Madagascar’s Central Highlands have experienced extensive deforestation and soil degradation, limiting the success of reforestation efforts. Poor soil fertility, particularly nitrogen limitation, constrains early seedling growth in degraded landscapes. This study evaluated the field performance of composted Black Soldier Fly frass (CBSFF) as a soil amendment for the native pioneer tree Dodonaea madagascariensis within the Ambohitantely Special Reserve. Four treatments were compared across four sites using a randomized complete block design: unfertilized control, cattle manure (4 g N), CBSFF one-fold (4 g N), and CBSFF two-fold (8 g N). The experiment was conducted on seedlings aged 16 months at the start of the study, and their growth was monitored over a six-month period. Growth responses were analyzed using generalized linear mixed-effects models with site included as a random factor. Seedling survival remained near 100% across all treatments, indicating no phytotoxic effects of composted frass under field conditions. Fertilization significantly enhanced both basal stem diameter and height growth. When standardized by nitrogen input, cattle manure and CBSFF produced comparable growth responses, indicating that nitrogen availability, rather than fertilizer identity, primarily drove early seedling performance. Height growth exhibited a clear dose-dependent response, with the double-dose CBSFF treatment producing the greatest increase. Planting method had a modest effect on height but did not alter the relative performance of fertilizer treatments. These findings demonstrate that composted BSF frass functions as an effective nitrogen source for early tree establishment in degraded tropical soils and performs comparably to traditional manure under field conditions. By validating insect-derived fertilizer within a restoration context, this study supports the integration of circular nutrient systems into sustainable reforestation strategies in biodiversity-rich yet resource-limited landscapes.
BackgroundLa Réunion Island (Mascarene Archipelago, south-western Indian Ocean) hosts a largely introduced ant fauna, shaped by historical and ongoing human-mediated introductions. Despite previous inventories, the ant fauna of the Island remains incompletely documented and updated faunistic records are needed to refine species checklists and improve knowledge of regional biodiversity. Documenting new occurrences contributes to a better understanding of species distributions, biogeographic patterns and invasion dynamics on oceanic islands, which are particularly vulnerable to biological invasions. New informationFollowing recent fieldwork, we report eight new species for La Réunion Island: Brachymymex australis Forel, 1901; Cardiocondyla obscurior Wheeler, 1929; Monomorium exiguum Forel, 1894; Pheidole parva Mayr, 1865; Solenopsis globularia Smith, 1858; Solenopsis gr. pygmaea, Stigmatomma cf. zwaluwenburgi Williams, 1946; and Strumigenys membranifera Emery, 1869. All are introduced species with varying invasiveness status. This work brings the total number of ants known from La Réunion Island to 62, although the presence and identification of some species cited in literature and databases needs verification. Further collections may uncover additional introduced species in urbanised and anthropogenised habitats and native species specific to La Réunion Island or the Mascarene Islands in natural ecosystems.
Insects have long been an integral part of Malagasy cuisine; however, their role in children’s lives outside traditional household contexts remains overlooked. This study examined the non-familial collection and consumption of edible insects among Malagasy children aged 6 to 15 years in rural communities of the Analamanga region in central Madagascar. Using two different assessment methods, direct questioning with adults and a game-based approach with children, we investigated children’s activities with edible insects during play. The findings revealed important differences between the two methods. Ethical compliance was confirmed by the institutional committee, which reviewed and approved the study design prior to data collection. Adults identified eight insect species consumed by children while playing, whereas children themselves reported consuming eleven species, suggesting that direct questioning may overlook certain aspects of children’s diets. The game-based approach proved to be a more effective tool in revealing hidden consumption patterns, as it allowed children to naturally show their interactions with insects. Overall, 35
Colour is a key trait involved in camouflage, physiological protection and thermoregulation. Yet environmental drivers of colour variation remain poorly understood at large spatial scales. Gloger's rule predicts animals should be darker in warmer and wetter climates, and in a complex version, redder in warmer and drier climates. Here, we present the first test of the complex Gloger's rule in insects using 34,331 images of 10,400 ant species across 586 assemblages worldwide. We decompose species mean colouration into two orthogonal axes, linked to darkness and redness. Assemblages were darker under high UV-B radiation and low dry-season precipitation, consistent with UV protection and desiccation resistance via melanisation. Canopy height increased both axes, suggesting camouflage. In contrast, higher mean temperature of the warmest quarter increased redness, as predicted by the complex Gloger's rule. Ant colouration cannot be explained by one macroecological rule but reflects environmental drivers acting independently on darkness and redness.
The big data era in biology is underway, but the study of organismal form has been slow to capitalize on advances in imaging and computation. Imaging approaches can digitize whole organisms, but low throughput has limited the effort to document morphological diversity. Here, within the open science initiative 'Antscan', we applied high-throughput synchrotron X-ray microtomography to capture phenotypes across a diverse and ecologically dominant insect group: ants. At https://www.antscan.info, we provide 2,193 whole-body three-dimensional ant datasets from 212 genera and 792 species to broadly cover the ant phylogeny with a global scope, also pairing phenomic data with genome sequencing projects. Scans acquired with standardized parameters facilitate automated analysis, and free access to data can broaden the audience and incentivize methods development. Antscan presents a scalable approach to create libraries of diverse anatomies, heralding an era of studies on the evolution, structure and function of organismal phenotypes.
The article is the fourth (and final) in a series of catalogues of ant type specimens from the collection of V. O. Karawajew (1864-1939), stored at the I.I. Schmalhausen Institute of Zoology of the National Academy of Sciences of Ukraine (SIZK, Kiev). This work includes data on 1530 type specimens (17 holotypes, one lecotype, 191 paralectotypes, one neotype and 1320 syntypes) from 220 taxa of the subfamily Myrmicinae, collected from around the world and described by Karawajew and other myrmecologists, which are currently assigned to 31 genera (Acromyrmex, Aphaenogaster, Apterostigma, Bondroitia, Cardiocondyla, Carebara, Cephalotes, Chelaner, Crematogaster, Goniomma, Lophomyrmex, Meranoplus, Messor, Monomorium, Mycocepurus, Myrmecina, Myrmica, Myrmicaria, Ocymyrmex, Oxyopomyrmex, Pogonomyrmex, Pheidole, Podomyrma, Pristomyrmex, Solenopsis, Stenamma, Strongylognathus, Strumigenys, Temnothorax, Tetramorium and Trichomyrmex).
Vietnam's diverse tropical habitats remain underexplored for ant biodiversity despite being part of the Indo-Burma biodiversity hotspot. This study presents the first detailed taxonomic revision of the ant genus Nylanderia Emery, 1906 in Vietnam, based on a combined approach using morphological characteristics and phylogenomic data. Using targeted enrichment of ultraconserved elements (UCEs) from 18 specimens and extensive morphological assessment of >200 individuals across all castes, we delimit 11 species from Vietnam. Three species are newly described: Nylanderia camon sp. nov., N. congtroi sp. nov., and N. didactica sp. nov., and one subspecies is elevated to species rank: N. donisthorpei (Forel, 1908) stat. nov. Our analysis reveals 3 new country-level records and clarifies the status of morphologically cryptic species through concordant genomic and morphological signals. All known Vietnamese Nylanderia species are diagnosed, with taxonomic accounts provided for each, including high-resolution montage images of all available castes and distribution maps. A revised illustrated key to the worker caste is also presented. This study highlights the utility of UCE-based phylogenetics for species delimitation within morphologically conservative taxa and contributes to a broader understanding of Nylanderia diversity in the Indomalayan region.
Male ants remain one of the least documented sources of morphological information in ant taxonomy, despite their frequent occurrence in biodiversity surveys and their potential value for systematic research. Within the ant subfamily Formicinae, identification resources for male ants are particularly limited in the Nearctic region. The only comprehensive treatment remains Smith’s (1943) key to male ants of the United States, which predates modern taxonomic revisions and advances in morphological terminology. In this work, a revised identification key to the genera of Nearctic Formicinae is presented, based exclusively on male morphology. An illustrated key to the 14 formicine genera recorded from the Nearctic region is provided, supported by updated diagnoses and figures of diagnostic characters, including genitalia and wing venation. This study provides the first modern synthesis of male morphological characters for Nearctic Formicinae and enables reliable identification of males, including specimens collected independently of workers in light traps, Malaise traps, and flight-intercept traps, thereby facilitating the broader integration of male morphology into taxonomic and biodiversity research on ants.
This study investigated whether cooked green beans, a discarded agro-industrial material, can partially replace commercial chicken feed to improve the performance of the edible cricket Gryllus madagascarensis, while accounting for the strong effects of rearing density. A two-by-two factorial experiment was conducted in which crickets were reared at low (500 individuals per box) or high density (2500 individuals per box) and fed either standard chicken feed or the same feed supplemented with cooked green beans, with twenty replicates per treatment. Survival, chicken feed consumption, biomass yield, frass production, efficiency of conversion of ingested feed, and approximate digestibility were measured over the rearing period. Survival increased from 30.0% to 32.9% at low density and from 11.7% to 13.2% at high density, while biomass yield increased from 117.2 g to 129.7 g and from 194.4 g to 231.2 g, respectively. Frass production also increased under supplementation. In contrast, individual body weight was not improved, indicating that higher biomass production resulted mainly from increased survival rather than faster growth. Rearing density remained a major determinant of performance, with low density favoring individual size and survival, and high density maximizing total biomass. Overall, partial replacement of commercial feed with cooked green beans improved survival, biomass yield, and frass production, supporting the use of plant-based wastes to enhance the sustainability of cricket farming.
In low-yield regions, yield intensification rather than land extensification offers the most efficient pathway to increase production and meet rising food demand. Yet, limited access to fertilizers remains a major constraint on crop productivity. The rapid expansion of insect farming has created a growing supply of an insect-derived organic fertilizer produced during insect rearing (frass); however, guidance on optimal field-scale application rates remains scarce. We conducted a large-scale field experiment in northern Madagascar to quantify yield responses of maize and soybean to Black Soldier Fly (Hermetia illucens) frass fertilizer and to identify optimal biophysical and economic application rates. Frass was applied at rates ranging from 0 to 30 t ha-1. For both crops, we did not identify a fertilizer rate that would maximize yield; instead, dry grain yield always increased with fertilizer rate and approached asymptotic maxima corresponding to 12.0- and 1.7-fold increases relative to unfertilized controls for maize and soybean, respectively. These rates are predicted to produce yields equivalent to 4.6- and 6.7-fold increases over national average yields, respectively. An economic analysis based on the marginal value-cost ratio (MVCR) indicated that the frass price is constrained by the profitability of soybean. Nevertheless, if using the national recommended dose of 3.6 t ha-1, a retail price of USD 13.8 t-1 would be highly attractive for soybean (MVCR = 3) and even more attractive for maize (MVCR = 5.6). This equates to just 2.2% of the current price of NPK fertilizer. Our results demonstrate that BSF frass can enhance crop yield and profitability in maize and soybean systems. While further research is necessary before these findings can be translated into large-scale application, this study establishes a foundational reference and framework toward achieving that goal. In Madagascar, intensifying maize production systems with organic fertilizers could increase yield and limit habitat destruction. Improved soybean productivity could reduce reliance on importation and diversify smallholder cropping systems. These findings highlight the promise of insect-derived fertilizers as a practical pathway toward sustainable intensification in low-yield agricultural contexts.
How communities are structured into functional groups and trophic layers is key to understanding ecosystem functioning. Nonetheless, we lack insights about spatiotemporal variation in guild composition of communities and its causes. To investigate spatial and temporal patterns and drivers of variation in insect feeding guilds, we combined data from a nationwide survey of Swedish insects using Malaise traps and DNA metabarcoding with a comprehensive trait database. We assigned species into one of three feeding guilds (phytophages, saprophages, predators) or into one of three associated parasitoid guilds. We then analysed patterns in species richness for each guild. Species richness declined with latitude in all guilds. Beyond this gradient, local variation in species richness matched between hosts and their parasitoids. Yet, hosts and their parasitoids responded differently to habitat. The phenological peak of parasitoid species richness appeared later than the peak of their hosts, but the length of time lags varied among guilds. Spatiotemporal patterns were driven by guild-specific responses to temperature, though much variation remained between seasons and locations even when controlling for temperature. Overall, these patterns suggest that shifts in both climate and land use may alter the synchrony of insect trophic layers, with unknown consequences.
Food insecurity remains a critical challenge for many farming communities in Madagascar, underscoring the need for sustainable strategies to improve crop yields. Black soldier fly frass (BSFF) is an emerging organic fertilizer, and composting may enhance its agronomic effectiveness. This study evaluated the effects of composted BSFF (CBSFF), fresh BSFF (FBSFF), cattle manure (CM), and a no-fertilizer control on maize (Zea mays L.) germination, height growth, grain yield, and agronomic efficiency of nitrogen (AEN) in southeastern Madagascar’s acidic sandy clay soils. Fertilizers were applied at 43 kg N/ha, reflecting actual farmer practices in the region. This rate represents one-third of the recommendation by FOFIFA (Madagascar’s national agricultural research institute). Farmers in this region rarely apply the full recommended dose due to limited access, high input costs, and low return on investment, factors that contribute to persistent yield gaps even where recommended rates are followed. Maize treated with FBSFF had significantly lower germination rate than all other treatments. In contrast, there were no statistically significant differences in final plant height or grain yield among CBSFF and CM. However, CBSFF-treated plants were on average 6 and 13% taller than those receiving CM and FBSFF, respectively, and produced 38% more grain than CM. All fertilizer treatments yielded significantly more grain than the unfertilized control, which produced no harvestable maize. CBSFF also showed the highest agronomic efficiency of nitrogen (AEN: 46 kg grain/kg N), although this difference was not statistically significant. These results suggest that BSFF, particularly in composted form, can supply sufficient nitrogen to support maize growth. Overall trends point to composted BSFF as the most agronomically effective option. Timing of application should depend on the frass form: composted BSFF is better suited for pre-planting use, while fresh BSFF may be more appropriate after germination. Even at sub-recommended doses, BSFF has the potential to enhance maize productivity and contribute to local food security. Further research across diverse agroecological zones, including trials comparing BSFF with the full recommended fertilizer dose, and economic analyses incorporating frass cost, labor, and transport are recommended to identify optimal application strategies and support widespread adoption by smallholder farmers.
Many insect groups have acquired obligate microbial symbionts, and the resulting associations can have important ecological and evolutionary consequences. A notable example among ants is the species‐rich tribe Camponotini, whose members derive nutritional benefits from a vertically inherited bacterial endosymbiont, Blochmannia . We generate ultraconserved element (UCE) phylogenomic data for 220 ingroup and 5 outgroup taxa to reconstruct a detailed evolutionary history of the Camponotini, including the inference of divergence times and dispersal events. Under multiple modes of analysis, including both concatenation and species‐tree approaches, we recover a well‐supported backbone phylogeny comprising eight lineages: three large genera ( Camponotus , Colobopsis , Polyrhachis ) and several smaller genera or clusters of genera. Three novel lineages are uncovered that cannot be placed in any existing genus: Lathidris gen. n ., from the mountains of Mesoamerica; Retalimyrma gen. n ., from the Indian Himalayas; and Uwari gen. n ., from eastern Asia. The species in these new genera were described and placed erroneously in Camponotus . The tribe Camponotini is estimated to have a crown origin in the Eocene (median age 38.4 Ma), with successively younger crown ages for Colobopsis (22.5 Ma), Camponotus (18.6 Ma) and Polyrhachis (18.5 Ma). We infer an Australasian or Indomalayan origin for the tribe, with multiple dispersal events to the Afrotropics, Palearctic region, and New World. Phylogenetic analysis of selected Blochmannia genes from a subset of 97 camponotine taxa yields results that are largely congruent with the ant host phylogeny, at least for well‐supported nodes, but we find evidence that Blochmannia from some old lineages—especially Lathidris —may have discordant histories, suggesting possible lability of this symbiosis in the early evolution of camponotine ants.
The genus-level classification of the ant subfamily Ponerinae (Hymenoptera: Formicidae) is revised based on a comprehensive phylogenomic analysis of more than 2,300 ultraconserved element (UCE) loci across 1,170 sampled specimens representing 1,020 taxa (600 valid species and 420 morphospecies) and all described ponerine genera known from workers. While most previously defined genus groups are recovered as monophyletic, several genera are shown to be polyphyletic or paraphyletic. To resolve these inconsistencies, four new genera are described: Boltonopone gen. nov., Makebapone gen. nov., Subiridopone gen. nov., and Sritoponera gen. nov. Xiphopelta stat. rev. is revalidated and Euponera is restricted by expanding Fisheropone to absorb a paraphyletic assemblage. Mesoponera is split into four lineages, resulting in transfers to Makebapone, Subiridopone, and Xiphopelta. Iroponera syn. nov. is synonymized under Cryptopone and additional new synonymies at both the generic and species levels are established. Morphological diagnoses are revised for each affected genus, and updated species lists and new combinations are provided. The updated classification recognizes 54 valid genera within Ponerinae and acknowledges an additional lineage that will be formally described in a subsequent publication. To support identification and comparative studies, revised keys to all extant Ponerinae genera are provided, presented by biogeographic region (African and Malagasy, Palearctic–Indomalaya–Australasia, and New World). This classification is intended to provide a stable, phylogenetically informed framework for future research on ponerine ants.
This comprehensive study examines the range and ecological interactions of Brachyponera sennaarensis (Mayr, 1862), a ponerine ant prevalent in sub-Saharan Africa and the Arabian Peninsula. Understanding the distribution patterns of ants is vital to developing sound long-term management strategies, as ants are critically involved in terrestrial ecosystems. This study employed species distribution modelling (SDM) via Maxent to forecast the present and future distributions of B. sennaarensis under two climate change scenarios: RCP 2.6, indicative of a climate change mitigation pathway, and RCP 8.5, denoting a high-emission scenario, for the years 2050 and 2070. The research utilized occurrence data from digital sources, implementing stringent filtration methods to guarantee data integrity. Crucial bioclimatic characteristics, such as annual mean temperature, mean diurnal range, minimum temperature of the coldest month, annual precipitation, and precipitation during the wettest quarter were recognized as key indicators of habitat compatibility. The findings demonstrate that the performance of the model is statistically dependable, as evidenced by an AUC value of 0.81 and a TSS value of 0.7. Our research predicts possible range changes and habitat degradation, especially in East Africa and the Horn of Africa, under the RCP 8.5 scenario, where rising temperatures and modified precipitation patterns may threaten the viability of this species. In contrast, the RCP 2.6 scenario indicates a slight northward extension of appropriate habitats in West Africa and the Sahel, underscoring the resilience of B. sennaarensis in more favorable climatic circumstances. This research highlights the significance of comprehending the ecological needs of B. sennaarensis in the context of climate change. It necessitates aggressive management strategies to alleviate the effects of climate change on this species and its habitats. The findings offer essential information for policymakers and management programs.
Edible insects are a significant component of traditional diets in Madagascar, where food insecurity and malnutrition persist. This study examines the production parameters and nutritional composition of four laboratory-farmed edible grasshopper species commonly consumed by Malagasy people with the aim of upscaling their farming to mitigate malnutrition. The grasshopper species include: vlei grasshopper (Paracinema tricolor), rice grasshopper (Oxya hyla), emerald-legged grasshopper (Eyprepocnemis smaragdipes), and Madagascan slant-faced grasshopper (Acrida madecassa). The study involved the assessment of production parameters (survival rate, developmental time, feed consumed, feed conversion ratio, biomass yield, fecundity, and hatchability). The study also involved analysis of the nutritional content (moisture, protein, fat, ash, fibre, carbohydrates, minerals, amino acids, fatty acids, and vitamins) to evaluate the potential dietary contribution of these grasshoppers. The result show P. tricolor had superior survival, faster development, low feed intake, and higher fecundity and hatchability when compared to other species of grasshoppers. Acrida madecassa showed the highest biomass yield and feed conversion ratio followed by P. tricolor. The results further show that all four species are rich in protein, essential fatty acids, and key minerals, particularly calcium, phosphorus, iron, and zinc. P. tricolor exhibited the highest protein and fat content. Moreover, P. tricolor showed the highest ash content, suggesting a superior mineral profile. Acrida madecassa showed the highest fibre content, reflecting its richness in chitin. These findings provide valuable insights into the nutritional role of grasshoppers in Malagasy diets. Furthermore, they offer reference values for selecting and optimizing the nutrient composition of insect species that are safe and easy to rear, which could serve as a sustainable alternative to wild collection. Future research should explore the bioavailability of nutrients in these species and identify suitable practices to mass rear these species to improve food security in Madagascar.
The tropical house cricket (Gryllodes sigillatus) can convert organic diets formulated from weeds and agro by-products into high-quality biomass. This study assessed the potential of diets developed from weeds and agro by-products as a feed source for G. sigillatus. We compared the development and nutritional value of crickets fed these alternative diets with control crickets fed chicken feed. Ten different diets with varying protein contents were used, including chicken feed (Control) with a protein content of 215 g/Kg dry matter (DM) basis), Cassava-Sugar Diet (250 g/Kg DM protein) Desmodium-Bran Diet (245 g/Kg DM protein), Morning Glory-Bean Diet (240 g/Kg DM protein), Morning Glory-Cassava Diet (235 g/Kg DM protein), Morning Glory-Cowpea Diet (225 g/Kg DM protein), Mixed Weed-Bran Diet (Optimal) (215 g/Kg DM protein) Cassava-Gallant Soldier Diet (200 g/Kg DM protein), Wheat-Bran Diet (145 g/Kg DM protein), and Maize-Cassava Diet (135 g/Kg DM protein). The weight and length of the crickets were measured for 9 weeks from day 1 after hatching to day 56. Then, the crickets were harvested and analyzed for dry matter, crude protein, fat, ash, fiber, minerals, and fatty acid composition. Cricket developmental time, survival rate, weight and length, yield, proximate components, and mineral and fatty acids differed depending on the diet provided. The Mixed Weed-Bran Diet (Optimal) resulted in the crickets developing faster (48.8 days), with a higher survival rate (88.1%), greater adult length (19.2 cm) and weight (0.44 g), and a nutrition content richer in minerals and unsaturated fatty acids when compared to other treatments. Oleic, linoleic, and palmitic acids were the major fatty acids. The highest protein content (64.4 g/100 g) was observed in the Mixed Weed-Bran Diet (Optimal) and Morning Glory-Cassava Diet treatments, while the Maize-Cassava Diet treatment crickets possessed the highest quantities of fats (19.1 g/100 g) and ash (15.4 g/100 g). The fatty acid profile of G. sigillatus revealed the cricket to have high unsaturated fatty acids except in crickets fed Morning Glory-Cowpea Diet and Wheat-Bran Diet. Generally, G. sigillatus grew best and had the most nutritious body composition on the Mixed Weed-Bran Diet (Optimal). The findings indicate that diets developed from weeds and agro by-products have great potential to be used as an alternative feed source for crickets and are capable of replacing expensive chicken feed, enhancing the circular farming potential of insect farming.
The delineation of zoogeographic regions is essential for understanding the evolution of biodiversity. Madagascar, characterized by high levels of endemism and habitat diversity, presents unique challenges and opportunities for such studies. Traditional global zoogeographic classifications, largely based on vertebrates, may overlook finer‐scale patterns of diversity. This study employs comprehensive ant distribution and phylogenomic datasets to propose a refined zoogeographic model for Madagascar. Utilizing phylogenetic Simpson's turnover, we identified three primary regions – Eastern, Northern, and Western – each characterized by distinct environmental and phylogenetic profiles. Further subdivision revealed nine subregions, reflecting variations in elevation, net primary productivity, and terrain ruggedness. Our findings highlight the importance of topographical and environmental barriers in shaping phylogenetic diversity and endemism. Notably, we observed significant phylogenetic clustering in lowland areas and distinct differences in net primary productivity and elevation across regions. This study underscores the value of integrating phylogenetic data in zoogeographic analyses and provides a nuanced framework for investigating biodiversity patterns in Madagascar, offering insights into the processes driving speciation and endemism on the island.