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More than 100 RNA chemical modifications to cellular RNA have been identified. N 6-methyladenosine (m6A) is the most prevalent modification of mRNA. RNA modifications have recently attracted significant attention due to their critical role in regulating mRNA processing and metabolism. tRNA and rRNA rank among the most heavily modified RNAs, and their modifications are essential for maintaining their structure and function. With our advanced understanding of RNA modifications, increasing evidence suggests RNA modifications are important in regulating various aspects of insect life. In this review, we will summarize recent studies investigating the impact of RNA modifications in insects, particularly highlighting the role of m6A in insect development, reproduction, and adaptation to the environment.

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Chagas disease affects around 8 million people globally, with Latin America bearing approximately 10,000 deaths each year. Combatting the disease relies heavily on vector control methods, necessitating the identification of new targets. Within insect genomes, genes harboring small open reading frames (smORFs - < 100 amino acids) present numerous potential candidates. In our investigation, we elucidate the pivotal role of the archetypal smORF-containing gene, mille-pattes/polished-rice/tarsalless (mlpt/pri/tal), in the post-embryonic development of the kissing bug Rhodnius prolixus. Injection of double-stranded RNA targeting mlpt (dsmlpt) during nymphal stages yields a spectrum of phenotypes hindering post-embryonic growth. Notably, fourth or fifth stage nymphs subjected to dsmlpt do not undergo molting. These dsmlpt nymphs display heightened mRNA levels of JHAMT-like and EPOX-like, enzymes putatively involved in the juvenile hormone (JH) pathway, alongside increased expression of the transcription factor Kr-h1, indicating changes in the hormonal control. Histological examination reveals structural alterations in the hindgut and external cuticle of dsmlpt nymphs compared to control (dsGFP) counterparts. Furthermore, significant changes in the vector's digestive physiology were observed, with elevated hemozoin and glucose levels in the posterior midgut of dsmlpt nymphs. Importantly, dsmlpt nymphs exhibit impaired metacyclogenesis of Trypanosoma cruzi, the causative agent of Chagas disease, underscoring the crucial role of proper gut organization in parasite differentiation. Thus, our findings constitute the first evidence of a smORF-containing gene's regulatory influence on vector physiology, parasitic cycle, and disease transmission.

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Host-microbe interactions underlie the development and fitness of many macroorganisms, including bees. Whereas many social bees benefit from vertically transmitted gut bacteria, current data suggests that solitary bees, which comprise the vast majority of species diversity within bees, lack a highly specialized gut microbiome. Here, we examine the composition and abundance of bacteria and fungi throughout the complete life cycle of the ground-nesting solitary bee Anthophora bomboides standfordiana. In contrast to expectations, immature bee stages maintain a distinct core microbiome consisting of Actinobacterial genera (Streptomyces, Nocardiodes) and the fungus Moniliella spathulata. Dormant (diapausing) larval bees hosted the most abundant and distinctive bacteria and fungi, attaining 33 and 52 times their initial copy number, respectively. We tested two adaptive hypotheses regarding microbial functions for diapausing bees. First, using isolated bacteria and fungi, we found that Streptomyces from brood cells inhibited the growth of multiple pathogenic filamentous fungi, suggesting a role in pathogen protection during overwintering, when bees face high pathogen pressure. Second, sugar alcohol composition changed in tandem with major changes in fungal abundance, suggesting links with bee cold tolerance or overwintering biology. We find that A. bomboides hosts a conserved core microbiome that may provide key fitness advantages through larval development and diapause, which raises the question of how this microbiome is maintained and faithfully transmitted between generations. Our results suggest that focus on microbiomes of mature or active insect developmental stages may overlook stage-specific symbionts and microbial fitness contributions during host dormancy.

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Blow flies (Diptera: Calliphoridae) are generally early colonisers of fresh cadavers, enabling the estimation of a minimum post-mortem interval (minPMI) based on an accurate aging of the oldest immature stages associated with a cadaver. In blow flies, the pupal stage and the subsequent development of the adult take place inside a protective case, the puparium, formed from the hardened and darkened cuticle of the third instar larva. Because the puparium is an opaque structure that shows virtually no external changes, qualitative analyses of the internal tissues can be very informative for determining reliable age-specific morphological markers. Those analyses can be performed using either non-invasive but expensive and not widely accessible techniques, or traditional histological methods, which are invasive as they require the serial sectioning of the sample. Histological methods are often readily available for forensic researchers and practitioners; however, the histological study of blow fly intra-puparial stages has traditionally been hampered by the poor paraffin infiltration of tissues due to the abundance of fat bodies, resulting in usually fragmented sections and the subsequent loss of relevant information. We present here an effective method for the preparation of histological sections of blow fly intra-puparial stages, maximising the paraffin infiltration while enabling the production of clean and entire sections that allow for the use of reliable age-specific morphological markers, thus improving the accuracy of minPMI estimations when access to more costly techniques is not feasible.

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In holometabolous insects, extensive reorganisation of tissues and cells occurs at the pupal stage. The remodelling of the external exoskeleton and internal organs that intervenes during metamorphosis has been traditionally studied in many insect species based on histological or ultrastructural methods. This study demonstrates the use of synchrotron X-ray phase-contrast micro-computed tomography as a powerful, non-destructive tool for in situ morphological observation of anatomical structures at the pupal stage in two Tenebrionid beetles, i.e. Tribolium castaneum and Tenebrio molitor, known as important pests, as well as emerging and promising models in experimental biology. Virtual sections and three-dimensional reconstructions were performed on both males and females at early, intermediate, and late pupal stage. The dataset allowed us to observe the remodelling of the gut and nervous system as well as the shaping of the female and male reproductive system at different pupal ages in both mealworm and red flour beetles. Moreover, we observed that the timing and duration pattern of organ development varied between the species analysed, likely related to the species-specific adaptations of the pre-imaginal stages to environmental conditions, which ultimately affect their life cycle. This research provides new knowledge on the morphological modifications that occur during the pupal stage of holometabolous insects and provides a baseline set of information on beetle metamorphosis that may support future research in forensics, physiology, and ecology as well as an image atlas for educational purposes.

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Hormones control developmental and physiological processes, often by regulating the expression of multiple genes simultaneously or sequentially. Crosstalk between hormones and epigenetics is pivotal to dynamically coordinate this process. Hormonal signals can guide the addition and removal of epigenetic marks, steering gene expression. Conversely, DNA methylation, histone modifications and non-coding RNAs can modulate regional chromatin structure and accessibility and regulate the expression of numerous (hormone-related) genes. Here, we provide a review of the interplay between the classical insect hormones, ecdysteroids and juvenile hormones, and epigenetics. We summarize the mode-of-action and roles of these hormones in post-embryonic development, and provide a general overview of epigenetic mechanisms. We then highlight recent advances on the interactions between these hormonal pathways and epigenetics, and their involvement in development. Furthermore, we give an overview of several 'omics techniques employed in the field. Finally, we discuss which questions remain unanswered and possible avenues for future research.

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Estrogen-related receptor (ERR) is a key regulator of insect growth, development, and metabolic processes in insects; however, the molecular mechanisms underlying its effects are not fully understood. We investigated roles of 20-hydroxyecdysone (20E) and insulin/insulin-like signaling/target of rapamycin (IIS/TOR) signaling pathways in the effects of PvERR on larval development, metamorphosis, and adult growth in ant Polyrhachis vicina Roger. PvFOXO expression levels depended on caste and developmental stage. PvERR RNAi significantly reduced the expression levels of IIS/TOR signaling pathway genes and 20E signaling pathway genes in fourth-instar larvae, pupae, females, and workers and significantly increased the expression levels of IIS/TOR signaling pathway genes PvFOXO and PvAkt in males. PvFOXO RNAi resulted in developmental defects and increased mortality. After PvFOXO RNAi, the expression of PvERR, 20E signaling pathway genes, and IIS/TOR signaling pathway genes decreased significantly in pupae, females, and workers and increased significantly in fourth-instar larvae. Exogenous 20E attenuated expression changes induced by PvFOXO RNAi in a sex- and stage-specific manner. These results indicate that ERR interacts with 20E and IIS/TOR signaling pathways to regulate caste determination, metamorphosis, and male fertility in P. vicina and that correlations between PvERR and PvFOXO are caste- and stage-specific.

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Terpenes are a major class of secondary metabolites present in all plants, and long hypothesized to have diversified in response to specific plant-herbivore interactions. Herbivory is a major biotic interaction that plays out across broad temporal and spatial scales that vary dramatically in temperature regimes, both due to climatic variation across geographic locations as well as the effect of seasonality. In addition, there is an emerging understanding that global climate change will continue to alter the temperature regimes of nearly every habitat on Earth over the coming centuries. Regardless of source, variation in temperature may influence herbivory, in particular via changes in the efficacy and impacts of plant defensive chemistry. This study aims to characterize temperature-driven variation in toxicological effects across several structural classes of terpenes in the model herbivore Vanessa cardui, the painted lady butterfly. We observed a general increase in monoterpene toxicity to larvae, pupa, and adults at higher temperatures, as well as an increase in development time as terpene concentration increased. Results obtained from this study yield insights into possible drivers of seasonal variation in plant terpene production as well as inform effects of rising global temperatures on plant-insect interactions. In the context of other known effects of climate change on plant-herbivore interactions like carbon fertilization and compensatory feeding, temperature-driven changes in plant chemical defense efficacy may further complicate the prediction of climate change impacts on the fundamental ecological process of herbivory.

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Ctenolepisma calvum (Ritter, 1910) (Zygentoma: Lepismatidae) is a primitive wingless insect that causes damage to paper, and it is regarded as a pest of collections in museums, archives, and libraries. This species was recently discovered in Japan for the first time and may have already spread over large areas of Japan, but, currently, no information is available on the biological characteristics of C. calvum in Japan. In this study, we observed the processes of development and reproduction of C. calvum found in Japan at room temperature. Oviposition was observed from April to November, with a peak in early June. The average egg period was 56.9 days at average temperatures above 24.0 °C, and was 72.4 days at average temperatures below 24.0 °C. The 1st, 2nd, and 3rd instars lasted 4.7 days, 13.2 days, and 26.1 days on average, respectively, at average temperatures above 22.0 °C. Average instar periods were 23-28 days in 4th-7th instars and tended to increase in later instars. Instar periods also increased when the average temperature was 22.0 °C or lower. In individual rearing, the longest-living individual lived for approximately two years, up to the 15th instar. The head width grew at an approximate ratio of 1.1 per molt. First oviposition occurred at the 10th or 11th instar. Individually observed females oviposited once or twice a year, laying 6-16 eggs at one time, but females at least two years old laid 78.2 eggs per year on average in a mass-culture cage. Through this study, only females were found, and the mature females produced their progenies parthenogenetically.

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MicroRNAs (miRNAs), a class of non-coding small RNAs, are crucial regulatory factors in plants and animals at the post-transcriptional level. These tiny molecules suppress gene expression by complementary oligonucleotide binding to sites in the target messenger. Recently, the discovery of plant-derived miRNAs with cross-kingdom abilities to regulate gene expression in insects has promoted exciting discussion, although some controversies exist regarding the modulation of insect development by plant-derived miRNAs. Here, we review current knowledge about the mechanisms of miRNA biogenesis, the roles of miRNAs in coevolution between insects and plants, the regulation of insect development by plant-derived miRNAs, the cross-kingdom transport mechanisms of plant-derived miRNAs, and cross-kingdom regulation. In addition, the controversy regarding the modulation of insect development by plant-derived miRNAs also was discussed. Our review provides new insights for understanding complex plant-insect interactions and discovering new strategies for pest management and even crop genetic improvement.

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Blow fly development rates have become a key factor in estimating the postmortem interval where blow flies are among the first decomposers to occur on a body. Because the use of blow fly development requires short time durations and high accuracy, stage transition distributions are essential for proper development modeling. However, detailed examinations of stage transitions are not available for any blow fly species. Consequently, we examined this issue in two blow fly species: Lucilia sericata and Phormia regina. Transitions for all life stages across all measured temperatures were normally distributed. Use of probit analysis allowed determination of 50% transition points and associated measures of variation (i.e., standard errors). The greatest variation was noted for the L2-L3, L3-L3m, and L3m-P stage transitions. These results invalidate the notion that largest maggots should be preferentially collected for determining current maggot population stage, and further call into question the relationship between intrinsic variation and potential geographic variation in development rates.

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Adipose tissue is a central organ for controlling systemic metabolism both in invertebrates and vertebrates. Here, we have investigated the developmental processes of the adult-type fat body (AFB) in Drosophila. We have established genetic tools that allow visualization and genetic manipulations of cells in the AFB lineage from early in metamorphosis. We identified precursor cells that give rise to the AFB and delineated dynamic cellular behaviors underlying AFB formation. These precursor cells displayed polarized cell shapes and oriented motility, with emigration from the thorax and subsequent dispersal to the abdomen and head. After the migration period, these cells adhered to each other, assembling into the AFB with a sheet-like architecture. Continuous cell proliferation occurred during and after the large-scale migration to achieve appropriate fat tissue mass. Homotypic cell fusion after the sheet formation contributed to the establishment of multinucleated cells in the AFB. We also examined candidate gene functions, and our results argue that ecdysone signaling and the transcription factor Serpent support adult fat body organogenesis.

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20-Hydroxyecdysone (20E) plays an essential role in coordinating developmental transitions in insects through responsive protein-coding genes and microRNAs (miRNAs). However, the interplay between 20E and miRNAs during insect metamorphosis is unknown. In this study, using small RNA sequencing, a comparative miRNA transcriptomic analysis in different development stages, and 20E treatment, we identified ame-bantam-3p as a key candidate miRNA involved in honeybee metamorphosis. Target prediction and in vitro dual-luciferase assays confirmed that ame-bantam-3p interacts with the coding region of the megf8 gene and promotes its expression. Meanwhile, temporal expression analysis revealed that the expression of ame-bantam-3p is higher in the larval stage than in prepupal and pupal stages, and that this expression pattern is similar to that of megf8. In vivo, we found that the mRNA level of megf8 was significantly increased after the injection of ame-bantam-3p agomir. A 20E feeding assay showed that 20E downregulated the expression of both ame-bantam-3p and its target gene megf8 on larval days five, six, and seven. Meanwhile, the injection of ame-bantam-3p agomir also reduced the 20E titer, as well as the transcript levels of essential ecdysteroid synthesis genes, including Dib, Phm, Sad, and Nvd. The transcript levels of 20E cascade genes, including EcRA, ECRB1, USP, E75, E93, and Br-c, were also significantly decreased after ame-bantam-3p agomir injection. However, ame-bantam-3p antagomir injection and dsmegf8 injection showed the opposite effect to ame-bantam-3p agomir injection. Ame-bantam-3p agomir treatment ultimately led to mortality and the failure of larval pupation by inhibiting ecdysteroid synthesis and the 20E signaling pathway. However, the expression of 20E signaling-related genes was significantly increased after megf8 knockdown, and larvae injected with dsmegf8 showed early pupation. Combined, our results indicate that ame-bantam-3p is involved in the 20E signaling pathway through positively regulating its target gene megf8 and is indispensable for larval-pupal development in the honeybee. These findings may enhance our understanding of the relationship between 20E signaling and small RNAs during honeybee development.

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Trehalose is a major circulatory sugar in the haemolymph of insects. It provides instant energy and protection against stress. Trehalose metabolism is associated with insect growth and development. The architecture and spatio-temporal expression dynamics of trehalose metabolism and transport genes are key for regulation. These genes are controlled by various transcription factors, largely linked to nutrition, insect development, and metamorphosis. Also, trehalose levels are affected by substrate affinities and modifications of enzymes involved in the pathway. A feedback mechanism involving the precursors and products can regulate trehalose metabolism. Further, the neuroendocrine system controls trehalose levels under normal and stressed conditions by producing different hormones. Hypotrehalosemic hormones work under surplus energy conditions to activate haemolymph trehalose uptake and degradation. In contrast, hypertrehalosemic hormones stimulate trehalose production in the fat body and its transport to the haemolymph. However, trehalose metabolism regulation in insects needs to be studied in detail. This review discusses aspects of trehalose synthesis, transport, and degradation dynamics in developmental transition and stress response. Unraveling the epigenetic factors, transcriptional control and chemical or genetic modulators can provide further insights into the intricate regulation of trehalose in a development- and tissue-specific manner. This molecular information about effectors and regulators of trehalose metabolism can be applied in developing diverse biotechnological applications.

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The isolated sub-Antarctic islands are of major ecological interest because of their unique species diversity and long history of limited human disturbance. However, since the presence of Europeans, these islands and their sensitive biota have been under increasing pressure due to human activity and associated biological invasions. In such delicate ecosystems, biological invasions are an exceptional threat that may be further amplified by climate change. We examined the invasion trajectory of the blowfly Calliphora vicina (Robineau-Desvoidy 1830). First introduced in the sub-Antarctic Kerguelen Islands in the 1970s, it is thought to have persisted only in sheltered microclimates for several decades. Here, we show that, in recent decades, C. vicina has been able to establish itself more widely. We combine experimental thermal developmental data with long-term ecological and meteorological monitoring to address whether warming conditions help explain its current success and dynamics in the eastern Kerguelen Islands. We found that warming temperatures and accumulated degree days could explain the species' phenological and long-term invasion dynamics, indicating that climate change has likely assisted its establishment. This study represents a unique long-term view of a polar invader and stresses the rapidly increasing susceptibility of cold regions to invasion under climate change.

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Hylurgus ligniperda (F.) and Hylastes ater (Paykull) are secondary bark beetles that have successfully spread beyond their native range, particularly into Pinus spp. plantations in the Southern Hemisphere. They feed on the phloem and cambial regions of highly stressed and recently dead Pinus spp. Here H. ligniperda and H. ater egg, larval, and pupal survival and development rates were modeled. Survival was variably influenced by temperatures depending on the life stage, but general trends were for H. ligniperda to tolerate warmer temperatures in comparison to H. ater. Nonlinear models showed 26, 29, and 34°C are the optimal temperature (maximum development rates) for the development of eggs, larvae, and pupae of H. ligniperda. In contrast, optimal temperature predictions were lower for H. ater, with estimates of 26, 22, and 23°C for the development of eggs, larvae, and pupae, respectively. H. ligniperda pre-imaginal stages were more tolerant to high temperatures, and H. ater pre-imaginal stages were more tolerant to low temperatures. Understanding the thermal requirements and limits for development for these two pests can assist in modeling emergence times, their current and potential species distribution and have potential phytosanitary applications.

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This study evaluated the effects of acute exposure of Aedes aegypti third instar (L3 ) larvae to the saline extract of Opuntia ficus-indica cladodes on the biological cycle and fertility of the emerging adults. For this, larvae were treated for 24 h with the extract at » LC50 (lethal concentration to kill 50% of larvae), ½ LC50 or LC50 ; the development and reproduction of the emerged adults were evaluated after a recovery period of 9 days. The resistance of proteins in the extract to hydrolysis by L3 digestive enzymes and histomorphological alterations in the larval midgut were also investigated. The extract contained lectin, flavonoids, cinnamic derivatives, terpenes, steroids, and reducing sugars. It showed a LC50 of 3.71% for 48 h. The data indicated mean survival times similar in control and extract treatments. It was observed development delay in extract-treated groups, with a lower number of adults than in control. However, the females that emerged laid similar number of eggs in control and treatments. Histological evaluation revealed absence of bacterial and fungal microorganisms in the food content in midguts from larvae treated with cladode extract. Electrophoresis revealed that three polypeptides in the extract resisted to hydrolysis by L3 digestive proteases for 90 min. The lectin activity was not altered even after 24-h incubation with the enzymes. In conclusion, the extract from O. ficus-indica can delay the development of Ae. aegypti larvae, which may be linked to induction of an axenic environment at larval midgut and permanence of lectin activity even after proteolysis.

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Microplastics, tiny plastic fragments from 1 µm to 5 mm, are widespread globally, even in remote environments. Due to their small sizes, they are easily ingested by organisms and contaminate the food chain. Recently, the biodegradation of some recalcitrant plastics by larva of Tenebrio molitor L. (mealworm) has been reported. However, the effects of microplastic feeding on them are limited. In our study, we selected rigid micro-polystyrene (MPS) as the model plastic to investigate the influences of particle size and larval age on plastic consumption and degradation, and the effects of microplastic feeding on the survival and development of mealworms at different larval ages. The smaller the microplastic fragment was, the more plastics the mealworms consumed, though there was a limit on particle size. Mealworms of three-month-old had the highest consumption rate. Both depolymerization and modification on the functional groups were only observed in frass excreted by three-month old mealworms. Additionally, mealworms cofed with wheat bran and MPS of this age had comparable mortality, larval growing curve and pupation distribution as the control group with wheat bran. Our results demonstrated that mealworms in this larval stage had the greatest resistance to high doses of microplastic feeding. We suggested that microplastic waste could be provided to three-month old mealworms as half replacement of bran diet to result in the greatest plastic consumption and degradation.

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Prothoracicotropic hormone (PTTH) is a central regulator of insect development that regulates the production of the steroid moulting hormones (ecdysteroids) from the prothoracic glands (PGs). Rhodnius PTTH was the first brain neurohormone discovered in any animal almost 100 years ago but has eluded identification and no homologue of Bombyx mori PTTH occurs in its genome. Here, we report Rhodnius PTTH is the first noggin-like PTTH found. It differs in important respects from known PTTHs and is the first PTTH from the Hemimetabola (Exopterygota) to be fully analysed. Recorded PTTHs are widespread in Holometabola but close to absent in hemimetabolous orders. We concluded Rhodnius PTTH likely differed substantially from the known ones. We identified one Rhodnius gene that coded a noggin-like protein (as defined by Molina et al., 2009) that had extensive similarities with known PTTHs but also had two additional cysteines. Sequence and structural analysis showed known PTTHs are closely related to noggin-like proteins, as both possess a growth factor cystine knot preceded by a potential cleavage site. The gene is significantly expressed only in the brain, in a few cells of the dorsal protocerebrum. We vector-expressed the sequence from the potential cleavage site to the C-terminus. This protein was strongly steroidogenic on PGs in vitro. An antiserum to the protein removed the steroidogenic protein released by the brain. RNAi performed on brains in vitro showed profound suppression of transcription of the gene and of production and release of PTTH and thus of ecdysteroid production by PGs. In vivo, the gene is expressed throughout development, in close synchrony with PTTH release, ecdysteroid production by PGs and the ecdysteroid titre. The Rhodnius PTTH monomer is 17kDa and immunoreactive to anti-PTTH of Bombyx mori (a holometabolan). Bombyx PTTH also mildly stimulated Rhodnius PGs. The two additional cysteines form a disulfide at the tip of finger 2, causing a loop of residues to protrude from the finger. A PTTH variant without this loop failed to stimulate PGs, showing the loop is essential for PTTH activity. It is considered that PTTHs of Holometabola evolved from a noggin-like protein in the ancestor of Holometabola and Hemiptera, c.400ma, explaining the absence of holometabolous-type PTTHs from hemimetabolous orders and the differences of Rhodnius PTTH from them. Noggin-like proteins studied from Hemiptera to Arachnida were homologous with Rhodnius PTTH and may be common as PTTHs or other hormones in lower insects.

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Artificial light at night (ALAN) is a recognised disruptor of biological function and ecological communities. Despite increasing research effort, we know little regarding the effect of ALAN on woody plants, including trees, or its indirect effects on their colonising invertebrates. These effects have the potential to disrupt woodland food webs by decreasing the productivity of invertebrates and their secretions, including honeydew and lerps, with cascading effects on other fauna. Here, we cultivated juvenile river red gums (Eucalyptus camaldulensis) for 40 weeks under experimentally manipulated light (ALAN) or naturally dark (control) conditions. To assess direct impacts on tree growth, we took multiple measures of growth at four time periods, and also measured physiological function, biomass and investment in semi-mature trees. To assess experimentally the direct and indirect (tree-mediated) impacts of ALAN on invertebrates, from 19 weeks onwards, we matched and mismatched trees with their original ALAN environments. We colonised trees with a common herbivore of E. camaldulensis, the red gum lerp psyllid (Glycaspis nr. brimblecombei) and then measured the effects of current and historic tree lighting treatment on the psyllid life cycle. Our data revealed direct effects of ALAN on tree morphology: E. camaldulensis trees exposed to ALAN shifted biomass allocation away from roots and into leaves and increased specific leaf area. However, while the intensity of ALAN was sufficient to promote photosynthesis (net carbon gain) at night, this did not translate into variation in tree water status or photosystem adaptation to dim night-time light for ALAN-exposed trees. We found some evidence that ALAN had broad-scale community effects-psyllid nymphs colonising ALAN trees produced more lerps-but we found no other direct or indirect impacts of ALAN on the psyllid life cycle. Our results suggest that trees exposed to ALAN may share morphological responses with trees under dim daylight conditions. Further, ALAN may have significant 'bottom-up' effects on Eucalyptus woodland food webs through both trees and herbivores, which may impact higher trophic levels including woodland birds, mammals and invertebrates.

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