The rapid spread of herbicide resistance in weeds, driven by gene flow through multiple pathways, poses an increasing challenge for agricultural systems. This review summarizes the extent and distance of pollen- and seed-mediated gene flow (PMGF/SMGF) in selected self-pollinated weeds and the environmental factors that influence PMGF. A comprehensive literature review was focused on assessing PMGF patterns, dispersal mechanisms, and influencing factors across self-pollinated weed species. Statistical analyses were conducted to evaluate correlations between PMGF and environmental variables, including temperature, precipitation, humidity, and elevation. Self-pollinated weeds in the Asteraceae family show the highest PMGF (av. 10.63%), with common groundsel showing 24% at 0.5 meters from the pollen source. Solanaceae and Chenopodiaceae are families with the second and third highest PMGF (av. 10.00% and 1.58%, respectively). Within Solanaceae, eastern black nightshade exhibited the maximum PMGF (17%), whereas in Chenopodiaceae, magenta spreen showed the highest gene flow, reaching 3% at 15 meters from the pollen source. In contrast, the lowest mean PMGF was observed in Poaceae and Brassicaceae (av. 1.87% and 0.33%, respectively). Furthermore, among environmental variables, only temperature showed a significant correlation (P < 0.05). Due to the limited number of studies, this correlation should be viewed cautiously, as it likely reflects general patterns rather than a causal link to PMGF. Bold-seeded grasses such as oat may disperse seeds at low frequency (14-18%), however, light-seeded species such as horseweed can disperse as high as 99% of their seeds. Understanding gene flow in self-pollinated weeds with high fecundity is vital to limiting herbicide resistance spread in such species.

Resource partitioning remains a central ecological issue for understanding the structures of animal communities. As members of the generalist New World Thraupidae bird family, tanagers can help reinforce knowledge of animal communities. Thus, to study bird trophic ecology and relationships, I followed six tanager species (Schistochlamys ruficapillus, Tangara desmaresti, Stilpnia cayana, Stephanophorus diadematus, Thraupis sayaca and Dacnis cayana) from the Ibitipoca State Park, south-eastern Brazil, for 12 months. I expected 1) more similarities among frugivorous behaviours when compared to the similarities in the frugivorous diet, 2) no differences between the predation behaviours on arthropods against the use of substrates to capture them, 3) no specialisation in the bird–fruit seed dispersal network and 4) similarities of frugivorous diet and substrate use equal those expected by chance due to the absence of specialisation in frugivory and limited use of substrates. Several ecological and behavioural factors seemed to determine the ways in which Ibitipoca tanagers consumed fruits to reduce similarities in diets and reveal network specialisation. On the other hand, the analysis of predation behaviour indicated that tanager species used similar tactics to reach arthropods randomly on substrates. In general, frugivory seems to be more relevant in trophic partitioning than foraging on arthropods in Ibitipoca tanagers.

Boneseed [Chrysanthemoides monilifera subsp. monilifera (L.) Norl.; syn. Osteospermum moniliferum subsp. moniliferum L.] is a perennial shrub native to the southwestern and southern coasts of South Africa. It was introduced to Australia in about 1852 and now represents a significant threat to natural ecosystems. Despite C. monilifera subsp. monilifera being listed as a Weed of National Significance, momentum on improving its management has dissipated at a national level, beginning in 2008 (when a national research initiative finished) and increasingly after 2013 (when funding for national coordination ceased). A recent synthesis of past management for C. monilifera subsp. monilifera and recommendations for guiding future priorities has rekindled interest in Western Australia. To complement this synthesis and to identify improvements for program efficiency and effectiveness, we reviewed research and management findings on this weed with a focus on the past two decades. We collated information across the ecology and biology of C. monilifera subsp. monilifera, and the near relative, bitou bush [Chrysanthemoides monilifera subsp. rotunda (DC.) J.C. Manning & Goldblatt; syn. Osteospermum moniliferum subsp. rotundatum (DC.)], as well as useful insight from C. monilifera subsp. monilifera management programs applied elsewhere. As part of this review, we assessed the classical biological control work that has been done on C. monilifera subsp. monilifera, focusing on likely explanations for why, despite nine agents and a naturalized fungus, biological control is not an effective management tool. Our synthesis suggests that for the limited populations with low-abundance plants in Western Australia, eradication from the state remains a realistic target. This objective, however, needs to build on the collated baseline of past management efforts and deploy a carefully planned management program over the coming two decades. Systematic surveillance using the latest techniques, combined with manual or herbicide removal and controlled burns where possible, remain the most suitable methods to deploy. The long-lived soil seedbank requires detailed monitoring following initial plant removals and long-term funding to ensure the sustained effort required to deliver the goal of eradication of C. monilifera subsp. monilifera in Western Australia.

Endozoochory, the dispersal of seeds through the animal gut passage, plays a significant role in vegetation dynamics. The success of endozoochorous seed dispersal depends on each stage of the process: ingestion by animals, gut passage, and post-dispersal events after defecation. After the deposition of seeds through feces, the effects of feces on the initial stages of seedling establishment, including seed germination and seedling growth, can significantly impact overall survival. The pattern of fecal effects on plant species depends on the animal species. In this study, we investigated the effects of feces presence on seed germination and early seedling growth using feces of the Korean water deer (Hydropotes inermis argyropus). We conducted a germination experiment on 12 plant species belonging to 10 plant families, which are known to germinate in the feces of Korean water deer. The study compared the seed germination rate and seedling length after germination between seeds sown with and without feces of the Korean water deer. In general, we found that the presence of deer feces per se had no significant effects on seed germination and early growth stages. However, additional research on post-dispersal events such as long-term growth, fecal type, and germination conditions is needed to fully understand the costs and benefits of endozoochory.

The persistence of seed-dispersing animals in degraded habitats could be critical for ensuring the long-term conservation value and restoration of forests. This is particularly important in Southeast Asia, where > 70% of the remaining forest areas are within 1 km of a forest edge, and many are degraded (e.g. logged). We synthesized information on the habitat associations of the binturong Arctictis binturong, a large, semi-arboreal, frugivorous civet and one of the most important seed dispersers in the region, especially for figs (Ficus spp). We adopted a multiscale approach by employing ensemble species distribution modelling from presence-only records, assessing landscape-scale variation in detection rates in published camera-trap studies and using hierarchical occupancy modelling to assess local (i.e. within-landscape) patterns observed from 20 new camera-trap surveys. Contrary to prior reports that binturongs are strongly associated with intact forests, the species was equally present in degraded forests and near forest edges where sufficient forest cover was maintained (> 40% forest cover within a 20-km radius). The species also tolerates moderate incursions of oil palm plantations (< 20% of the area within a 20-km radius covered by oil palm plantations). The relative resilience of binturongs to habitat degradation could be in part because of behavioural adaptations towards increased nocturnal activity. These results support the notion that key seed dispersers can persist and maintain their ecological function in degraded forests.

Monoon liukiuense (Annonaceae) is an endangered tree species distributed in Iriomote Island and Hateruma Island in the Ryukyu (Nansei) Islands, Japan, and in Orchid Island (Lanyu) in Taiwan. While its habitat is confined to small areas surrounded by human-altered landscapes, the matured trees bear abundant fruits, and many offspring grow under the mother trees. M. liukiuense is hypothesised to have lost effective seed dispersers. To test this hypothesis, fate of its seeds and the behaviour of frugivores were observed using time-lapse photography during three fruiting seasons from June 2015 to August 2016 in Iriomote Island, Japan. Although several animal taxa were observed to consume the fruit pulp, only two volant animals, namely the Yaeyama flying fox and large-billed crow, were proposed as seed dispersal agents for M. liukiuense. The present study shows that an average of 82% of the fruits in the canopies fell directly beneath the fruiting trees and an average of 90% of the seeds on the forest floor remained in their original positions. These results suggest that M. liukiuense has lost most of its seed dispersal agents and the chance to expand its distribution.

Frugivorous mammals play an important role in maintaining biodiversity and are considered one of the main dispersers of large seeds. In this study, we describe the structure of the interaction network between non-flying mammals and seven plant species with large fruits in a megadiverse savanna-forest mosaic in the Brazilian Cerrado. We also evaluated the individual contribution of each species to the organization of the interaction network and tested whether body mass determined the mammals’ role in the network. To record frugivory events of mammals with arboreal and terrestrial habits, camera traps were installed at ground and canopy levels. We identified 18 mammal species interacting with seven plant species in 515 frugivory events. Our observations highlight an interaction network with a modular and non-nested topology and the important role of large mammals in the network structure, which reflects the importance of the group in potential seed dispersal. The extinction of large frugivorous mammals can cause several damages to ecosystem services in the Brazilian Cerrado through changes in network structure, especially threatening the survival of plant species with large fruits.

Branched broomrape, an obligate root parasitic weed, has recently re-emerged in tomato fields in several California counties. California produces more tomato than any other state, and the outbreak of this noxious weed could potentially wreak havoc on the industry’s economy. Preventive measures must be taken to stop or reduce the spread of branched broomrape seeds to other areas. Branched broomrape can produce thousands of tiny seeds, which can easily spread with farm machinery over short and long distances. To prevent branched broomrape seed dispersal, sanitation and disinfection of farm equipment are necessary before entering a new farm. We tested the effectiveness of various ammonium compounds, including didecyl dimethyl ammonium chloride (DDAC), alkyl dimethyl benzyl ammonium chloride (ADBC), didecyl dimethyl ammonium bromide (DDAB), ammonium bromide (AB), and ammonium chloride (AC) on prevention of branched broomrape seed germination. Dose-response analysis showed that three chemical products, ADBC, DDAB, and DDAC, could completely inhibit branched broomrape seeds (potentially making them nonviable) at 1%, 1%, and 10% wt/vol concentrations, respectively. These three compounds were further tested in an exposure duration experiment that additionally included Egyptian broomrape. Only 10 min of exposure to these compounds was needed to prevent germination of both branched and Egyptian broomrape seeds at 1% (ADBC, DDAB) and 10% wt/vol (DDAC). Lower concentrations can provide similar inhibition effects when combined with longer exposure times. Egyptian broomrape seeds were more sensitive than branched broomrape seeds. Findings suggest that quaternary ammonium compounds could be used as potential sanitation agents to disinfect agriculture machinery from branched and Egyptian broomrape seeds.

Species-rich ecosystems as tropical forests are extremely vulnerable to anthropogenic destruction. Most tropical plant species rely on animals to disperse their seeds. However, patterns of mutualistic interactions have rarely been explored, and seed dispersal networks are still poorly studied in Africa. Here, we examine how forest edges’ (FE) seed dispersal networks differ from the mature forest (MF) at a West African sub-humid tropical forest within the National Park of Cantanhez (Guinea-Bissau). Additionally, we explore species’ roles within the network. MF had higher fruit availability, more frugivore visitors, and plant–frugivore interactions. Network structure was quite similar between habitats, showing signs of redundancy, and some robustness to species’ extinction. FE was more nested, modular, and specialized, whereas MF had higher connectance, interaction evenness, and robustness to extinction. Most species were generalists, but large-bodied frugivores prevailed at MF. FE showed a higher vulnerability, mostly to the loss of trees. Trees are key, keeping the structure of both networks. Large-bodied frugivores and fruiting-tree species that work as network connectors should thus be the focus of active conservation management in these forests. Only viable populations of these species will ensure a good performance of the seed dispersal network, promoting the natural regeneration of the ecosystem.

Large herbivores can disperse seeds over long distances through endozoochory. The Korean water deer (Hydropotes inermis argyropus), an internationally vulnerable species but locally considered a vermin, is a potential endozoochorous seed dispersal vector. In this study, feeding experiments were conducted to test the efficiency of seed dispersal through gut ingestion by the Korean water deer, its temporal pattern and the effect of gut passage on seed recovery and germination rate. Eight plant species, including species that formerly germinated from its faeces, were used to feed three Korean water deer. Once the deer had consumed all the provided seeds, their faeces were collected after 24, 48, 72 and 96 h. The collected faeces were air-dried, and the number of seeds retrieved from the faeces was counted every 24 h (0–24, 24–48, 48–72 and 72–96 h). Among the eight plant species, six species were retrieved with intact seeds. Panicum bisulcatum had the highest recovery rate of 33.7%, followed by Amaranthus mangostanus (24.5%) and Chenopodium album (14.4%). Most of the seeds were recovered within the 24–48 h time interval. Germination tests were conducted on the ingested and uningested seeds for the four species which had a sufficient recovery rate. The effects of gut passage on seed germination differed according to plant species. The germination rate substantially decreased after gut passage. The results suggest that the Korean water deer can disperse seeds, potentially over long distances albeit at a high cost of low seed recovery and germination rate.

Myrmecochory, the dispersal of seeds with lipid-rich appendages by ants, is a significant ant–plant interaction. Less well understood is the potential for ant dispersal of non-myrmecochorous seeds. Here we investigate ant–diaspore interactions in a West African montane habitat. We combine observation with depot experiments to determine ant species that move diaspores and distance moved across a forest-edge-grassland gradient. We recorded seed cleaning by ants using a bird/mammal dispersed Paullinia pinnata to determine whether seed cleaning improved plant fitness. We found that two out of a total of 17 ant species (Pheidole sp. 1 and Myrmicaria opaciventris) interacted with 10 species of non-myrmecochorous diaspores across nine plant families. Diaspores were from large canopy trees, understorey trees and vines. Both ant species interacted with small (≤0.24 g) and large (≥0.24 g) diaspores. Ants individually moved small diaspores up to 1.2 m and worked together to clean larger ones. Our experiments with P. pinnata showed that ants removed the pulp of 70% of fruit over 5 days. Cleaned seeds germinated significantly faster and produced seedlings with significantly longer shoot length and higher fresh weight than seedlings from intact seeds. Together our results suggest that ant dispersal may be less significant than seed cleaning in Afromontane forests. However, given the decline in vertebrate frugivores across Africa, a small dispersal advantage may become increasingly important to plant fitness.

In central Europe, barnyardgrass [Echinochloa crus-galli (L.) P. Beauv.], has commonly been found in humid lowland areas. As a result of the introduction of new crops and farming practices, in the northwest Carpathians, E. crus-galli has spread from lowland (<200 m altitude) to highland (>400 m altitude) areas. We collected seed samples from local populations lying at a distance of approximately 5 km from each other and lined up along transects following the flows of two rivers. The rivers first flow through the valleys separated by mountain ridges and eventually flow into a common lowland. After ripening, the seeds of all populations were germinated at 25 C under long-day conditions. Only the seeds of some lowland populations germinated up to 75%. The frequency of germinated seeds decreased as the altitude where the population was collected increased, and above 200 m above sea level, germination was mostly zero. We then studied the phenological and morphological differentiation of plants from the original (lowland) and recently occupied (highland) areas. Seeds of the lowest and the highest localities lying on the transect of each river were sown in a common garden experiment. In plants from the highland localities, heading and seed dispersal were earlier, while tiller height and tiller mass were lower than in plants from the lowland localities. Seed mass produced per tiller in the lowland and highland plants was similar, and as a result, highland plants allocated a larger proportion of body mass to seed production than did lowland plants. Echinochloa crus-galli populations from highland localities thus produce their progeny earlier and at a lower energy cost than populations from lowland localities. The plasticity of phenological characters likely facilitated adaptation during E. crus-galli spread from lowlands to highlands. Similar adaptations in plant phenology may contribute to the spread of E. crus-galli in other geographic areas.

Cultivated rice (Oryza sativa L.) was domesticated from the Asian wild species O. rufipogon. Compared with cultivated rice, wild rice has spikelets/seeds with long barbed awns. In order to evaluate the role of awns in wild rice, four seed groups with different awn lengths (full, half, quarter and no awns) were prepared, and the following seed dispersal movements were investigated under simulated natural conditions as observed in the tropical Asian habitat: (1) seed detachment from the parent plant; (2) falling angle of mature seed; (3) ability to slip into small spaces; (4) horizontal movement on the ground and (5) horizontal movement in water. As a result, awns were found to enhance the detachment of matured seeds from the panicles in the initial seed dispersal step. They regulated vertical orientation in the air, and the vertical form may enable seeds to squeeze to the ground. The awned seeds also showed advantages in horizontal movements on the ground and in the water. In most of the experiments, seeds with full awns showed the best performance for seed dispersal, suggesting that wild rice keeps long awns to survive under natural conditions. Since seed awning is dominantly controlled by wild functional alleles at several loci, wild rice may be able to prevent a drastic reduction of awn length.

The great fruit-eating bat (Artibeus lituratus) is a large-sized species that forages primarily on fruits. This species is widespread throughout the Neotropics, where it is common in natural areas and also occupies forest patches and cities. In this study, we review the composition of Artibeus lituratus diet in Brazil as well as the size of fruits and seeds, plant geographic origin, and sampling methods used in natural versus urban habitats. We show that Artibeus lituratus is able to consume a higher proportion of exotic fruits with large seeds in urban environments than in natural areas. Fruit diameter was not statistically different between environments, but both fruit and seed diameters are smaller when detected by fecal sampling than by other methods. This difference is likely due to the fact that in natural habitats studies are predominantly based on fecal samples, which hinders the detection of large unswallowed seeds. Consequently, we recommend the use of complementary sampling methods (not only the widely used technique of fecal sorting) in order to produce more accurate descriptions of frugivorous bats’ diets. We suggest that the ability to exploit fruits of exotic plant species including the ones with large seeds may be a key trait for the persistence of A. lituratus in urban habitats.

In 2016 and 2017, 98 separate commercially available bird feed mixes were examined for the presence of weed seed. All weed seed contaminants were counted and identified by species. Amaranthus species were present in 94 of the 98 bags of bird feed. Amaranthus species present in bird feed mixes included waterhemp [Amaranthus tuberculatus (Moq.) Sauer], redroot pigweed (Amaranthus retroflexus L.), Palmer amaranth (Amaranthus palmeri S. Watson), smooth pigweed (Amaranthus hybridus L.), and tumble pigweed (Amaranthus albus L.). Amaranthus palmeri was present in 27 of the 98 mixes. Seed of common ragweed (Ambrosia artemisiifolia L.), kochia [Bassia scoparia (L.) A.J. Scott], grain sorghum [Sorghum bicolor (L.) Moench], wild buckwheat (Fallopia convolvulus L., syn: Polygonum convolvulus), common lambsquarters (Chenopodium album L.), large crabgrass [Digitaria sanguinalis (L.) Scop.], and Setaria species were also present in bird feed mixes. A greenhouse assay to determine Amaranthus species seed germinability and resistance to glyphosate revealed that approximately 19% of Amaranthus seed in bird feed mixes are readily germinable, and five mixes contained A. tuberculatus and A. palmeri seed that were resistant to glyphosate. Results from linear regression and t-test analysis indicate that when proso millet (Panicum miliaceum L.), grain sorghum, and corn (Zea mays L.) were present in feed mixes, Amaranthus seed contamination was increased. The presence of proso millet and grain sorghum also increased contamination of grass weed species, while sunflower (Helianthus annuus L.) increased A. artemisiifolia contamination and safflower (Carthamus tinctorius L.) increased contamination of Bassia scoparia.

Seed dispersal is an important ecological process that structures plant communities and influences ecosystem functioning. Loss of animal dispersers therefore poses a serious threat to forest ecosystems, particularly in the tropics where zoochory predominates. A prominent example is the near-total extinction of seed dispersers on the tropical island of Guam following the accidental introduction of the invasive brown tree snake (Boiga irregularis), negatively impacting seedling recruitment and forest regeneration. We investigated frugivory by a remnant population of Såli (Micronesian starling – Aplonis opaca) on Guam and two other island populations (Rota, Saipan) to evaluate their ecological role as a seed disperser in the Mariana archipelago. Using a combination of behavioural observations, nest contents and fecal samples, we documented frugivory of 37 plant species. Native plants comprised the majority (66%) of all species and 90% of all seeds identified in fecal and nest contents. Diet was highly similar across age classes and sampling years. In addition, plant species consumed by Såli comprised 88% of bird-dispersed adult trees and 54% of all adult trees in long-term forest monitoring plots, demonstrating the Såli’s broad diet and potential for restoring native forests. Overall, we provide the most comprehensive assessment to date of frugivory by the Såli and confirm its importance as a seed disperser on Guam and throughout the Marianas.

Large frugivores provide critical seed dispersal services for many plant species and their extirpation from forested ecosystems can cause compositional shifts in regenerating plant cohorts. Yet, we still poorly understand whether large seed-dispersers have complementary or redundant roles for forest regeneration. Here, to assess the functional complementarity of large-bodied frugivores in forest regeneration, we quantified the effects of varying abundance of hornbills, primates and the forest elephant on the density, species richness and the mean weighted seed length of animal-dispersed tree species among seedlings in five sites in a forest–savanna mosaic in D. R. Congo, while accounting for percentage forest cover and the local presence of fruiting trees. We found that the abundance of primates was positively associated with species richness of seedlings, while percentage forest cover was negatively associated (R2 = 0.19). The abundance of hornbills, the presence of elephants and percentage forest cover were positively associated with mean seed length of the regenerating cohort (R2 = 0.13). Spatially explicit analysis indicated that some additional processes have an important influence on these response indices. Primates would seem to have a preponderant role for maintaining relatively high species richness, while hornbills and elephant would seem to be predominantly responsible for the recruitment of large-seeded trees. Our results could indicate that these taxa of frugivores play complementary functional roles for forest regeneration. This suggests that the extirpation of one or more of these dispersers would likely not be functionally compensated for by the remaining taxa, hence possibly cascading into compositional shifts.

Many plant traits might explain the different ecological and network roles of fruit-eating birds. We assessed the relationship of plant productivity, fruit traits (colour, seed size and nutritional quality) and dietary specialization, with the network roles of fruit-eating birds (number of partners, centrality and selectivity) in the Atlantic Forest, Brazil. We classified bird species according to their dietary specialization into three categories: obligate, partial and opportunistic fruit-eating birds. To test if network roles changed according to dietary specialization, fruit productivity and traits, we used a generalized linear model analysis. The selected 14 species of plant interacted with 52 bird species, which consumed 2199 fruits. The most central and generalist fruit-eating bird, Turdus albicolis, interacted with plants that produced more fruits, such as Miconia cinerascens, and had, on average, larger seeds, such as Myrcia splendens. The most selective birds interacted with fruits with a higher concentration of lipids and less intense colour, and plants that produced fewer fruits. Obligate fruit-eating birds, such as Patagioenas plumbea, were more selective than partial and opportunistic birds. Different plant traits are therefore related to the different network roles of fruit-eating birds in the Atlantic Forest, which are also dependent on bird dietary specialization.

Large animals tend to disperse seeds over long distances via ingestion and defecation due to their large home range and capacity to move among different habitats for feeding. The aim of this study was to investigate the potential of endozoochorous seed dispersal by five herbivores: Ovis vignei, Capra aegagrus, Gazella subgutturosa, Cervus elaphus, Capreolus capreolus and two omnivores: Sus Scrofa and Ursus arctos in the Golestan National Park, northeast of Iran, by a greenhouse germination experiment. A total of 3107 seedlings belonging to 154 different plant taxa were germinated from 655 dung samples collected in three different habitats. Plant families that most frequently germinated were Poaceae and Brassicaceae. Urtica dioica was the most abundant germinating seed, accounting for 20% of all the seedlings recorded in our dung samples, whereas the most frequently observed species was Portulaca oleracea, which occurred in 24% of our samples. We showed that 54% of the seeds germinating were dispersed by only one of the mammals studied. Herbs and graminoids were the most frequently dispersed growth forms by the herbivores and the wild boar, whereas brown bears mostly dispersed shrubs. The seedling composition in the dung samples was strongly correlated with the local flora especially for non-selective feeders, like red deer. The differences observed in the number of plants and frequency of different growth forms dispersed among the studied mammals reflect their body size, digestive physiology, and dietary and habitat preferences. Our findings highlight the different and complementary roles of large herbivores and omnivores as long-distance seed dispersal vectors.

Harvest weed seed control (HWSC) techniques have been implemented in Australian cropping systems to target and reduce the number of weed seeds entering the seedbank and thereby reduce the number of problematic weeds emerging in subsequent years to infest subsequent crops. However, the influence of HWSC on ameliorating herbicide-resistance (HR) evolution has not been investigated. This research used integrated spatial modeling to examine how the frequency and efficacy of HWSC affected the evolution of resistance to initially effective herbicides. Herbicides were, in all cases, better protected from future resistance evolution when their use was combined with annual HWSC. Outbreaks of multiple HR were very unlikely to occur and were nearly always eliminated by adding annual, efficient HWSC. The efficacy of the HWSC was important, with greater reductions in the number of resistance genes achieved with higher-efficacy HWSC. Annual HWSC was necessary to protect sequences of lower-efficacy herbicides, but HWSC could still protect herbicides if it was used less often than once per year, when the HWSC and the herbicides were highly effective. Our results highlight the potential benefits of combining HWSC with effective herbicides for controlling weed populations and reducing the future evolution of HR.