates as the beetle travels is inversely proportional to the step size of the insect, and that both beetle species weigh the two sources of directional information in a similar fashion.