Beyond the Black Box: Reproductive Strategies of the Black Soldier Fly as a Model for Bridging Evolutionary Biology and Applied Entomology
Open accessAbstract
ABSTRACT The black soldier fly (BSF; Hermetia illucens ) is rapidly emerging as a model for evolutionary biology and insect biotechnology. Although larval biology has been extensively characterised, the reproductive biology of adults remains comparatively understudied. In this review, we synthesise the most recent empirical work on physiology, behavioural and chemical ecology to open the ‘black box’ of BSF reproduction, focusing on processes that span eclosion to senescence. We highlight pre‐ and post‐mating mechanisms that determine overall reproductive fitness: from mating latency, lekking dynamics, courtship and copulation, to sperm transfer, storage and oviposition. We discuss these processes within the framework of sexual selection theory. Several notable characteristics of BSF reproduction differ from traditional insect models. These include a hybrid capital‐income breeding strategy (adults do not need to feed but can benefit from supplemental nutrition), protandry (early male emergence), sex‐specific longevity that varies with mating status and a lek‐like mating system. In addition, females possess morphologically complex sperm‐storage organs, providing ample opportunity for intense post‐copulatory sexual selection. Recent work shows that environmental factors such as light, humidity, temperature, substrate volatiles and rearing design strongly influence reproductive output in industrial settings, highlighting the potential for BSF to bridge fundamental and applied research. We propose a novel conceptual framework that integrates these elements and outline key unresolved questions (e.g., mechanisms of sperm precedence, female control of fertilization, reproductive barriers, drivers of speciation etc.). This interdisciplinary model supports both fundamental insights into the evolution of reproductive traits and provides practical improvements for optimizing industrial mass‐rearing.
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TL;DR
This review synthesizes recent empirical work on adult black soldier fly reproduction, arguing that BSF can bridge evolutionary biology and applied entomology. It proposes a conceptual framework spanning emergence to senescence and identifies unresolved questions in pre- and post-copulatory sexual selection, reproductive barriers, and mass-rearing optimization.
Key findings
- BSF adults show a hybrid capital-income breeding strategy: they do not require feeding for gametogenesis, but supplemental nutrition can extend lifespan and enhance reproduction; larval nutrition remains the primary driver. Females carry 300–900 eggs, and fecundity is correlated with adult body size (Spearman’s ρ = 0.73).
- The mating system is lek-like and polygynandrous with protandry. Copulation involves a genital lock lasting approximately 33 min; initial matings last 32.5 ± 13.4 min and fifth matings 50.2 ± 26.6 min. Sperm transfer occurs late in copulation (e.g., after ~25 min), males possess ca. 3000–11,000 spermatozoa at eclosion with a lifetime limit of approximately 50,000, and BSF sperm are about 6 times longer than the average for internally fertilizing animals.
- Reproductive skew is pronounced: in equal-sex-ratio populations of 500 males and 500 females, only 43–120 mating events were observed over a week; in groups of 15 males and 15 females, nearly half of males never mated and half of successful males mated repeatedly. Females have complex sperm-storage organs, a single ejaculate can fill sperm reservoirs in excess, and females can lay two to three fertile clutches after even a single mating.
- Genomic and domestication evidence indicates global BSF COI divergences of 4.3%–4.9%, exceeding the standard 2%–3% barcode gap; 66% of allelic diversity resides in Neotropical strains, and domesticated or selectively bred lines show reduced heterozygosity and differentiation, raising the possibility of a cryptic species complex.
Methods
- Design: a review and synthesis of recent empirical work on BSF physiology, behavioural ecology, and chemical ecology, framed within sexual selection theory and culminating in a conceptual framework.
- Data or sample: synthesizes published studies plus some unpublished data, preprints, and personal observations across wild, laboratory, and industrial BSF populations; no new primary data are collected.
- Main analysis: qualitative integration of pre-copulatory, copulatory, and post-copulatory processes, environmental influences, and industrial implications, with identification of research gaps and future priorities.
Limitations
- The authors state that adult reproductive biology remains largely underexplored and that many mechanisms are unresolved, including sperm precedence, female control of fertilization, mate recognition, lek formation, reproductive barriers, and drivers of speciation.
- The authors note conflicting results and a lack of conclusive evidence in several areas, such as lekking in captivity, wing-buzzing and mating success, and inbreeding effects; much of what is known comes from captive or industrial populations and may not generalize to wild BSF.
- Evident from the text: the review relies on cited studies including preprints, unpublished data, personal observations, and small-scale behavioural assays, and it does not report a systematic review or meta-analysis.
Model-written summary. Check it against the paper before relying on it.
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Cites
Statistics
Claims the paper makes
84 of 242 cited sources checked · 27 supported · 2 flagged · accuracy not yet measured
- Premisecitation§ WILEY · p. 2
Environmental factors such as light spectra and intensity, specific substrate volatiles, and the physical design of mating cages can significantly influence mating success and fecundity in captive BSF.
“factors such as like light spectra and intensity (Zhang et al. 2010), specific substrate volatiles that attract females for oviposition (Zheng et al. 2013) and even the physical design of mating cages (Grosso et al. 2025) can significantly influence mating success and fecundity in captivity.”
Quote in the paperNot a substantial claim2 of 3 sources checked- cites Zhang et al. 2010 source
The Results and Discussion (pages unavailable in our copy) show that light source and intensity affected captive BSF mating, and that oviposition occurred under the quartz-iodine lamp. However, the Results report no statistically significant difference in egg masses between the sunlight and quartz-iodine treatments, so the claim of a significant effect on fecundity overstates this source. The substrate-volatiles and cage-design claims have separate citations and are not judged here. (LLM, conf=0.84)
Experimental — accuracy not yet measured - cites Zheng et al. 2013 source
The Discussion (page unavailable) reports that bacteria on decomposing material attract gravid black soldier flies, but says this occurs “presumably” through emitted volatiles; it does not identify specific attractive volatiles. The Results section on conspecific eggs and substrate preference reports differences in egg deposition, not mating success or fecundity. The separate claims about lighting and cage design are attributed to other citations. (LLM, conf=0.86)
Experimental — accuracy not yet measured - cites Grosso et al. 2025 source
- Premisecitation§ WILEY · p. 2
The asynchronous emergence of both sexes and of entire cohorts creates complex reproductive dynamics in BSF.
“The asynchronous emergence of both sexes and of entire cohorts creates complex reproductive dynamics (Lemke, Li, Dickerson, et al. 2025).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke, Li, Dickerson, et al. 2025 source
The Abstract reports that mixed-age breeding cohorts differ from same-age cohorts in mating frequency, egg production and fertility, supporting the point that age structure affects BSF reproduction. It does not say that asynchronous emergence of both sexes or entire cohorts creates that age structure. (LLM, conf=0.84)
Experimental — accuracy not yet measured
- Premisecitation§ WILEY · p. 2
Industrial studies suggest that reproduction in BSF can be responsive to environmental cues.
“Moreover, industrial studies suggests that reproduction in BSF can be responsive to environmental cues (Chia et al. 2018; Addeo et al. 2022).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Chia et al. 2018 source
- cites Addeo et al. 2022
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 2.1.3 | Courtship and Mounting · p. 5
In captivity, female BSF have been observed to momentarily leave their perch to enter male swarms occurring near artificial light sources before either returning to perch or descending in copula.
“In captivity, females have been observed to momentarily leave their perch (on artificial plants or walls) to enter male swarms occurring near artificial light sources (Lemke et al. 2024) before either returning to perch or descending in copula .”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke et al. 2024 source
The Discussion (§4.3) reports casual observations of females joining aerial swarms and then returning to their perches; §4.4 describes male mating swarms near lights in the captive cages. Our copy does not document females descending in copula after entering a swarm or identify walls as the perches they left. (LLM, conf=0.86)
Experimental — accuracy not yet measured
- Premisecitation§ 2.1.2 | Navigation and Lekking · p. 5
Observations of captive BSF populations suggest that flies may be guided by gradients in light, because newly eclosed adults from darkened pupation areas will be attracted towards illuminated zones.
“Observations of captive populations suggest that flies may be guided by gradients in light because newly eclosed adults from darkened pupation areas (Ferdousi and Sultana 2021) will be attracted towards illuminated zones (Dortmans et al. 2017; Coudron et al. 2025).”
Quote in the paperNot a substantial claim1 of 3 sources checked- cites Ferdousi and Sultana 2021
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Dortmans et al. 2017
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Coudron et al. 2025 source
The Good practices per stage section (p. 12) says pupae can be placed in an adjoining dark cage, and the Reproduction room section (p. 10) discusses light needed for mating. Neither passage says newly eclosed flies are attracted toward illuminated zones or guided by light gradients; searches for that movement found no supporting passage in our copy. (LLM, conf=0.70)
Experimental — accuracy not yet measured
- Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 5
Inadequate larval nutrition in BSF can generate smaller males with shorter sperm.
“Although inadequate larval nutrition in BSF can generate smaller males with shorter sperm (Zhang, Henawy, et al. 2025)”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Zhang, Henawy, et al. 2025 source
The Discussion (page unavailable) mentions nutritional supplements for BSF adults, while the Results and Conclusion address artificial light and oviposition. Searches for larval nutrition, male size, and sperm length found no passage in our copy addressing whether inadequate larval nutrition produces smaller males with shorter sperm. (LLM, conf=0.75)
Experimental — accuracy not yet measured
- Premisecitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6
BSF exhibit sexual dimorphism in wing interference patterns created by differences in structural colouration.
“BSF exhibit sexual dimorphism in wing interference patterns created by differences in structural colouration (Butterworth et al. 2021; Rebora et al. 2024).”
Quote in the paperNot a substantial claim1 of 2 sources checked - Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8
Female BSF can lay two to three fertile clutches after even a single mating.
“Originally these additional clutches were thought to be completely infertile (Nakamura et al. 2016) but female BSF can lay two to three fertile clutches after even a single mating (Manas et al. 2024).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Manas et al. 2024 source
- Interpretationcitation§ BOX 6 | (Continued) · p. 9
Housing mixed ages of BSF adults can reduce the efficiency of reproduction at the industrial scale.
“This means that housing mixed ages of BSF adults can reduce the efficiency of reproduction at the industrial scale (Lemke, Li, Dickerson, et al. 2025).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Lemke, Li, Dickerson, et al. 2025 source
The Abstract reports that mixed-age cohorts produced fewer eggs and less fertile eggs than same-age cohorts, supporting the concern about reproductive efficiency. But the experiment was conducted in greenhouse breeding cages; our copy does not establish the effect at industrial scale. (LLM, conf=0.84)
Experimental — accuracy not yet measured
- Findingcitation§ 2.2.2 | Sperm Storage · p. 7
Females possess complex sperm storage organs: three spherical spermathecae, attached to three sclerotised rods that end in hinges, connected to three separate fishnet canals containing sperm.
“Females possess complex sperm storage organs: three spherical spermathecae, attached to three sclerotised rods that end in hinges, which are connected to three separate fishnet canals (that contain sperm) (Munsch- Masset et al. 2023; Bruno et al. 2025).”
Quote in the paperNot a substantial claim2 of 2 sources checked- cites Munsch-Masset et al. 2023 source
The female-tract section describes three separate spermathecae, each with a sperm-containing reservoir, a rigid rod, an elbow, a ringed canal, and a fishnet canal that contains sperm after copulation. It calls the reservoirs ovoid rather than spherical and describes an elbow rather than a hinge; it also places a ringed canal between each elbow and fishnet canal. Page numbers were not supplied. (LLM, conf=0.86)
Experimental — accuracy not yet measured - cites Bruno et al. 2025 source
The 'Male and female reproductive tracts' section (p. 7) describes three spermathecae, fishnet canals that receive sperm, and rigid tubes forming an elbow before the sperm-storage reservoirs. It does not describe the spermathecae as spherical or the tubes as three sclerotised rods ending in hinges; it also places the reservoirs, rather than the fishnet canals, at the end of the storage pathway. (LLM, conf=0.86)
Experimental — accuracy not yet measured
- Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10
The highly specialised sperm storage organs of BSF females are capable of retaining more sperm than required to fertilise a single clutch.
“The highly specialised sperm storage organs of BSF females are capable of retaining more sperm than required to fertilise a single clutch (Manas et al. 2024).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Manas et al. 2024 source
- Interpretationcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10
Last-male sperm precedence may arise not just from males removing or displacing rivals' sperm, but if the female herself digests/ejects sperm from her genital tract prior to a subsequent mating.
“Last- male sperm precedence may arise-not just males remove or displace their rivals' sperm-but if the female herself digests/ ejects sperm from her genital tract prior to a subsequent mating (Luck et al. 2007; Schnakenberg et al. 2012).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Luck et al. 2007 source
- cites Schnakenberg et al. 2012 source
The sections “Female influences on sperm precedence, cryptic female choice” and “Storage of multiple ejaculates” describe female sperm dumping as a possible contributor to sperm precedence, alongside male-driven displacement. But the reported dumping is triggered by a second mating, not shown to occur before it, and the cited passages do not establish that females digest the sperm. No page numbers are available in our copy. (LLM, conf=0.88)
Experimental — accuracy not yet measured
- Interpretationcitation§ 3 | Implications for Evolutionary Biology · p. 10
The interaction between industrial selection pressures and ancestral reproductive behaviours such as the separation of mating sites from oviposition sites enables research to test how evolutionary 'holdovers' persist or decay in closed systems by comparing patterns in captivity to those in the wild.
“The interaction between industrial selection pressures and ancestral reproductive behaviours such as the separation of mating sites from oviposition sites (Tomberlin and Sheppard 2001) enables research to test of how evolutionary 'holdovers' persist or decay in closed systems (Lemke et al. 2024) by comparing patterns in captivity to those in the wild (Lemke, Smith, Smink, et al. 2025).”
Quote in the paperSubstantial claim1 of 3 sources checked - Interpretationcitation§ BOX 6 | (Continued) · p. 9
Older, less competitive males may interfere with females mating with younger mates, as observed in tephritid flies.
“For instance, older, less competitive males may interfere with females mating with younger mates, as observed in tephritid flies (Papanastasiou et al. 2011), which in these species has necessitated habitat design that features distinct areas for each sex (Meza et al. 2025).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Papanastasiou et al. 2011 source
The Discussion reports that older male medflies are less competitive and lose mating contests to younger males. It does not report that older males interfere with females mating with younger males; instead, it says older males fail to be chosen when young rivals are present. No page number is available in our copy. (LLM, conf=0.85)
Experimental — accuracy not yet measured - cites Meza et al. 2025 source
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 12
Overrepresentation of old or non-competitive males appears to depress productivity in BSF.
“overrepresentation of old or non- competitive males appears to depress productivity (Dickerson et al. 2024; Lemke, Li, Dickerson, et al. 2025)”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Dickerson et al. 2024 source
- cites Lemke, Li, Dickerson, et al. 2025 source
The Abstract reports that same-aged BSF cohorts mated more frequently and produced more eggs than mixed-age cohorts, and recommends removing old adults to improve yields. It does not establish that an overrepresentation of old or non-competitive males, specifically, depresses productivity; it also suggests older females and misplaced egg laying as possible contributors. (LLM, conf=0.86)
Experimental — accuracy not yet measured
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 11
Globally, 66% of allelic diversity resides within Neotropics BSF strains.
“Globally, 66% of allelic diversity resides within Neotropics strains”
Quote in the paperNot a substantial claim2 of 3 sources checked- cites Ståhls et al. 2020 source
The Results section on non-cultured flies reports that Neotropical field-collected flies had the highest number of distinct COI haplotypes: 32 of 56, or about 57%. This supports substantial Neotropical diversity, but not the claimed 66% of global allelic diversity in Neotropical strains; the reported measure is mitochondrial haplotypes in field-collected flies. (LLM, conf=0.86)
Experimental — accuracy not yet measured - cites Sandrock et al. 2021
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Kaya et al. 2021 source
The Results section (page unavailable) reports that South American BSF populations, considering both wild and captive provenances, contained 65% of the overall allelic variation. This supports the claim’s general point about high regional diversity, but the paper gives 65%, not 66%, and specifies South American populations rather than Neotropical strains as a whole. (LLM, conf=0.88)
Experimental — accuracy not yet measured
- Premisecitation§ 4.3 | Sex Ratios and Habitat Design · p. 12
Lek-like behaviours persist among captive BSF.
“Lek- like behaviours that persist among captive BSF (Tomberlin and Sheppard 2001; Lemke et al. 2023, 2024)”
Quote in the paperSubstantial claim1 of 3 sources checked - Premisecitation§ 4.1 | Genetics and Population Structure · p. 12
Size differences increase variation in mating outcomes in BSF.
“Size differences increase variation in such outcomes (Jones and Tomberlin 2021)”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Jones and Tomberlin 2021 source
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 11
High genetic diversity is present among global BSF strains.
“High genetic diversity is present among global BSF strains (Ståhls et al. 2020; Sandrock et al. 2021; Kaya et al. 2021; Generalovic et al. 2023)”
Quote in the paperNot a substantial claim2 of 3 sources checked - Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
BSF strains have divergences of 4.3%-4.9%.
“BSF strains have divergences of 4.3%-4.9% (Ståhls et al. 2020), which far exceeds the standard 2%-3% for a 'barcode gap' (Hebert et al. 2003).”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Ståhls et al. 2020 source
The Results (COI barcode dataset and analyses; page unavailable) report a maximum COI divergence of 4.9% across the full dataset. But the Results (COI haplotypes: cultured flies; page unavailable) give a maximum of 4.1% for rearing-culture samples, so the cited passage does not support a 4.3%–4.9% range specifically for BSF strains. (LLM, conf=0.88)
Experimental — accuracy not yet measured - cites Hebert et al. 2003 source
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 11
Industrial BSF strains often harbour a narrowed gene pool due to shared provenance, founding events, and repeated genetic bottlenecks.
“industrial strains often harbour a narrowed gene pool due to breeding stocks having shared provenance, founding events from colony establishments and repeated genetic bottlenecks (Ståhls et al. 2020; Sandrock et al. 2021; Kaya et al. 2021).”
Quote in the paperNot a substantial claim2 of 3 sources checked- cites Ståhls et al. 2020 source
The Background says many cultures began with Phoenix Worms, and the Results (COI haplotypes: cultured flies; page unavailable) find two shared mitochondrial haplotypes in cultures traced to that provider. This supports shared provenance for some strains, but the paper does not establish that industrial strains generally have a narrowed gene pool or attribute it to repeated genetic bottlenecks. (LLM, conf=0.86)
Experimental — accuracy not yet measured - cites Sandrock et al. 2021
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Kaya et al. 2021 source
The Discussion section ‘Demography and genetic signatures of domesticated BSF’ reports low allelic diversity and a pronounced founder effect in North American captive populations, with related strains sharing genetic footprints worldwide. It does not support repeated bottlenecks as a general explanation: the same section reports an absence of severe genetic bottlenecks across the domesticated BSF farms investigated. (LLM, conf=0.90)
Experimental — accuracy not yet measured
- Premisecitation§ 4.4 | Chemical Ecology and Oviposition Control · p. 12
BSF symbiotic microbiota community undergoes successive shifts throughout its ontogeny.
“the BSF symbiotic microbiota community undergoes successive shifts throughout its ontogeny (Klammsteiner et al. 2026).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Klammsteiner et al. 2026 source
- Premisecitation§ 4.4 | Chemical Ecology and Oviposition Control · p. 12
Synthetic blends combining multiple VOCs elicit stronger responses than individual components alone.
“Because synthetic blends combining multiple VOCs elicit stronger responses than individual components alone (Thomas et al. 2024)”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Thomas et al. 2024 source
In “Behavioural responses to artificial mixes” (p. 8), the five-compound Mixture 1 attracted oviposition, while neither of its two smaller submixtures was significantly attractive. The authors infer that more than one compound is needed, but they did not report a direct comparison with each individual component; the result also concerns black soldier fly oviposition rather than VOC blends and responses generally. (LLM, conf=0.84)
Experimental — accuracy not yet measured
- Interpretationreasoning§ 4.4 | Chemical Ecology and Oviposition Control · p. 12
A precise blend can be engineered as part of a push-pull strategy to direct egg-laying away from cage materials and towards the trap.
“this means that a precise blend can be engineered as part of a push-pull strategy (Menger et al. 2015; Cui et al. 2022) to more precisely direct egg- laying both away from cage materials and towards the trap.”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Menger et al. 2015 source
The Materials and Methods section describes a five-compound attractant blend used with a repellent in a push-pull system, supporting that general strategy. But the study concerns mosquitoes seeking human hosts and entering houses, not egg-laying on cage materials. The Discussion also says mosquitoes repelled from a house did not appear more likely to enter the trap, so it does not establish the claimed coordinated redirection. (LLM, conf=0.88)
Experimental — accuracy not yet measured - cites Cui et al. 2022 source
- Findingcitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6
Courtship in BSF involves the multimodal integration of visual, acoustic, vibratory and (presumably) olfactory and gustatory signals.
“Courtship in BSF involves the multimodal integration of visual, acoustic, vibratory and (presumably) olfactory and gustatory signals (Wicker- Thomas 2007).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Wicker-Thomas 2007 source
- Findingcitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6
These visual displays are accompanied by vibratory 'songs' produced by wing-buzzing.
“These visual displays are accompanied by vibratory 'songs' produced by wing- buzzing (Giunti et al. 2018; Laudani et al. 2024; Kortsmit et al. 2025), similar to many other groups of insects (e.g., crickets grasshoppers) (Baker et al. 2019).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Findingcitation§ BOX 3 | (Continued) · p. 7
If a female is receptive, successful pairs 'lock' their flexible genitalia, allowing them to rotate into a stable, tail-to-tail posture maintained during copulation while perched on a surface.
“If a female is receptive, successful pairs 'lock' their flexible genitalia (Rollinson et al. 2025), allowing them to rotate their position into a stable, tail- to- tail posture that is maintained during copulation while perched on a surface.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Rollinson et al. 2025 source
- Findingcitation§ BOX 3 | (Continued) · p. 7
When males mount females, they typically attempt genital engagement from behind.
“When males mount females, they typically attempt genital engagement from behind (Julita et al. 2020).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Julita et al. 2020 source
- Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7
Copulation in BSF is marked by genital engagement lasting approximately 33 min.
“Copulation in BSF is marked by genital engagement lasting approximately 33 min (Manas, Labrousse, and Bressac 2025), the onset of which is defined by a physical 'lock' between the male and female genitalia.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Manas, Labrousse, and Bressac 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7
The genital lock is made possible due to the morphology of the male terminalia (lateral extensions of the gonostylus) that bend medially inwards, functioning like a claw and allowing a male to grasp the female's terminalia.
“This is made possible due to the morphology of the male terminalia (i.e., the lateral extensions of the gonostylus) that bend medially inwards, effectively functioning like a claw and allowing a male to grasp the female's terminalia (Rollinson et al. 2025).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Rollinson et al. 2025 source
- Findingcitation§ BOX 3 | Debates on lekking in BSF. · p. 6
Mating in lone pairs of BSF has recently been described.
“and indeed mating in lone pairs of BSF has recently been described (Jensen et al. 2025).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Jensen et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Findingcitation§ 2.3.1 | Oviposition · p. 8
Females lay eggs in small, dry crevices near the actual oviposition substrate that is often wet and humid.
“Females lay eggs in small, dry crevices near the actual oviposition substrate that is often wet and humid (Sheppard et al. 2002).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Sheppard et al. 2002 source
- Findingcitation§ 2.3.2 | Aging and Senescence · p. 8
In BSF, aging can be estimated based on the decreasing opacity of the abdominal 'windows', through which one can observe changes in stored nutrient reserves.
“In BSF, because such physical changes are quite conspicuous, aging can be estimated based on the decreasing opacity of the abdominal 'windows' (Harjoko et al. 2023), through which one can observe changes in the stored nutrient reserves (which also include changes in colour).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Harjoko et al. 2023 source
- Findingcitation§ BOX 4 | Reproductive skews in BSF. · p. 7
Mating success in BSF is far from evenly distributed.
“Mating success in BSF is far from evenly distributed.”
Quote in the paperSubstantial claim - Findingreasoning§ 2.3.2 | Aging and Senescence · p. 8
Aging in adult BSF is often marked by visible changes in physical condition including wing damage, limb loss and desiccation.
“Aging in adult BSF is often marked by visible changes in the physical condition of flies including wing damage, limb loss and desiccation.”
Quote in the paperNot a substantial claim - Findingcitation§ 2.2.2 | Sperm Storage · p. 7
After copulation, BSF females store sperm in multiple spermathecae, enabling fertilization of several egg clutches over time.
“After copulation, BSF females store sperm in multiple spermathecae, enabling fertilization of several egg clutches over time.”
Quote in the paperNot a substantial claim - Findingcitation§ BOX 4 | Reproductive skews in BSF. · p. 8
The social environment or presence of rival males appears to influence sperm storage because females that mate in the presence of conspecifics retained more sperm than those isolated post-copulation.
“In addition, the social environment or presence of rival males appears to influence sperm storage because females that mate in the presence of conspecifics also retained more sperm than those isolated post- copulation (Manas, Labrousse, and Bressac 2025) (Box 5).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Manas, Labrousse, and Bressac 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Findingcitation§ 2.3.1 | Oviposition · p. 8
The timing of oviposition can be both genetically and environmentally influenced.
“The timing of oviposition itself can be both genetically and environmentally influenced.”
Quote in the paperNot a substantial claim - Findingcitation§ BOX 4 | Reproductive skews in BSF. · p. 7
No studies have addressed how genetic diversity might be maintained over time for BSF in light of the lekking paradox.
“Thus far, no studies have addressed how genetic diversity might be maintained over time for BSF in light of the lekking paradox (Kotiaho et al. 2001).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Kotiaho et al. 2001 source
- Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10
This often occurs in response to perceived sperm competition risk.
“This often occurs in response to perceived sperm competition risk (Manas, Labrousse, and Bressac 2025).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Manas, Labrousse, and Bressac 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10
These seminal fluid proteins act to increase male fitness at the cost of female fitness.
“These act to increase male fitness at the cost of female fitness (Wigby and Chapman 2005).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Wigby and Chapman 2005 source
- Premisecitation§ WILEY · p. 2
BSF exhibit protandry, with males emerging several days before females.
“Black soldier flies also exhibit protandry. That is, males emerge several days before females (Tomberlin et al. 2002)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Tomberlin et al. 2002
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ ABSTRACT · p. 1
The black soldier fly (BSF; Hermetia illucens) is rapidly emerging as a model for evolutionary biology and insect biotechnology.
“The black soldier fly (BSF; Hermetia illucens ) is rapidly emerging as a model for evolutionary biology and insect biotechnology.”
- Also stated in: § 1 | Introduction · p. 1 · plain
“BSF is increasingly recognised as a model organism for fundamental studies in evolutionary biology and behavioural ecology (Tomberlin et al. 2025)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tomberlin et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Also stated in: § 1 | Introduction · p. 1 · plain
- Premisereasoning§ ABSTRACT · p. 1
The reproductive biology of adult BSF remains comparatively understudied relative to larval biology.
“Although larval biology has been extensively characterised, the reproductive biology of adults remains comparatively understudied.”
- Also stated in: § 1 | Introduction · p. 1 · plain
“the adult reproductive biology remains largely underexplored”
Quote in the paperSubstantial claim - Also stated in: § 1 | Introduction · p. 1 · plain
- Premisecitation§ 2.1.3 | Courtship and Mounting · p. 5
Courtship in BSF is typically initiated by female flight or entry into a swarm.
“Courtship is typically initiated by female flight or entry into a swarm.”
Quote in the paperNot a substantial claim - Premisecitation§ 2.1.2 | Navigation and Lekking · p. 5
A recent study in Costa Rica failed to find any mating BSF within a radius of 30 m, despite the prevalence of female flies within 7.5 m of an oviposition site.
“a recent study in Costa Rica failed to find any mating BSF within a radius of 30 m, despite the prevalence of female flies within 7.5 m of an oviposition site (Lemke, Smith, Smink, et al. 2025).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Lemke, Smith, Smink, et al. 2025 source
- Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4
Larval nutritional legacy remains the primary driver of BSF reproductive potential.
“But despite any love of sugar the adults might possess, larval nutritional legacy remains the primary driver of BSF reproductive potential (Lemke et al. 2023; Harjoko et al. 2023).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Lemke et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Harjoko et al. 2023 source
- Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 5
In response to perceived sperm competition risk from other males, BSF males can adjust their sperm investment, indicating an adaptive allocation of ejaculates.
“In response to perceived sperm competition risk from other males, BSF males can adjust their sperm investment, indicating an adaptive allocation of ejaculates (Manas, Labrousse, and Bressac 2025)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Manas, Labrousse, and Bressac 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 2.1.2 | Navigation and Lekking · p. 5
The exact mechanisms by which BSF locate mating sites in the wild remain largely unknown.
“To date, the exact mechanisms by which BSF locate mating sites in the wild remain largely unknown (Lemke et al. 2023).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Lemke et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation§ 2.1.2 | Navigation and Lekking · p. 5
Spatial segregation between larval media/oviposition sites and mating sites is a key feature of BSF reproductive ecology.
“A spatial segregation between the larval media/oviposition sites with mating sites is a key feature of BSF reproductive ecology”
Quote in the paperSubstantial claim - Premisenone given§ ABSTRACT · p. 1
Recent work shows that environmental factors such as light, humidity, temperature, substrate volatiles and rearing design strongly influence reproductive output in industrial settings.
“Recent work shows that environmental factors such as light, humidity, temperature, substrate volatiles and rearing design strongly influence reproductive output in industrial settings”
Quote in the paperNot a substantial claim - Premisecitation§ 2.1.2 | Navigation and Lekking · p. 5
Natural BSF lekking sites have been described as occurring hundreds to potentially thousands of meters from emergence zones.
“Although natural lekking sites have been described as occurring hundreds to potentially thousands of meters from emergence zones (Tomberlin and Sheppard 2001; Lemke et al. 2023)”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Tomberlin and Sheppard 2001 source
- cites Lemke et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation§ 2.1.3 | Courtship and Mounting · p. 5
The observed female BSF behaviour of leaving perch to enter male swarms appears to mirror natural history descriptions in which females visit the lek where males are already present.
“This observed behaviour appears to mirror natural history descriptions in which females visit the lek where males are already present (Tomberlin and Sheppard 2001; Lemke et al. 2023).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Tomberlin and Sheppard 2001 source
- cites Lemke et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4
As the size difference between female and male BSF increases, so does the degree of sexual bimaturism.
“As the size difference between females and males increases, so does the degree of SBM (Teder et al. 2021; Kortsmit et al. 2025).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Interpretationreasoning§ 2.1.2 | Navigation and Lekking · p. 5
Because females are readily attracted to rotting substrates, field observations are often biased towards ovipositing females near anthropogenic wastes.
“Because females are readily attracted to rotting substrates, field observations are often biased towards these ovipositing females near anthropogenic wastes.”
Quote in the paperNot a substantial claim - Interpretationreasoning§ 2.1.1 | Latency and Sexual Maturation · p. 5
BSF's synovogenic egg development strategy is a likely adaptation to stochastic nutritional environments that BSF larvae often inhabit.
“This strategy in BSF is a likely adaptation to stochastic nutritional environments that BSF larvae often inhabit (i.e., in heterogenous wastes).”
Quote in the paperNot a substantial claim - Premisecitation§ 2.1.2 | Navigation and Lekking · p. 5
Mating BSF adults are conspicuously absent from locations where females are attracted to rotting substrates.
“However, mating adults are conspicuously absent from these same locations.”
Quote in the paperSubstantial claim - Interpretationreasoning§ BOX 2 | Debates on mate discrimination in BSF. · p. 6
For BSF, SSS behaviours are largely an artifact of their captivity and of laboratory studies using small enclosures.
“This suggests that for BSF, SSS behaviours are largely an artifact of their captivity and of laboratory studies using small enclosures, e.g., as small as 30 × 30 × 30 cm (Nakamura et al. 2016).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Nakamura et al. 2016
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation§ BOX 3 | Debates on lekking in BSF. · p. 6
BSF should still be able to mate in the absence of competitors.
“Instead, it follows that BSF should still be able to mate in the absence of competitors and indeed mating in lone pairs of BSF has recently been described (Jensen et al. 2025).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Jensen et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ BOX 3 | Debates on lekking in BSF. · p. 6
Under high population densities and when males are unable to control access to females, this leads to scramble competition polygyny.
“For instance, under high population densities and when males are unable to control access to females, this leads to scramble competition polygyny (Herberstein et al. 2017).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Herberstein et al. 2017
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7
BSF might not engage in mate guarding.
“suggesting that BSF might not engage in mate guarding, which in other species can arise as a mating tactic to prevent females from mating with rivals.”
Quote in the paperNot a substantial claim- Rests on an assumption: Once the genital lock commences, rival males cease interruption attempts.
- Interpretationreasoning§ WILEY · p. 2
BSF's largely capital breeding (but hybrid) strategy has critical evolutionary implications.
“This largely capital breeding (but hybrid) strategy has critical evolutionary implications”
Quote in the paperNot a substantial claim - Interpretationreasoning§ BOX 5 | Re- mating in BSF. · p. 8
These findings highlight the potential for strong postcopulatory sexual selection that influence ejaculate-female interactions and overall reproductive fitness in BSF.
“These findings highlight the potential for strong postcopulatory sexual selection that influence ejaculate- female interactions and overall reproductive fitness in BSF.”
Quote in the paperNot a substantial claim - Interpretationnone given§ 3 | Implications for Evolutionary Biology · p. 10
Reproductive barriers may be driven by local adaptations to different feedstocks, rearing conditions and selection regimes that produce phenotypic differences (e.g., microbiota, morphology, behaviour).
“Besides the obvious isolation of these strains from one another in time and space, reproductive barriers may be driven by local adaptations to different feedstocks, rearing conditions and selection regimes that produce phenotypic differences (e.g., microbiota, morphology, behaviour).”
Quote in the paperSubstantial claim - Interpretationcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10
Retention duration, sperm viability and selective sperm use suggest potential for cryptic female choice in BSF, as described in other stratiomyids.
“Retention duration, sperm viability and selective sperm use suggest potential for cryptic female choice as described in other stratiomyids (Barbosa 2009, 2011, 2012, 2015).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
The study identified signatures of selection across eleven BSF populations.
“the study identified clear population structure, reduced heterozygosity in long- term domesticated lines and signatures of selection across eleven BSF populations”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
COI haplotype diversity in BSF is high enough to possibly warrant considering BSF a species complex.
“These results echo earlier work using mitochondrial DNA, indicating that cytochrome oxidase subunit I ( COI ) haplotype diversity is high enough to possibly warrant considering BSF a species complex.”
Quote in the paperSubstantial claim - Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
The study identified clear population structure across eleven BSF populations.
“the study identified clear population structure, reduced heterozygosity in long- term domesticated lines and signatures of selection across eleven BSF populations”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
Hybrids between BSF strains could suffer reduced fitness, especially under intense selection regimes typical of industrial rearing.
“the observed genomic divergence and reduced heterozygosity in domesticated lines raise the possibility that hybrids could suffer reduced fitness, especially under the intense selection regimes typical of industrial rearing (Silvaraju et al. 2026).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 11
Some studies document inbreeding depression and reduced progeny in captive BSF populations.
“although some studies document inbreeding depression and reduced progeny in captive populations (Rhode et al. 2020; Laudani et al. 2024)”
Quote in the paperNot a substantial claim0 of 2 sources checked - Interpretationreasoning§ 4.2 | Nutritional Management · p. 12
Supplementing the nutrition of adult BSF breeding stocks may be necessary to sustain high fecundity, especially when larvae are fed on abundant but low-quality feed stocks.
“supplementing the nutrition of adult breeding stocks may be necessary to sustain high fecundity, especially to compensate for the fact that waste remediation efforts by industry will necessarily require larvae to be fed on”
Quote in the paperSubstantial claim - Interpretationcitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
Selective breeding and prolonged captive rearing potentially accelerate the emergence of reproductive barriers such as mechanical isolation or reduced hybrid fitness in BSF.
“potentially accelerating the emergence of reproductive barriers such as mechanical isolation or reduced hybrid fitness.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 11
Reduced heterozygosity is detected in captive BSF colonies.
“The detection of reduced heterozygosity in captive colonies (Rhode et al. 2020; Generalovic 2023; Silvaraju et al. 2026)”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Rhode et al. 2020 source
- cites Generalovic 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 4.2 | Nutritional Management · p. 12
Larval diet can influence adult reproductive strategies in BSF.
“because larval diet can influence adult reproductive strategies (Kortsmit et al. 2025)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Kortsmit et al. 2025 source
- Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
Male BSF genitalia are hypo-allometric.
“a recent study shows that male BSF genitalia are hypo- allometric (Rollinson et al. 2025)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Rollinson et al. 2025 source
- Interpretationreasoning§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
BSF may consist of multiple genetically distinct lineages that are on independent evolutionary trajectories.
“Together, the mitochondrial and nuclear evidence suggest that BSF may consist of multiple genetically distinct lineages that are on independent evolutionary trajectories.”
Quote in the paperSubstantial claim - Premisecitation§ 4 | Implications for Industrial Applications · p. 11
Adult reproduction is the rate-limiting step in closed-cycle BSF rearing.
“adult reproduction is the rate- limiting step in closed- cycle rearing (Boller 1972)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Boller 1972 source
- Premisecitation§ 4.1 | Genetics and Population Structure · p. 11
Larval nutritional legacy affects downstream reproductive outcomes in BSF.
“given the effects of larval nutritional legacy on downstream reproductive outcomes.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Kortsmit et al. 2025 source
- Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
A recent population genetic study of wild and industrial BSF strains revealed extensive rapid differentiation that can occur under domestication.
“A recent population genetic study of wild and industrial BSF strains revealed extensive rapid differentiation that can occur under domestication (Silvaraju et al. 2026).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationcitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
Genital size is unlikely to be a physical barrier to reproduction in BSF.
“and thus is unlikely to be a physical barrier to reproduction.”
Quote in the paperSubstantial claim - Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11
The study identified reduced heterozygosity in long-term domesticated BSF lines.
“the study identified clear population structure, reduced heterozygosity in long- term domesticated lines and signatures of selection across eleven BSF populations”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation§ 4.2 | Nutritional Management · p. 12
Providing BSF adults supplemental nutrition pushes them slightly towards the middle of the capital-income continuum.
“Although classically described as capital breeders, providing BSF adults supplemental nutrition pushes them slightly towards the middle of the capital- income continuum (Davis et al. 2016).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Davis et al. 2016 source
- Premisecitation§ 4.3 | Sex Ratios and Habitat Design · p. 12
BSF microhabitat preference is related to their age.
“A recent report indicates that BSF microhabitat preference is related to their age”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Salari and De Goede 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisenone given§ 6 | Conclusions and Future Directions · p. 14
BSF displays a capital-income breeding hybrid strategy.
“BSF displays a capital- income breeding hybrid strategy, allowing the study of reproductive outcomes that are mediated strictly by larval nutrition, as well as enhanced or otherwise affected by adult nutrition.”
Quote in the paperNot a substantial claim - Interpretationreasoning§ 6 | Conclusions and Future Directions · p. 14
BSF is well-poised to be used as a research model to investigate questions in both fundamental evolutionary and applied sciences such as sustainable agriculture, waste management, biotechnology and insect welfare.
“BSF is well- poised to be used as a research model to investigate questions in both fundamental evolutionary and applied sciences such as sustainable agriculture, waste management, biotechnology and insect welfare.”
Quote in the paperNot a substantial claim
- Classified as probably not a substantial claim. 39
- May combine two claims. 82
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8Several recent studies suggest female BSF can oviposit multiple times, though others argue BSF oviposit only once.
“Several recent studies suggest female BSF can oviposit multiple times (Jones and Tomberlin 2021; Hoffmann et al. 2021; Chiabotto et al. 2024; Laudani et al. 2024), though others argue BSF oviposit only once (Sibonje 2024).”
Quote in the paperNot a substantial claim2 of 3 sources checked- cites Jones and Tomberlin 2021 source
The Oviposition and hatch rate section reports egg counts by day for populations, not repeated oviposition by individual females. The Effects of body size on oviposition and hatch rate section discusses multiple mating, which is distinct from laying eggs multiple times. Searches for repeated oviposition and oviposition frequency found no passage addressing the attributed claim; page numbers were unavailable in our copy. (LLM, conf=0.76)
Experimental — accuracy not yet measured - cites Hoffmann et al. 2021 source
The Introduction (page unavailable) says BSF females produce only one viable egg clutch in their lifetime. The Discussion (page unavailable) finds evidence of multiple mating and paternity, not multiple oviposition; the claim appears to conflate mating with egg-laying. (LLM, conf=0.84)
Checked by a reasoning model — may be wrong.
Experimental — accuracy not yet measured - cites Chiabotto et al. 2024 source
May combine two claims.
Premisecitation§ 4.3 | Sex Ratios and Habitat Design · p. 12Increased perching area provided by plants increases mating rates in BSF, with magnitude varying with plant density.
“the increased perching area provided by the plants has been shown to increase mating rates, although the magnitude of the effects varies with the density of the plants provided (Lemke et al. 2024; Grosso et al. 2025).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Lemke et al. 2024 source
The Methods (§2.6) identify mating events as a fitness metric tested against plant treatment and leaf area. The Results (§3.1) report no statistical effect of treatment on any tested fitness metric, contrary to the claimed increase in mating rates with plant density. Page numbers are not available in our copy. (LLM, conf=0.89)
Checked by a reasoning model — may be wrong.
Experimental — accuracy not yet measured - cites Grosso et al. 2025 source
May combine two claims.
Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 5Female fecundity in BSF is tightly correlated with adult body size (Spearman's ρ = 0.73), which depends on larval diet.
“female fecundity in BSF is instead tightly correlated with adult body size (Spearman's ρ = 0.73) which depends on larval diet (Gobbi et al. 2013; Shrestha et al. 2025).”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Gobbi et al. 2013 source
The Discussion (page not supplied) says larger BSF females were more fertile, and the Adult size section reports that adult size differed by larval diet. But the paper does not report a Spearman correlation of ρ = 0.73 between fecundity and body size: its Statistical analysis section specifies Pearson correlations for body size and ovarian measurements, and Table 3’s 0.73 values concern those measurements. (LLM, conf=0.90)
Experimental — accuracy not yet measured - cites Shrestha et al. 2025 source
May combine two claims.
Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4BSF are not strict capital breeders and supplemental nutrition can benefit adult fitness, though this is not always consistently observed.
“However, BSF are not strict capital breeders and supplemental nutrition can benefit adult fitness (Thinn and Kainoh 2022; Klüber et al. 2023; Barrett et al. 2025), though this is not always consistently observed (Lemke et al. 2023; Coudron et al. 2025).”
Quote in the paperNot a substantial claim2 of 3 sources checkedMay combine two claims.
Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4Neurobehavioural assays confirm that BSF can taste and respond to sugar, with females responding more strongly than males.
“Neurobehavioural assays confirm that BSF can indeed taste and respond to sugar, with females responding more strongly than males (Merle et al. 2026).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Merle et al. 2026 source
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation§ 1 | Introduction · p. 1BSF larvae are widely studied for converting organic waste into protein and fertiliser, making BSF a keystone species in sustainable agriculture and circular food systems.
“As an efficient decomposer of organic waste with a high degree of nutritional flexibility, BSF larvae are widely studied for converting organic waste into protein and fertiliser, making it a keystone species in sustainable agriculture and circular food systems (Tomberlin and van Huis 2020; Tomberlin et al. 2022).”
Quote in the paperNot a substantial claim2 of 2 sources checked- cites Tomberlin and van Huis 2020 source
The Introduction (pp. 1–2) describes BSF as a widely studied model for recycling organic waste into animal feed while producing frass usable as fertiliser. The Conclusions (p. 3) call it a key asset for a circular economy. The paper does not establish the claimed high nutritional flexibility or describe BSF as a keystone species in sustainable agriculture and circular food systems. (LLM, conf=0.85)
Experimental — accuracy not yet measured - cites Tomberlin et al. 2022 source
The Abstract names black soldier fly among the three insects prominent in mass production and describes insect-agriculture outputs including protein and frass. It also links the broader sector to sustainability and circular economies. It does not specifically say that BSF larvae convert organic waste into protein and fertiliser, have high nutritional flexibility, or are a keystone species. (LLM, conf=0.84)
Experimental — accuracy not yet measured
May combine two claims.
Premisecitation§ 2.1.3 | Courtship and Mounting · p. 5Male BSF flight activity peaks in the late morning and early afternoon, followed by a peak in female activity later in the day.
“The timing of these interactions is structured, with male flight activity peaking in the late morning and early afternoon, followed by a peak in female activity later in the day (Lemke et al. 2024).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke et al. 2024 source
The Abstract links early-hour perching patterns to males engaging in aerial contests, and Discussion §4.3 says mating intensity generally peaks by 12:00 under artificial light or 15:00 under natural light. But the paper measured perching and mating events, not sex-specific flight-activity peaks. It only suggests that females might leave perches to join mating swarms later in the day; it does not establish a later female flight peak. (LLM, conf=0.82)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6Visual neural pathways are more developed in male BSF than in females.
“Evidence suggests that visual neural pathways are more developed in males than females (Barrett, Godfrey, et al. 2022)”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Barrett, Godfrey, et al. 2022 source
May combine two claims.
Premisecitation§ 2.1.2 | Navigation and Lekking · p. 5BSF larvae are negatively phototropic and often seek dark crevices prior to pupation; the subsequent positive phototaxis of adults would guide them out of emergence sites and into open, illuminated areas where mating can occur.
“BSF larvae are negatively phototropic and often seek dark crevices prior to pupation; the subsequent positive phototaxis of adults would guide them out of emergence sites and into open, illuminated areas where mating can occur (Giannetti et al. 2022).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Giannetti et al. 2022 source
The Introduction (page unavailable) says prepupae avoid light and seek shelter to pupate, supporting the larval portion of the claim. The cited paper does not establish that adults are positively phototactic or that light guides them from emergence sites to mating areas. (LLM, conf=0.88)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6BSF's wing interference patterns putatively are detected visually thanks to high photoreceptive sensitivity of the ommatidia in BSF compound eyes.
“These honest signals putatively are detected visually thanks to high photoreceptive sensitivity of the ommatidia in BSF compound eyes (Oonincx et al. 2016).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Oonincx et al. 2016 source
Our copy contains only the paper’s title and bibliographic details (no section or page). The title identifies a study of BSF compound-eye photoreceptor spectral sensitivity, but the available text does not address whether ommatidia are highly sensitive or whether BSF visually detect wing interference patterns. Searches for those points returned no further passages. (LLM, conf=0.75)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing).
Interpretationcitation§ BOX 3 | (Continued) · p. 7BSF might alter the harmonics of their buzzing by rapidly modulating the frequency of their wingbeats, as is the case in Aedes aegypti.
“BSF might alter the harmonics of their buzzing by rapidly modulating the frequency of their wingbeats, as is the case in Aedes aegypti (Diptera: Culicidae) (Aldersley and Cator 2019).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Aldersley and Cator 2019 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingcitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6Males perform aerial displays of wing-fanning, which combined with wing-interference patterns (WIPs) putatively serve as visual signals for conspecifics.
“Males perform aerial displays of wing- fanning, which combined with wing- interference patterns (WIPs) putatively serve as visual signals for conspecifics (Rebora et al. 2024).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Rebora et al. 2024 source
The Methods (Behavioural experiments; page unavailable) record males fanning their wings on coloured cardboard disks, not performing aerial wing-fanning displays. The Discussion (page unavailable) suggests that wing structural colours, including WIPs, may serve as mating signals, but it does not establish that male wing-fanning and WIPs act together as visual signals to conspecifics. (LLM, conf=0.86)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7The duration of the genital lock appears mostly unaffected by adult size but varies with age.
“Moreover the duration of the genital lock appears mostly unaffected by adult size but varies with age (Manas, Labrousse, and Bressac 2025; Manas, Venon, et al. 2025), suggesting that BSF might not engage in mate guarding, which in other species can arise as a mating tactic to prevent females from mating with rivals.”
Quote in the paperSubstantial claim1 of 2 sources checkedMay combine two claims.
Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7Initial matings might last between 32.5 ± 13.4 (mean ± SD) minutes, and mating duration generally increases with each successive mating up to 50.2 ± 26.6 min for the fifth mating.
“Initial matings might last between 32.5 ± 13.4 (mean ± SD) minutes, though research suggests mating duration generally increases with each successive mating up to 50.2 ± 26.6 min for the fifth mating (Manas, Venon, et al. 2025).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Manas, Venon, et al. 2025 source
The Copulation duration section reports 32.5 ± 13.4 minutes for the first male mating and an increase through successive matings. But it assigns 50.2 ± 26.6 minutes to the fourth mating, not the fifth; Table 1 notes that no male mated five times. (LLM, conf=0.94)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ BOX 3 | Debates on lekking in BSF. · p. 6Some studies report a clear behavioural segregation of the sexes, but other aspects of lekking are often not reported because most detailed behavioural studies in BSF are typically done at small scale, because manual observations become impossible even at modest densities.
“Although some studies report a clear behavioural segregation of the sexes (Lemke et al. 2024), other aspects of lekking are often not reported because most detailed behavioural studies in BSF are typically done at small scale, because manual observations of BSF behaviour become impossible even at modest densities.”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke et al. 2024 source
Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 3 | Debates on lekking in BSF. · p. 6The mating system of BSF is described as 'lekking' or 'lek-like', in which males aggregate at spatially distinct sites which females visit strictly to mate.
“The mating system of BSF is described as 'lekking' (Tomberlin and Sheppard 2001) or 'lek- like' (Birrell 2018), in which males aggregate at spatially distinct sites which females visit strictly to mate (Alcock 1990).”
Quote in the paperNot a substantial claim1 of 3 sources checked- cites Tomberlin and Sheppard 2001 source
- cites Birrell 2018
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Alcock 1990 source
The Introduction and “Resource-defense or lek polygyny?” sections describe males of Hermetia comstocki defending landmark plants that females visit to mate, probably without obtaining other resources. But the cited study concerns H. comstocki, not the black soldier fly (BSF), and its conclusion about females obtaining no resources is qualified rather than strict. (LLM, conf=0.87)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ BOX 3 | Debates on lekking in BSF. · p. 6Some field and lab observations document stable male aggregations, whilst other reports mention no aggregation except within ephemeral mating balls involving multiple males.
“Some field and lab observations document stable male aggregations (Tomberlin and Sheppard 2001; Lemke et al. 2024), whilst other reports mention no aggregation except within ephemeral mating balls involving multiple males (Permana et al. 2020), although these might be incorrectly mistaken for leks.”
Quote in the paperNot a substantial claim2 of 3 sources checked- cites Tomberlin and Sheppard 2001 source
- cites Lemke et al. 2024 source
The Discussion (§4.4, page unavailable) reports male aerial swarms in captivity and prolonged bouts of mating behavior near lights, supporting the observation of male aggregations. It does not establish that those aggregations were stable, rather than recurring or changing over time. The separate claim about ephemeral mating balls is attributed to Permana et al., not Lemke et al. (LLM, conf=0.81)
Experimental — accuracy not yet measured - cites Permana et al. 2020 source
The Abstract describes mating success and mate virginity in semi-outdoor cages, but does not discuss male aggregations. Searches for mating balls, multiple-male groups, and aggregations returned no relevant passage from our copy, so the claim attributed to Permana et al. cannot be verified. (LLM, conf=0.72)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingcitation§ BOX 3 | Debates on lekking in BSF. · p. 6Lekking behaviour has been described in one other congener, the Agave soldier fly, Hermetia comstockii.
“This behaviour has been described in one other congener, the Agave soldier fly, Hermetia comstockii (Diptera: Stratiomyidae) (Alcock 1990).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Alcock 1990 source
May combine two claims.
Findingcitation§ BOX 3 | Debates on lekking in BSF. · p. 6More distantly related soldier flies do not lek: Merosargus cingulatus mate near their oviposition site of decomposing vegetation and Inopus rubiceps engages in scramble competition.
“However, more distantly related soldier flies do not lek: Merosargus cingulatus , which mate near their oviposition site of decomposing vegetation (Barbosa 2011), and Inopus rubiceps (Macquart) (Diptera: Stratiomyidae) which engages in scramble competition (Alcock 1990).”
Quote in the paperNot a substantial claim1 of 2 sources checkedMay combine two claims.
Findingcitation§ 2.3.1 | Oviposition · p. 8Following copulation, female BSF typically begin laying eggs between 4 and 6 days of their introduction into the breeding environment, though the pre-oviposition period can be greatly lengthened at temperatures below 30°C or shortened with increasing temperatures up to BSF's maximum threshold.
“Following copulation, female BSF typically begin laying eggs between 4 and 6 days of their introduction into the breeding environment, though the pre- oviposition period can be greatly lengthened at temperatures below 30°C or shortened with increasing temperatures up to BSF's maximum threshold (Chia et al. 2018).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Chia et al. 2018 source
Split into 2 atoms — 1 confirmed, 1 partially supported, 0 disputed, 0 misattributed, 0 unverifiable. Expand the claim to see per-atom verdicts.
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8Delaying mating is suspected to cause females to haphazardly lay eggs far away from the larval substrate, as can environmental stress.
“Likewise, delaying mating (e.g., as an artifact of experimental set- up) is suspected to cause females to haphazardly lay eggs when females lay single (or additional) clutches (Dickerson et al. 2024; Muraro et al. 2024; Lemke et al. 2024) far away from the larval substrate, as can environmental stress (Lemke, Li, Dickerson, et al. 2025).”
Quote in the paperNot a substantial claim1 of 3 sources checked- cites Dickerson et al. 2024 source
- cites Muraro et al. 2024 source
- cites Lemke et al. 2024 source
The Discussion (§4.3; page unavailable) speculates that delayed mating might explain higher female mortality, not where females lay eggs. The Methods (§2.5; page unavailable) describe clutches collected in egg traps but do not link delayed mating to haphazard laying or laying away from larval substrate. I searched for delayed mating, additional clutches, and oviposition away from substrate; the environmental-stress clause is cited to a different paper. (LLM, conf=0.75)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8For BSF in captivity, the majority of matings occur on the first or second day after being introduced to mating cages, but continue at a low rate after this, reaching a plateau between days 4 and 6.
“For BSF in captivity, the majority of matings occur on the first or second day after being introduced to mating cages, but continue to occur at a low rate after this, reaching a plateau between days 4 and 6 (Tomberlin et al. 2002; Dickerson et al. 2024; Lemke et al. 2024).”
Quote in the paperNot a substantial claim1 of 3 sources checked- cites Tomberlin et al. 2002
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Dickerson et al. 2024 source
- cites Lemke et al. 2024 source
Methods §2.5 reports that mating events were counted for 6–7 days and that their distribution plateaued after that period. Discussion §4.3 describes mating intensity as generally peaking by the third day, but the paper does not establish that most matings occurred on days 1–2 or that mating reached a plateau between days 4 and 6. Page numbers were not available in our copy. (LLM, conf=0.83)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ 2.3.1 | Oviposition · p. 8Oviposition timing varies based on biological age and mating experience.
“This timing varies based on biological age (Dickerson et al. 2024) and mating experience (Permana et al. 2020).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites Dickerson et al. 2024 source
- cites Permana et al. 2020 source
Permana et al. are cited for the mating-experience part, not the biological-age part. The Abstract (page unavailable) reports that mates’ virginity significantly influenced oviposition frequency, but it does not establish that the timing of oviposition changed. (LLM, conf=0.76)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ BOX 4 | Reproductive skews in BSF. · p. 7In equal sex ratio populations of 500 male and 500 female BSF, only 43-120 mating events were observed over a week, indicating that a minority of males likely secure the bulk of matings.
“In fact, in equal sex ratio populations of 500 male and 500 female BSF, only 43-120 mating events were observed over a week, indicating that a minority of males likely secure the bulk of matings (Meyermans et al. 2025).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Meyermans et al. 2025 source
BSF rearing (p. 3) confirms that each tent held 500 males and 500 females, with matings recorded during eight days of periodic observations. Reproduction parameters (p. 4) reports 43 matings for one strain, but also reports counts up to 200, so 43–120 does not describe the full range. The paper does not establish which males mated or whether a minority secured most matings. (LLM, conf=0.88)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ BOX 6 | (Continued) · p. 9Seminal components transferred by males to females may contribute to female longevity, potentially acting as nuptial gifts for females to digest by supplying amino acids and/or lipids that may support egg development and/or metabolic function.
“Interestingly, the seminal components transferred by males to females may contribute to female longevity, potentially acting as nuptial gifts (Harjoko et al. 2023) for females to digest (Manas et al. 2024) by supplying amino acids and/or lipids that may support egg development and/or metabolic function.”
Quote in the paperNot a substantial claim1 of 2 sources checkedMay combine two claims.
Findingcitation§ BOX 4 | Reproductive skews in BSF. · p. 7Mating success in BSF can be dependent on biotic factors (sex ratio, density) and abiotic factors (light, temperature, stressors), and mating frequencies can vary dramatically across populations and colonies.
“It can be dependent on both biotic-e.g., sex ratio, density (Hoc et al. 2019)-and abiotic factors (e.g. light, temperature, stressors (Dearlove et al. 2025)) and experimental studies suggest that mating frequencies can vary dramatically across populations and colonies (Jones and Tomberlin 2021; Dickerson et al. 2024; Lemke et al. 2024; Lemke, Li, Dickerson, et”
Quote in the paperNot a substantial claim3 of 3 sources checked- cites Hoc et al. 2019 source
- cites Dearlove et al. 2025 source
The section “Stressors in mass-reared insects” identifies temperature and light as environmental conditions to standardize and discusses how stressors can affect insect health. It does not establish that these factors affect mating success in black soldier flies. The biotic factors and variation in mating frequency are attributed to other citations in the citing sentence. (LLM, conf=0.76)
Experimental — accuracy not yet measured - cites Jones and Tomberlin 2021 source
May combine two claims.
Findingcitation§ BOX 4 | Reproductive skews in BSF. · p. 7In much smaller groups of 15 male and 15 female BSF, nearly half of the males never mated; half of the successful males mated repeatedly, producing a pronounced reproductive skew.
“In much smaller groups of 15 male and 15 female BSF, nearly half of the males never mated. Half of the successful males mated repeatedly, producing a pronounced reproductive skew (Manas, Venon, et al. 2025).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Manas, Venon, et al. 2025 source
May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8Fecundity and fertility can trade-off at industrial scale, and larval nutrition can induce phenotypic plasticity in both female fecundity and fertility.
“Fecundity and fertility (i.e., the proportion of larvae that hatch) can trade- off at industrial scale (Hoc et al. 2019) and recent study suggests that larval nutrition can induce phenotypic plasticity in both female fecundity as well as fertility (Zhang and Puniamoorthy 2025) (see Box 6).”
Quote in the paperNot a substantial claim2 of 2 sources checked- cites Hoc et al. 2019 source
In Results > Density and sex, female-dominant cages produced more egg mass at higher densities but had a lower hatching rate (93.5% versus 96.5%), supporting a trade-off in the tested cages. The paper does not establish that this trade-off occurs at industrial scale. The larval-nutrition claim is attributed to Zhang and Puniamoorthy (2025), not to this source. (LLM, conf=0.83)
Experimental — accuracy not yet measured - cites Zhang and Puniamoorthy 2025 source
May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8The temporal pattern of mating appears generally consistent across experiments and is highly correlated (98%) to oviposition events 2-3 days later.
“Importantly, recent work has shown this temporal pattern appears to generally be consistent across experiments and are highly correlated (98%) to oviposition events 2-3 days later (Lemke, Li, and De Smet 2025).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke, Li, and De Smet 2025 source
The Cross Correlation of Time Series Data section (page unavailable) reports a 97.54% average percent correlation between mating and oviposition in traps at a two-day lag, reasonably rounded to 98%. But oviposition outside traps reached only about 72% at a three-day lag. The analysis used two trials from one experiment, so it does not establish that the 98% relationship is consistent across experiments. (LLM, conf=0.86)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ 2.2.2 | Sperm Storage · p. 7The total number of stored sperm can range from a few hundred to several thousand, and it may take about 48 h for approximately 50% of transferred sperm to reach the sperm storage reservoirs.
“The total number of stored sperm can range from a few hundred to several thousand (Manas et al. 2024) and it may take about 48 h for approximately 50% of transferred sperm to reach the sperm storage reservoirs.”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Manas et al. 2024 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingcitation§ 2.3.2 | Aging and Senescence · p. 8These translucent windows are thought to have evolved to mimic the petiole of Polistes paper wasps.
“These translucent windows are thought to have evolved to mimic the petiole of Polistes paper wasps (James 1935; Alcock 1990).”
Quote in the paperNot a substantial claim1 of 2 sources checked- cites James 1935
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Alcock 1990 source
The Introduction (page not supplied) says Hermetia comstocki appears to be a Batesian mimic of Polistes paper wasps. It does not say that translucent windows mimic the wasps’ petiole or that those windows evolved for that purpose; searches for those features found no supporting passage. (LLM, conf=0.85)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ 2.2.2 | Sperm Storage · p. 7A single ejaculate is sufficient to fill a female's sperm reservoirs in excess, challenging the previous hypothesis that females were sperm limited; a single mating can support multiple egg clutches.
“Recent work demonstrated that a single ejaculate is sufficient to fill a female's sperm reservoirs in excess, which challenges the previous hypothesis that females were sperm limited (Permana et al. 2020). Instead, a single mating can support multiple egg clutches.”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Permana et al. 2020 source
The Abstract describes an experiment on mate virginity, remating, and oviposition frequency, but does not report ejaculate volume or sperm-reservoir capacity. Searches for sperm limitation and multiple egg clutches found no passage establishing that one mating supplies enough sperm for multiple clutches. (LLM, conf=0.70)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 10Intentional selection on larval feed conversion rates, adult body weights and fecundity can yield rapid responses but increase the risk of inbreeding depression, especially as a consequence of unintentional artificial selection.
“Intentional selection on larval feed conversion rates, adult body weights and fecundity can yield rapid responses but increase the risk of inbreeding depression (Meyermans et al. 2025), especially as a consequence of unintentional artificial selection (e.g., for BSF that perform well in captivity, such as those that survive abiotic shocks, those that oviposit in traps and those”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Meyermans et al. 2025 source
The Introduction (p. 2) reports that long-term selection increased BSF larval weight, supporting the general potential for a response to selection. It does not establish rapid responses from selection on larval feed conversion, adult body weight, and fecundity, or say that such selection increases inbreeding-depression risk. The unintentional-selection explanation follows the citation in the citing sentence. (LLM, conf=0.82)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10The degree to which sperm precedence follows 'last male wins' patterns in BSF is unknown, nor how this pattern may break down with high amounts of remating or as flies senesce.
“However, the degree to which sperm precedence follows 'last male wins' patterns, as seen in other Diptera, is unknown, nor how this pattern may subsequently break down with high amounts of remating (Zeh and Zeh 1997; Laturney et al. 2018) or as flies senesce (Mack et al. 2003).”
Quote in the paperNot a substantial claim1 of 3 sources checkedMay combine two claims.
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Sperm length in BSF is unusually long for internally fertilizing animals and is sensitive to larval diet quality, suggesting potential trade-offs between sperm quality and quantity that could shape outcomes among sperm competing within the female genital tract and sperm storage organs.
“Moreover, as mentioned, sperm length in BSF (Malawey et al. 2019, 2020) is unusually long for internally fertilizing animals and is sensitive to larval diet quality (Zhang and Puniamoorthy 2025), suggesting potential trade- offs between sperm quality and quantity that could shape outcomes among sperm competing within the genital tract of the female and her sperm storage organs.”
Quote in the paperNot a substantial claim1 of 3 sources checkedClassified as probably not a substantial claim. May combine two claims.
Interpretationreasoning§ 3 | Implications for Evolutionary Biology · p. 10This raises the possibility of BSF actually being a cryptic species complex and provides an avenue to examine potential mechanisms governing reproductive isolation under domestication.
“This raises the possibility of BSF actually being a cryptic species complex (Ståhls et al. 2020; Generalovic et al. 2023; Athanassiou et al. 2024) and provides an avenue to examine potential mechanisms governing reproductive isolation under domestication.”
Quote in the paperSubstantial claim1 of 3 sources checked- cites Ståhls et al. 2020 source
- cites Generalovic et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Athanassiou et al. 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 10Recent genomic surveys indicate substantial diversity among global BSF populations, including rapid genomic differentiation between strains under artificial selection.
“Recent genomic surveys indicate substantial diversity among global BSF populations (Ståhls et al. 2020), including rapid genomic differentiation between strains under artificial selection (Silvaraju et al. 2026).”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Ståhls et al. 2020 source
The Discussion (‘High COI barcode diversity’) reports very high mitochondrial COI variability in BSF sampled across regions, supporting the diversity portion attributed to Ståhls et al. It also says that this variability was not mirrored by the two nuclear markers studied, so describing the finding as one from genomic surveys overstates what this paper establishes. The artificial-selection clause is cited to a different source and is not part of this verdict. (LLM, conf=0.84)
Experimental — accuracy not yet measured - cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 10Phenotypic plasticity driven by larval diet influences key reproductive traits in BSF, including adult body size, fecundity and sperm length, and interacts with heritable variation to shape evolutionary potential.
“Phenotypic plasticity (West- Eberhard 1989) driven by larval diet influences key reproductive traits in BSF, including adult body size, fecundity and sperm length, that also interacts with heritable variation to shape evolutionary potential (Gobbi et al. 2013; Shrestha et al. 2025; Zhang and Puniamoorthy 2025).”
Quote in the paperSubstantial claim1 of 3 sources checked- cites West-Eberhard 1989 source
- cites Gobbi et al. 2013 source
The Discussion reports variation in BSF body size and fecundity among flies reared on different larval diets. It does not report effects on sperm length or test whether diet-driven plasticity interacts with heritable variation to shape evolutionary potential. Its general observation that wing size can reflect genetic and environmental variables does not establish that interaction. (LLM, conf=0.88)
Experimental — accuracy not yet measured - cites Shrestha et al. 2025 source
May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 9In BSF, recent studies document polygynandry, steep reproductive skews, extended sperm storage and possible cryptic female choice, including ejaculate digestion and selective sperm retention.
“In BSF, recent studies document polygynandry, steep reproductive skews, extended sperm storage and possible cryptic female choice, including ejaculate digestion and selective sperm retention (Muraro et al. 2024; Manas et al. 2024; Manas, Labrousse, and Bressac 2025).”
Quote in the paperSubstantial claim1 of 3 sources checked- cites Muraro et al. 2024 source
- cites Manas et al. 2024 source
The Introduction reports a polygynandrous mating system in BSF, and the Discussion reports prolonged sperm storage and evidence suggesting possible sperm digestion. The paper does not document steep reproductive skews or selective sperm retention in BSF; its discussion of sperm-based cryptic female choice describes mechanisms in other species, not a demonstrated BSF finding. No page numbers are provided in our copy. (LLM, conf=0.87)
Experimental — accuracy not yet measured - cites Manas, Labrousse, and Bressac 2025 source
May combine two claims.
Premisecitation§ 4.1 | Genetics and Population Structure · p. 11Continuous production makes it difficult to identify and control for variation and perpetuates variation because eggs are fertilized by a heterogeneous group of flies.
“it has the significant drawbacks of making it difficult to precisely identify and control for variation that accrues throughout various steps in the industrial production cycle, as well as perpetuating this variation because any collected eggs will be fertilized and laid by a heterogenous group of flies (Lemke, Li, Dickerson, et al. 2025).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Lemke, Li, Dickerson, et al. 2025 source
The Abstract (page unavailable) says introducing flies before older flies are removed creates a mixed-age breeding population, and it reports differences in egg yield and fertility between mixed- and same-age cohorts. It does not say that continuous industrial production makes variation across production steps difficult to identify or control, or that eggs perpetuate that variation. (LLM, conf=0.82)
Experimental — accuracy not yet measured
May combine two claims.
Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11BSF of multiple size-classes successfully mate, although with dramatically increased variability.
“Indeed BSF of multiple size- classes successfully mate, although with dramatically increased variability (Jones and Tomberlin 2021).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Jones and Tomberlin 2021 source
The Mating observations section reports successful pairings between large and small black soldier flies, with different numbers of pairs across size combinations. The Oviposition and hatch rate section reports dramatically greater variability specifically in egg hatch rate in heterogeneous populations; it does not establish dramatically increased variability in mating success, as the citing sentence could imply. (LLM, conf=0.86)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ 4.1 | Genetics and Population Structure · p. 11Another study shows increased production traits as a result of inbreeding in BSF.
“another shows the opposite, with increased production traits as a result of inbreeding (Cai et al. 2022).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Cai et al. 2022 source
The Abstract reports that overall egg production in the selectively inbred BSF strain was 1.5 times that of the Wuhan strain. But the Discussion (§4.3) says the mechanism remains unclear, and the Conclusions say the advantage *may* reflect inbreeding selection. The claim presents inbreeding as an established cause and describes the improvement more broadly than the measured egg-production result. (LLM, conf=0.83)
Experimental — accuracy not yet measured
May combine two claims.
Premisecitation§ 4.2 | Nutritional Management · p. 12Adult BSF access to water and nutrient supplements can extend lifespan, shorten pre-oviposition intervals, and increase egg production under several conditions.
“Adult BSF's access to water and nutrient supplements (e.g., honey, pollen, or sugar water) can extend lifespan, shorten pre- oviposition intervals and increase egg production under several conditions (Thinn and Kainoh 2022; Klüber et al. 2023; Barrett et al. 2025).”
Quote in the paperSubstantial claim2 of 3 sources checked- cites Thinn and Kainoh 2022 source
The Results (§§1–2, page unavailable) show that adults given sugar-, pollen-, or honey-containing diets lived longer and produced more eggs than adults given water alone. The paper reports oviposition duration and peak egg-laying days, but not a shortened pre-oviposition interval. It also does not experimentally compare access to water with no water. (LLM, conf=0.88)
Experimental — accuracy not yet measured - cites Klüber et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Barrett et al. 2025 source
The Abstract reports that adults given a preferred yeast–molasses diet rather than water alone lived 4–6 days longer and laid more eggs on days 2–5 after eclosion. It does not report a shorter pre-oviposition interval, and its tested supplement was not honey, pollen, or sugar water. (LLM, conf=0.86)
Experimental — accuracy not yet measured
May combine two claims.
Interpretationcitation§ 4.3 | Sex Ratios and Habitat Design · p. 12Providing perches may allow BSF, especially females, to conserve energy and reduce negative effects of crowding.
“Providing perches may allow BSF (and especially females) to conserve energy and help to reduce any negative effects of crowding (Lemke et al. 2024).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke et al. 2024 source
May combine two claims.
Interpretationcitation§ 4.3 | Sex Ratios and Habitat Design · p. 12Spatial separation of mating and oviposition zones, provisioning of microclimates, and control of light gradients can enhance reproductive success in captive BSF.
“suggest that spatial separation of the mating and oviposition zones within cages, proper provisioning of microclimates and a precise control of light gradients can enhance reproductive success (Lemke, Li, Dickerson, et al. 2025).”
Quote in the paperSubstantial claim1 of 1 sources checked- cites Lemke, Li, Dickerson, et al. 2025 source
The Abstract reports that oviposition-attractant timing affected egg yield and hypothesizes that limited spatial segregation in small cages may reduce mate quality. It does not say that separating mating and oviposition zones, provisioning microclimates, or controlling light gradients enhances reproductive success; its spatial-segregation hypothesis concerns mixed-age flies, not distinct activity zones. (LLM, conf=0.77)
Experimental — accuracy not yet measured
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Interpretationcitation§ 4.3 | Sex Ratios and Habitat Design · p. 12There is an optimum plant density for BSF mating, with too little not beneficial and too much potentially hindering mating.
“Theoretically there should be an optimum: too little is not beneficial, but too much may be a physical or visual hinderance to mating (Lemke et al. 2024).”
Quote in the paperNot a substantial claim1 of 1 sources checked- cites Lemke et al. 2024 source
May combine two claims.
Premisecitation§ 4.4 | Chemical Ecology and Oviposition Control · p. 12BSF symbiotic microbes release VOCs that are attractive to mature females and drive detection and oviposition behaviour.
“It is these microbes (i.e., colonizing the surface of eggs and larvae, as well as living in the gut and substrates of the larval media) (Heussler et al. 2023) which release VOCs that have evolved to be attractive to mature females, driving detection and oviposition behaviour (Zheng et al. 2013).”
Quote in the paperSubstantial claim1 of 2 sources checked- cites Heussler et al. 2023 source
Checked before — its earlier verdict is held back until a new check runs.
- cites Zheng et al. 2013 source
The Discussion (page not provided) says bacteria isolated from conspecifics on decomposing material attract gravid black soldier flies, presumably through emitted volatiles; the Abstract reports effects on oviposition preference. The paper does not establish that the bacteria release the proposed VOCs or that those VOCs evolved to attract females. Its evolutionary statement concerns flies evolving to detect resources, not microbes evolving to attract flies. (LLM, conf=0.88)
Experimental — accuracy not yet measured
May combine two claims.
Findingcitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6In 111 other insects and arthropods, same-sex sexual (SSS) behaviour is more common between males than females and is highly associated with captivity, high-rearing densities and the presence of female pheromones.
“Interestingly, the examination of same- sex sexual (SSS) behaviours in 111 other insects and arthropods revealed that SSS is more common between males than females and is highly associated with captivity, high- rearing densities and the presence of female pheromones (Scharf and Martin 2013).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Scharf and Martin 2013 source
May combine two claims.
Findingcitation§ BOX 3 | (Continued) · p. 7The relationship between duration of wing buzzing and mating success in BSF is inconsistent across studies, with some reporting a positive correlation, some the opposite trend, and some no correlation at all.
“Some experiments suggest a positive correlation between duration of wing buzzing and mating success (Giunti et al. 2018), others have reporting the opposite trend (Laudani et al. 2024) or no correlation at all (Kortsmit et al. 2025).”
Quote in the paperNot a substantial claim0 of 3 sources checkedMay combine two claims.
Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7Sperm transfer is not a continuous process; sperm is delivered only in the lattermost portion of copulation (e.g., after ~25 min), following the transmission of seminal fluids.
“Crucially, sperm transfer is not a continuous process. Dissection of females at different times throughout mating suggests that sperm is delivered only in the lattermost portion of copulation (e.g., after ~25 min), following the transmission of seminal fluids (Manas, Labrousse, and Bressac 2025).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Manas, Labrousse, and Bressac 2025 source
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7After copulation, both sexes perform grooming behaviours, with females typically grooming for a longer period than males (~300 versus 100 s).
“Copulation then ceases when both sexes disengage, after which both sexes perform grooming behaviours, with females typically grooming for a longer period than males (~300 versus 100 s) (Giunti et al. 2018; Laudani et al. 2024) (Box 4).”
Quote in the paperNot a substantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ BOX 3 | Debates on lekking in BSF. · p. 6There exists debate as to whether lekking as a whole is preserved in captivity and whether large aggregations are a prerequisite for mating.
“Overall, there exists debate as to whether lekking as a whole is preserved in captivity (Lemke et al. 2023) and whether large aggregations are a prerequisite for mating (Laudani et al. 2024).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Lemke et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Laudani et al. 2024 source
May combine two claims.
Findingcitation§ 2.2.1 | Copulatory Mechanics, Duration and Sperm Transfer · p. 7Overall mating duration is prolonged in sibling crosses and amongst individuals that experienced poor larval nutrition.
“Additionally, overall mating duration is prolonged in sibling crosses (Laudani et al. 2024) and amongst individuals that experienced poor larval nutrition (Zhang, Ng, et al. 2025).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Laudani et al. 2024 source
- cites Zhang, Ng, et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ 2.3.2 | Aging and Senescence · p. 8Observing colour changes in the abdominal windows can serve as a non-invasive indicator of biological aging and has been successfully adopted for population management in industrial settings.
“Observing colour changes in the abdominal windows can serve as a non- invasive indicator of biological aging and has been successfully adopted for population management in industrial settings (Salari and De Goede 2024).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Salari and De Goede 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8Examination of egg clutches from breeding cage experiments has revealed that both fertility and neonate viability can vary substantially, with portions of eggs not developing, developing eyespots but not hatching, hatching but not surviving, or hatching and surviving and reproducing.
“Examination of egg clutches collected from breeding cage experiments has revealed that both fertility and neonate viability can vary in substantial ways, with a portion of eggs (from multiple clutches) not developing at all, a portion developing eyespots but otherwise not hatching, a portion hatching but not surviving, as well as a portion that hatches, survives and successfully reproduces (A.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites A. Dickerson, unpublished data
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8Male BSF have been observed to mate up to four, five or six times, and females up to twice, but higher female polyandry up to five times has been supported by parentage assignments.
“Behavioural assays have observed male BSF to mate up to four (Chiabotto et al. 2024; Laudani et al. 2024), five (Jensen et al. 2025) or six times (Manas, Venon, et al. 2025) and females up to twice (Chiabotto et al. 2024) but higher female polyandry, up to five times, has been supported by parentage assignments (Hoffmann et al. 2021).”
Quote in the paperNot a substantial claim0 of 3 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Findingnone given§ 2.3.2 | Aging and Senescence · p. 8Observations from both laboratory and industrial settings report older BSF frequently crashing to the cage floor during flight or spinning erratically after having broken one of their wings, which can explain some of their injuries.
“Observations from both laboratory and industrial settings report older BSF frequently crashing to the cage floor during flight or even spinning erratically after having broken one of their wings, which can explain some of their injuries.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingcitation§ BOX 6 | (Continued) · p. 9Older males exhibit reduced mating frequency and fertility due to declining sperm viability.
“Older males exhibit reduced mating frequency (Dickerson et al. 2024) and fertility due to declining sperm viability (Malawey et al. 2020).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Dickerson et al. 2024 source
- cites Malawey et al. 2020
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ BOX 5 | Re- mating in BSF. · p. 8Off-target egg laying can quickly snowball because females are attracted to VOCs released by eggs, which might then signal to additional females to lay their eggs in off-target locations.
“Off- target egg laying can quickly snowball because females are attracted to VOCs released by eggs (Klüber et al. 2024), which might then signal to additional females to lay their eggs in off- target locations.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Klüber et al. 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 9BSF is a (hybrid) capital breeder; adult reproduction relies almost entirely on larval-derived reserves, decoupling gametogenesis and courtship investment from adult foraging.
“As mentioned, being a (hybrid) capital breeder (Stephens et al. 2009), adult reproduction relies almost entirely on larval- derived reserves, thereby decoupling gametogenesis and courtship investment from adult foraging.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Stephens et al. 2009 source
May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 10Laboratory studies show BSF can adapt within generations to suboptimal diets, supporting their use in experimental evolution.
“Laboratory studies show BSF can adapt within generations to suboptimal diets (Jiggins 2024), supporting their use in experimental evolution.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Jiggins 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Findingcitation§ 3 | Implications for Evolutionary Biology · p. 9Comparative studies show how sperm traits evolve under such competition: Drosophila bifurca produces the longest sperm in the animal kingdom, whereas lepidopterans package sperm into nutrient-rich spermatophores.
“Comparative studies show how sperm traits evolve under such competition: Drosophila bifurca produces the longest sperm in the animal kingdom, whereas lepidopterans package sperm into nutrient- rich spermatophores (Pitnick et al. 1995; Vahed 1998).”
Quote in the paperNot a substantial claim0 of 2 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Insects such as Drosophila, crickets and butterflies can bias sperm use through differential storage or ejection of sperm from less-preferred males.
“For instance, insects such as Drosophila , crickets and butterflies can bias sperm use through differential storage or ejection of sperm from less- preferred males (Eberhard 1991).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Eberhard 1991 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10In insects, dozens of mechanisms of cryptic female choice have been uncovered.
“In insects, dozens of such mechanisms have been uncovered (Shuker and Simmons 2014).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Shuker and Simmons 2014
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10BSF males can mate up to 4-5 times, whereas females might only remate twice.
“BSF males can mate up to 4-5 times (Jensen et al. 2025), whereas females might only remate twice (Chiabotto et al. 2024).”
Quote in the paperNot a substantial claim0 of 2 sources checkedMay combine two claims.
Findingnone given§ BOX 6 | (Continued) · p. 9Female clutch size and egg viability decline with age, and egg collections from mixed age cohorts will exhibit greater variability in survival and overall fitness.
“Similarly, female clutch size and egg viability decline with age and egg collections from mixed age cohorts will undoubtedly exhibit a greater variability in survival and overall fitness.”
Quote in the paperNot a substantial claimMay combine two claims.
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Males may transfer seminal fluid proteins (e.g., sex peptides) that influence female re-mating latency or stimulate reproductive development.
“In addition, males may transfer seminal fluid proteins (e.g., sex peptides) that influence female re- mating latency or stimulate reproductive development (Kubli 1992).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Kubli 1992 source
May combine two claims.
Findingcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Mating in BSF has been shown to reduce male longevity whilst increasing females', suggesting a potential trade-off between mating frequency and fitness for each sex.
“But conversely, mating in BSF has been shown to reduce male longevity whilst increasing females', suggesting a potential trade- off between mating frequency and fitness for each sex (Harjoko et al. 2023).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Harjoko et al. 2023 source
May combine two claims.
Premisecitation§ WILEY · p. 2Adult foraging can extend lifespan and enhance reproductive output in BSF.
“Adult foraging can nevertheless extend lifespan and enhance reproductive output (Zhang, Ng, et al. 2025; Kortsmit et al. 2025).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Zhang, Ng, et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Kortsmit et al. 2025 source
May combine two claims.
Premisecitation§ WILEY · p. 2The timing of BSF emergence is sensitive to larval nutrition and is linked with size dimorphism.
“though the timing of emergence is sensitive to larval nutrition (Zhang, Ng, et al. 2025) and is linked with size dimorphism (Generalovic et al. 2025).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Zhang, Ng, et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Generalovic et al. 2025 source
May combine two claims.
Premisecitation§ WILEY · p. 2BSF adults do not require feeding for gametogenesis; successful adult reproduction relies almost entirely on fat body reserves accumulated during the larval stage.
“By contrast, BSF adults do not require feeding for gametogenesis. Instead, successful adult reproduction relies almost entirely on fat body reserves accumulated during the larval stage occur (Tomberlin and Sheppard 2002).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Tomberlin and Sheppard 2002
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4The duration of sexual bimaturism in BSF is modulated by larval nutrition and interacts with body size and longevity; larvae that take longer to develop become larger adults.
“The duration of this developmental gap between male and female emergence, also known as sexual bimaturism (SBM), is modulated by larval nutrition and interacts with body size and longevity (Generalovic et al. 2025); larvae that take longer to develop become larger adults.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Generalovic et al. 2025 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6Acoustic signals may play a role in sex discrimination in BSF once mounted.
“Recent work in BSF suggests that acoustic signals may play a role in sex discrimination once mounted”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 2 | Debates on mate discrimination in BSF. · p. 6BSF lack sexual dimorphism in their cuticular hydrocarbon (CHC) profiles.
“Supporting evidence for this hypothesis comes from the lack of sexual dimorphism in their cuticular hydrocarbon (CHC) profiles (Lemke et al. 2023)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Lemke et al. 2023
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Interpretationreasoning§ WILEY · p. 2Because adult nutrition cannot compensate for the nutritional history experienced as larvae, any variation in larval diet is directly reflected in adult reproductive performance, especially in captive or industrial systems where many producers do not feed adults.
“Because adult nutrition cannot compensate for the nutritional history experienced as larvae, any variation in larval diet is directly reflected in adult reproductive performance, especially in captive or industrial systems where many producers do not feed adults.”
Quote in the paperNot a substantial claimMay combine two claims.
Premisereasoning§ BOX 1 | What makes BSF a model organism in the classic sense? · p. 3In captivity, BSF generation time is generally between 6 and 7 weeks, meaning 7 generations can occur per year.
“The generation time is thus generally between 6 and 7 weeks, meaning 7 generations can occur per year”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationreasoning§ WILEY · p. 3BSF are an ideal system to address questions about nutritional legacies, abiotic conditions, mating system evolution under domestication, pre- and post-copulatory sexual selection, and reproductive barriers.
“Specifically, we believe that BSF are an ideal system to address:”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4Protandry is thought to allow males to monopolise access by being present and ready to mate as soon as females become receptive, thereby minimizing the time females spend unmated.
“It is thought to allow males to monopolise access by being present and ready to mate as soon as females become receptive, thereby minimizing the time females spend unmated (Zhang, Henawy, et al. 2025; Fagerström and Wiklund 1982).”
Quote in the paperNot a substantial claim0 of 2 sources checked- cites Zhang, Henawy, et al. 2025
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Fagerström and Wiklund 1982 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisenone given§ BOX 1 | What makes BSF a model organism in the classic sense? · p. 3Female BSF carry between 300 and 900 eggs.
“Females carry between 300 and 900 eggs”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Interpretationcitation§ 2.1.2 | Navigation and Lekking · p. 5In an industrial setting, dedicated mating arenas and oviposition areas within the cage could enhance reproductive success.
“This suggests that in an industrial setting, dedicated mating arenas and oviposition areas within the cage could enhance reproductive success (Salari and De Goede 2024)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Salari and De Goede 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Interpretationreasoning§ WILEY · p. 2The wide global distribution, pronounced larval nutritional flexibility, rapid genomic divergence and high industrial relevance of BSF together provide a powerful system for investigating phenotypic plasticity and the mechanisms between selection, genetic drift and reproductive isolation.
“Altogether, the wide global distribution, pronounced larval nutritional flexibility, rapid genomic divergence and high industrial relevance of BSF together provide a powerful system for investigating phenotypic plasticity and the mechanisms between selection, genetic drift and reproductive isolation.”
Quote in the paperSubstantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ 2.1.1 | Latency and Sexual Maturation · p. 4Protandry occurs in ~36% of investigated insect species.
“This strategy is common in butterflies and other insects, occurring in ~36% of investigated insect species (Teder et al. 2021).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Teder et al. 2021 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Interpretationcitation§ 2.1.3 | Courtship and Mounting · p. 5Temporal patterns in BSF perching and flight activity mirror that of other diurnal lekking insects.
“Such temporal patterns in perching and flight activity mirror that of other diurnal lekking insects (Michiels and Dhondt 1989)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Michiels and Dhondt 1989
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 3 | Debates on lekking in BSF. · p. 6Lekking itself is no longer viewed within a strict definition.
“However, lekking itself is no longer viewed within a strict definition (Alcock 1987).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Alcock 1987 source
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation§ BOX 3 | Debates on lekking in BSF. · p. 6When females become rare, the mating system can shift to either resource-defence polygyny or female-defence polygyny once males start to monopolise female resources or the females themselves.
“Conversely, when females become rare, the mating system can shift to either resource- defence polygyny or female- defence polygyny once males start to monopolise female resources or the females themselves (Buzatto and Machado 2008), respectively.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Buzatto and Machado 2008 source
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Premisecitation§ BOX 3 | Debates on lekking in BSF. · p. 6The lek mating system can be modelled along a gradient of increasing resource dependence and mate monopolization and can shift dynamically if underlying factors change.
“Moreover, the lek mating system itself also can be modelled along a gradient of increasing resource dependence and mate monopolization (Emlen and Oring 1977; Parker 1978; Thornhill and Alcock 1983) and can shift dynamically if these underlying factors change.”
Quote in the paperNot a substantial claim0 of 3 sources checkedProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ BOX 3 | Debates on lekking in BSF. · p. 6Each criterion of lekking can be considered as an independent continuum in multidimensional parameter space.
“Each of these criteria can be considered as an independent continuum in multidimensional parameter space similar to the Hutchinsonian niche (Hutchinson 1957), enabling a more holistic view of lekking systems.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Hutchinson 1957 source
Probably not a substantial claim: may not pass S3 (load-bearing).
Interpretationcitation§ BOX 3 | (Continued) · p. 7Mating 'songs' might need to be performed with the proper rhythm and duration to be accepted by prospective mates.
“Besides pitch and tone, mating 'songs' might need to be performed with the proper rhythm (Eberhard and Gelhaus 2009) and duration to be accepted by prospective mates.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Eberhard and Gelhaus 2009 source
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Interpretationown data§ BOX 3 | (Continued) · p. 7Although some pairs remain mounted from behind, the ability to rotate between the two positions may allow the pair to take flight and relocate in response to disturbance.
“Although some remain mounted from behind (Chiabotto et al. 2024), the ability to rotate between the two positions may allow the pair to take flight and relocate in response to disturbance (pers. obs.).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Chiabotto et al. 2024 source
Probably not a substantial claim: may not pass S3 (load-bearing), S4 (based). May combine two claims.
Interpretationreasoning§ 2.3.2 | Aging and Senescence · p. 8These physical changes likely occur from repeated attempts to mate or oviposit despite declining energy reserves, as well as inadequate humidity and/or water resources in the breeding environment.
“These likely occur from repeated attempts to mate or oviposit despite declining energy reserves, as well as inadequate humidity and/or water resources in the breeding environment.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationreasoning§ BOX 4 | Reproductive skews in BSF. · p. 7This pattern suggests population substructures (driven via non-random mating) and a low effective population size, which are hallmarks of lek-type mating systems in which a single or a few dominant males monopolise fertilizations.
“This pattern suggests population substructures (driven via non- random mating) and a low effective population size, which are hallmarks of lek- type mating systems in which a single or a few dominant males monopolise fertilizations.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Males can differentially adjust their sperm transfer, which is often called cryptic male choice.
“Likewise, males can differentially adjust their sperm transfer, which is often called cryptic male choice (Shuker and Simmons 2014).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Shuker and Simmons 2014
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Interpretationreasoning§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Females may regulate sperm uptake or storage or bias fertilization toward preferred males, but such mechanisms remain largely untested in BSF.
“Females may regulate sperm uptake or storage or bias fertilization toward preferred males, but such mechanisms remain largely untested in BSF.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationcitation§ 3 | Implications for Evolutionary Biology · p. 9This capital breeding reshapes sexual selection dynamics relative to other income-breeding insects and anautogenous blood-feeding Diptera, creating potential trade-offs among larval resource allocation, somatic growth, longevity and reproductive output.
“This reshapes sexual selection dynamics relative to other income- breeding insects (Stephens et al. 2009) and anautogenous blood- feeding Diptera (which require a blood meal to produce eggs), creating potential trade- offs among larval resource allocation, somatic growth, longevity and reproductive output (Miller 2005).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Stephens et al. 2009 source
- cites Miller 2005
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S3 (load-bearing).
Premisecitation§ 3 | Implications for Evolutionary Biology · p. 9Multiple mating by females influences sperm competition among males, a ubiquitous force in insect reproduction.
“Multiple mating by females influences sperm competition among males, a ubiquitous force in insect reproduction (Parker 1970).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Parker 1970 source
May combine two claims.
Interpretationcitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Male harassment to secure extra matings may interfere with female oviposition or induce injury, such that increased exposure to males may be negatively associated with female lifetime fitness.
“Male harassment to secure these extra matings may interfere with female oviposition or even induce injury (Pitnick and García-González 2002), such that increased exposure to males may be negatively associated with female lifetime fitness (Fowler and Partridge 1989; Morrow and Gage 2001).”
Quote in the paperNot a substantial claim0 of 3 sources checkedMay combine two claims.
Premisecitation§ 3.1 | Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10Often, the number of matings that optimizes fitness for males is higher than it is for females, driving such conflicts.
“Often, the number of matings that optimizes fitness for males is higher than it is for females (Wigby and Chapman 2005), driving such conflicts as additional matings for females may be less advantageous for their fitness, but obtaining too few matings will be sub- optimal for male fitness.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Wigby and Chapman 2005 source
May combine two claims.
Interpretationcitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11Male BSF genitalia hypo-allometry allows males to mate with females across a wide range of body sizes and increases their pool of available mates.
“allowing males to mate with females across a wide range of body sizes and increase their pool of available mates (Eberhard et al. 1998)”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Eberhard et al. 1998 source
May combine two claims.
Premisecitation§ 4.1 | Genetics and Population Structure · p. 11Domesticated and selectively bred BSF lines can exhibit markedly reduced heterozygosity and form distinct population clusters.
“recent work demonstrates that domesticated and selectively bred lines can exhibit markedly reduced heterozygosity and form distinct population clusters, highlighting the genetic consequences of intensive breeding programmes (Silvaraju et al. 2026).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11Selective breeding and prolonged captive rearing can rapidly erode genetic diversity and shrink effective population size in BSF.
“The genomic data from Silvaraju et al. (2026) further reveal that selective breeding and prolonged captive rearing can rapidly erode genetic diversity and shrink effective population size”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Silvaraju et al. 2026
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
May combine two claims.
Premisecitation§ 3.2 | Reproductive Barriers and Potential Drivers of Speciation · p. 11Mating experiments between Singapore wild-caught and Spanish commercial BSF strains produced successful hybrids, indicating reproductive incompatibilities were not absolute between those strains.
“An earlier study noted that mating experiments between Singapore wild- caught flies and Spanish commercial strains ( COI distance ~3.7%-4.0%) produced successful hybrids, indicating that reproductive incompatibilities were not yet absolute between those two strains.”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationcitation§ 4.3 | Sex Ratios and Habitat Design · p. 12Cage compartmentalization to physically isolate different BSF cohorts should help maintain proper iOSR and age-structure, increasing fertile egg production.
“different cohorts of flies may be physically isolated through cage compartmentalization to (Salari and De Goede 2024) which ultimately should help to maintain a proper iOSR and age- structure of the breeding zone, increasing fertile egg production.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Salari and De Goede 2024
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim: may not pass S4 (based). May combine two claims.
Interpretationreasoning§ 6 | Conclusions and Future Directions · p. 14BSF is a non-pest, lekking species that is easily culturable, and provides an avenue to study reproductive processes in a wide range of species with similar life history and reproductive traits.
“As a non- pest, lekking species that is easily culturable, meaning BSF provides an avenue to study reproductive processes that are happening in a wide range of species with similar life history and reproductive traits.”
Quote in the paperNot a substantial claimMay combine two claims.
Premisenone given§ 4.4 | Chemical Ecology and Oviposition Control · p. 12Some VOCs, such as decanoic acid, delay oviposition behaviour in BSF, while others increase off-target laying.
“some compounds (e.g., decanoic acid) have been shown to delay oviposition behaviour in BSF, whereas others increase off target laying.”
Quote in the paperSubstantial claim
The authors state this as an assumption.
BSF exist in the wild in a wide range of ecological contexts.
“Yet, BSF exist worldwide (Kaya et al. 2021), in presumably a wide range of ecological contexts.”
The paper relies on this without support.
Once the genital lock commences, rival males cease interruption attempts.
“Once the lock commences, rival males apparently cease interruption attempts.”
The paper relies on this without support.
Industrial BSF production depends on predictable, high-output reproduction.
“Industrial BSF production depends on predictable, high- output reproduction”
The paper relies on this without support.
Adult reproductive biology often remains a major bottleneck in industrial BSF production, especially for maintaining genetic diversity and performance over time.
“yet adult reproductive biology often remains a major bottleneck, especially when it comes to maintaining genetic diversity and performance over time.”
The paper relies on this without support.
Continuous breeding cycles lead to interactions between BSF of mixed ages, sizes, conditions, and reproductive statuses.
“Continuous breeding cycles (whereby new flies are introduced into cages prior to the removal/death of old flies) lead to the interactions (or lack thereof) between BSF of mixed ages, sizes, conditions and reproductive statuses”
The paper relies on this without support.
BSF adult ontogeny begins with protandrous emergence, where earliest males emerge several days before earliest females and establish lek-like aggregations.
“The onset of adult black soldier fly, Hermetia illucens , ontogeny begins with their protandrous emergence, where typically the earliest males emerge several days before the earliest females and establish lek- like aggregations.”
The paper relies on this without support.
Females visit leks, where males initiate aerial courtship displays involving wing-buzzing.
“Females subsequently visit these leks, where males initiate aerial courtship displays involving wing- buzzing.”
The paper relies on this without support.
Successful courtship culminates in a genital lock lasting approximately 33 min, with high variability.
“Successful courtship culminates in a genital lock lasting approximately 33 min, though this duration is highly variable.”
The paper relies on this without support.
Following copulation, intense post-copulatory sexual selection ensues within the female's complex reproductive tract.
“Following copulation, a period of intense post- copulatory sexual selection ensues within the female's complex reproductive tract.”
The paper relies on this without support.
Post-reproductive females become highly responsive to VOCs that guide them to oviposition sites.
“Post reproductive females somehow then become highly responsive to volatile organic compounds that guide them to oviposition sites”
The paper relies on this without support.
Females use gustatory and hygroreceptors on their ovipositor to probe substrates.
“Although females use gustatory and hygroreceptors on their ovipositor to probe substrates”
Cites
Ecology, Sexual Selection, and the Evolution of Mating Systems
Emlen, S. T., & Oring, L. W. (1977) · Science · doi:10.1126/science.327542 · Cited by 6,230 · cited 1× · see citing works
“Moreover, the lek mating system itself also can be modelled along a gradient of increasing resource dependence and mate monopolization (Emlen and Oring 1977; Parker 1978; Thornhill and Alcock 1983) and can shift dynamically if these underlying factors change.”
Hutchinson, G. E. (1957) · Cold Spring Harbor Symposia on Quantitative Biology · doi:10.1101/sqb.1957.022.01.039 · Cited by 6,228 · see citing works
Hebert, P. D. N., Ratnasingham, S., & de Waard, J. R. (2003) · Proceedings of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rsbl.2003.0025 · Cited by 4,466 · cited 1× · see citing works
“BSF strains have divergences of 4.3%–4.9% (Ståhls et al. 2020), which far exceeds the standard 2%–3% for a ‘bar-code gap’ (Hebert et al. 2003).”
InfluentialIntra-sexual selection in Drosophila
Bateman, A. J. (1948) · Heredity · doi:10.1038/hdy.1948.21 · Cited by 3,643 · see citing works
SPERM COMPETITION AND ITS EVOLUTIONARY CONSEQUENCES IN THE INSECTS
PARKER, G. A. (1970) · Biological Reviews · doi:10.1111/j.1469-185x.1970.tb01176.x · Cited by 3,596 · cited 1× · see citing works
“Multiple mating by females influences sperm competition among males, a ubiquitous force in insect reproduction (Parker 1970).”
Phenotypic Plasticity and the Origins of Diversity
West-Eberhard, M. J. (1989) · Annual Review of Ecology and Systematics · doi:10.1146/annurev.es.20.110189.001341 · Cited by 2,110 · cited 1× · see citing works
“Phenotypic plasticity (West-Eberhard 1989) driven by larval diet influences key reproductive traits in BSF, including adult body size, fecundity and sperm length, that also interacts with heritable variation to shape evolutionary potential (Gobbi et al. 2013; Shrestha et al. 2025; Zhang and…”
The Evolution of Insect Mating Systems
Thornhill, R., & Alcock, J. (1983) · doi:10.4159/harvard.9780674433960 · Cited by 1,954 · see citing works
The function of nuptial feeding in insects: a review of empirical studies
VAHED, K. (1998) · Biological Reviews · doi:10.1111/j.1469-185x.1997.tb00025.x · Cited by 645 · cited 1× · see citing works
“Comparative studies show how sperm traits evolve under such competition: Drosophila bifurca produces the longest sperm in the animal kingdom, whereas lepidopterans package sperm into nutrient-rich spermatophores (Pitnick et al. 1995; Vahed 1998).”
A cost of mating in female fruitflies
Fowler, K., & Partridge, L. (1989) · Nature · doi:10.1038/338760a0 · Cited by 570 · cited 1× · see citing works
“Male harassment to secure these extra matings may interfere with female oviposition or even induce injury (Pitnick and García–González 2002), such that increased exposure to males may be negatively associated with female lifetime fitness (Fowler and Partridge 1989; Morrow and Gage 2001).”
Dongen, S. V. (2006) · Journal of Evolutionary Biology · doi:10.1111/j.1420-9101.2006.01175.x · Cited by 560 · see citing works
Chapman, T., Bangham, J., Vinti, G., Seifried, B., Lung, O., Wolfner, M. F., Smith, H. K., & Partridge, L. (2003) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.1631635100 · Cited by 550 · cited 1× · see citing works
“Moreover, it is unclear whether seminal fluid proteins modulate such a switch in females; molecules similar to sex peptide in Drosophila that are transferred in the male ejaculate, are famous for triggering a cascade of behavioural and physiological changes in females (Chapman et al. 2003).”
Höglund, ., & Alatalo, R. V. (1995) · Princeton University Press eBooks · doi:10.1515/9781400864157 · Cited by 514 · see citing works
Sex Peptide Causes Mating Costs in Female Drosophila melanogaster
Wigby, S., & Chapman, T. (2005) · Current Biology · doi:10.1016/j.cub.2005.01.051 · Cited by 498 · cited 3× · see citing works
“Often, the number of matings that optimizes fitness for males is higher than it is for females (Wigby and Chapman 2005), driving such conflicts as additional matings for females may be less advantageous for their fitness, but obtaining too few matings will be sub-optimal for male fitness.”
InfluentialRearing Methods for the Black Soldier Fly (Diptera: Stratiomyidae): Table 1
Sheppard, D. C., Tomberlin, J. K., Joyce, J. A., Kiser, B. C., & Sumner, S. M. (2002) · Journal of Medical Entomology · doi:10.1603/0022-2585-39.4.695 · Cited by 479 · cited 4× · see citing works
“Females lay eggs in small, dry crevices near the actual oviposition substrate that is often wet and humid (Sheppard et al. 2002).”
InfluentialThe function of nuptial feeding in insects: a review of empirical studies
VAHED, K. (1998) · Biological Reviews of the Cambridge Philosophical Society · doi:10.1017/s0006323197005112 · Cited by 457 · see citing works
Surendra, K. C., Tomberlin, J. K., van Huis, A., Cammack, J. A., Heckmann, L.-H. L., & Khanal, S. K. (2020) · Waste Management · doi:10.1016/j.wasman.2020.07.050 · Cited by 456 · cited 2× · see citing works
“Freitas Spindola 2019; Macavei et al. 2020; Malawey et al. 2020; Surendra et al. 2020; Awal et al. 2022).”
Nitrogen Journey in Plants: From Uptake to Metabolism, Stress Response, and Microbe Interaction
Zayed, O., Hewedy, O. A., Abdelmoteleb, A., Ali, M., Youssef, M. S., Roumia, A. F., Seymour, D. K., & Yuan, Z. (2023) · Biomolecules · doi:10.3390/biom13101443 · Cited by 436 · cited 1× · see citing works
“Although not necessary for BSF, live plants can help modulate ambient humidity via transpiration of water vapor, as well as to ameliorate air quality by assimilating ammonia (NH3) into their leaf tissues (Zayed et al. 2023).”
Eberhard, W. G., Huber, B. A., S., R. L. R., Briceño, R. D., Salas, I., & Rodriguez, V. (1998) · Evolution · doi:10.1111/j.1558-5646.1998.tb01642.x · Cited by 389 · see citing works
What’s so special about model organisms?
Ankeny, R. A., & Leonelli, S. (2011) · Studies in History and Philosophy of Science Part A · doi:10.1016/j.shpsa.2010.11.039 · Cited by 387 · cited 1× · see citing works
“Although invaluable, D. melanogaster represents a narrow slice of insect diversity, and its utility has often stemmed from laboratory tractability (Ankeny and Leonelli 2011).”
Capital breeding and income breeding: their meaning, measurement, and worth
Stephens, P. A., Boyd, I. L., McNamara, J. M., & Houston, A. I. (2009) · Ecology · doi:10.1890/08-1369.1 · Cited by 382 · cited 2× · see citing works
“As mentioned, being a (hybrid) capital breeder (Stephens et al. 2009), adult reproduction relies almost entirely on larval-derived reserves, thereby decoupling gametogenesis and courtship investment from adult foraging.”
Tomberlin, J. K., Sheppard, D. C., & Joyce, J. A. (2002) · Annals of the Entomological Society of America · doi:10.1603/0013-8746(2002)095[0379:slhtob]2.0.co;2 · Cited by 363 · cited 6× · see citing works
“This latency likely serves to synchronise their reproductive readiness with female oocyte maturation (Tomberlin et al. 2002; Meyermans et al. 2025) and allows them time to establish positions within lekking aggregations (Lemke, Li, Dickerson, et al. 2025).”
InfluentialDelayed male maturity is a cost of producing large sperm in Drosophila.
Pitnick, S., Markow, T. A., & Spicer, G. S. (1995) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.92.23.10614 · Cited by 345 · see citing works
Towards a resolution of the lek paradox
Kotiaho, J. S., Simmons, L. W., & Tomkins, J. L. (2001) · Nature · doi:10.1038/35070557 · Cited by 301 · cited 1× · see citing works
“Thus far, no studies have addressed how genetic diversity might be maintained over time for BSF in light of the lekking paradox (Kotiaho et al. 2001).”
Zhan, S., Fang, G., Cai, M., Kou, Z., Xu, J. Q., Cao, Y., Bai, L., Zhang, Y., Jiang, Y., Luo, X., et al. (2019) · Cell Research · doi:10.1038/s41422-019-0252-6 · Cited by 261 · cited 2× · see citing works
“Characterised genetics : BSF have 7 chromosomes with a genome size of approximately 1.01 Gb with roughly 67% of this comprised of repetitive, noncoding, or transposable elements (Zhan et al. 2020).”
InfluentialWhy do males emerge before females? protandry as a mating strategy in male and female butterflies
Fagerström, T., & Wiklund, C. (1982) · Oecologia · doi:10.1007/bf00363830 · Cited by 245 · cited 1× · see citing works
“It is thought to allow males to monopo-lise access by being present and ready to mate as soon as females become receptive, thereby minimizing the time females spend unmated (Zhang, Henawy, et al. 2025; Fagerström and Wiklund 1982).”
Temperature and Insect Development
Ratte, H. T. (1984) · Springer eBooks · doi:10.1007/978-3-642-70020-0_2 · Cited by 245 · see citing works
COPULATORY COURTSHIP AND CRYPTIC FEMALE CHOICE IN INSECTS
EBERHARD, W. G. (1991) · Biological Reviews · doi:10.1111/j.1469-185x.1991.tb01133.x · Cited by 240 · cited 1× · see citing works
“For instance, insects such as Drosophila , crickets and butterflies can bias sperm use through differential storage or ejection of sperm from less-preferred males (Eberhard 1991).”
Tomberlin, J. K., & Sheppard, D. C. (2002) · Journal of Entomological Science · doi:10.18474/0749-8004-37.4.345 · Cited by 238 · cited 8× · see citing works
“Longevity in both sexes can be extended by provisioning adults with water (Tomberlin and Sheppard 2002) and food (Lemke For BSF in captivity, the majority of matings occur on the first or second day after being introduced to mating cages, but continue to occur at a low rate after this, reaching a…”
InfluentialEvolution of Competitive Mate Searching
Parker, G. A. (1978) · Annual Review of Entomology · doi:10.1146/annurev.en.23.010178.001133 · Cited by 237 · cited 1× · see citing works
“Moreover, the lek mating system itself also can be modelled along a gradient of increasing resource dependence and mate monopolization (Emlen and Oring 1977; Parker 1978; Thornhill and Alcock 1983) and can shift dynamically if these underlying factors change.”
GOBBI, P., MARTÍNEZ-SÁNCHEZ, A., & ROJO, S. (2013) · European Journal of Entomology · doi:10.14411/eje.2013.061 · Cited by 233 · cited 3× · see citing works
“…(West-Eberhard 1989) driven by larval diet influences key reproductive traits in BSF, including adult body size, fecundity and sperm length, that also interacts with heritable variation to shape evolutionary potential (Gobbi et al. 2013; Shrestha et al. 2025; Zhang and Puniamoorthy 2025).”
InfluentialThe secret lives of Drosophila flies
Markow, T. A. (2015) · eLife · doi:10.7554/elife.06793 · Cited by 229 · see citing works
Chia, S. Y., Tanga, C. M., Khamis, F. M., Mohamed, S. A., Salifu, D., Sevgan, S., Fiaboe, K. K. M., Niassy, S., van Loon, J. J. A., Dicke, M., et al. (2018) · PLOS ONE · doi:10.1371/journal.pone.0206097 · Cited by 221 · cited 4× · see citing works
“Moreover, industrial studies suggests that reproduction in BSF can be responsive to environmental cues (Chia et al. 2018; Addeo et al. 2022).”
InfluentialHarm to females increases with male body size in<i>Drosophila melanogaster</i>
Pitnick, S., & García–González, F. (2002) · Proceedings of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rspb.2002.2090 · Cited by 218 · see citing works
How sexual selection can drive the evolution of costly sperm ornamentation
Lüpold, S., Manier, M. K., Puniamoorthy, N., Schoff, C., Starmer, W. T., Luepold, S. H. B., Belote, J. M., & Pitnick, S. (2016) · Nature · doi:10.1038/nature18005 · Cited by 196 · see citing works
Self-citationCapital Breeders and Income Breeders among Lepidoptera: Consequences to Population Dynamics
Tammaru, T., & Haukioja, E. (1996) · Oikos · doi:10.2307/3545946 · Cited by 194 · cited 2× · see citing works
“Together, these processes place BSF among the minority of capital-breeding insects (e.g., Lepidoptera of the families Notodontidae, Arctiidae, Lymantriidae, Saturniidae and Lasiocampidae (Tammaru and Haukioja 1996) and others) in which both sperm competition and cryptic female choice can be studied under experimental…”
Nyakeri, E., Ogola, H. J. O., Ayieko, M. A., & Amimo, F. (2017) · Journal of Insects as Food and Feed · doi:10.3920/jiff2017.0004 · Cited by 178 · cited 1× · see citing works
“Most accounts of wild-trapped females are typically associated with anthropogenic waste (Sripontan et al. 2017; Nyakeri et al. 2017; Ewusie et al. 2019; Ferdousi et al. 2022; Purkayastha and Sarkar 2023; Sable and Chavan 2024; Yaseen et al. 2025).”
Behavioral aspects of mass-rearing of insects
Boller, E. (1972) · Entomophaga · doi:10.1007/bf02371070 · Cited by 157 · cited 1× · see citing works
“Although larval traits dominate economic models (Zaalberg et al. 2024), adult reproduction is the rate-limiting step in closed-cycle rearing (Boller 1972), leading some industry producers to specialise strictly in BSF egg/neonate production and selling these to other producers who then can…”
Tomberlin, J. K., & van Huis, A. (2020) · Journal of Insects as Food and Feed · doi:10.3920/jiff2020.0003 · Cited by 154 · see citing works
Kaya, C., Generalovic, T. N., Ståhls, G., Hauser, M., Samayoa, A. C., Nunes-Silva, C. G., Roxburgh, H., Wohlfahrt, J., Ewusie, E. A., Kenis, M., et al. (2021) · BMC Biology · doi:10.1186/s12915-021-01029-w · Cited by 149 · cited 3× · see citing works
“Yet, BSF exist worldwide (Kaya et al. 2021), in presumably a wide range of ecological contexts.”
Nakamura, S., Ichiki, R. T., Shimoda, M., & Morioka, S. M. (2015) · Applied Entomology and Zoology · doi:10.1007/s13355-015-0376-1 · Cited by 141 · cited 4× · see citing works
“…dynamic physiological and behavioural adaptations that challenge earlier assumptions of BSF as a monogamous or behaviourally uniform species (Nakamura et al. 2016; Park et al. 2017; Giunti et al. 2018; Short lifespan : In captivity, BSF take ~5 days to hatch from eggs, ~2–3 weeks to develop…”
InfluentialThe Evolution of Insect Mating Systems
Simmons, L., & Shuker, D. M. (2014) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780199678020.001.0001 · Cited by 133 · see citing works
Advances in deciphering the genetic basis of insect cuticular hydrocarbon biosynthesis and variation
Holze, H., Schrader, L., & Buellesbach, J. (2020) · Heredity · doi:10.1038/s41437-020-00380-y · Cited by 130 · cited 1× · see citing works
“For instance, the CHCs of Drosophila serve as short-range signals indicating sex, reproductive status, age and social rank, which together mediate attractiveness and can be manipulated to dupe rivals and mates (Laturney and Billeter 2016; Holze et al. 2021).”
Leks and hilltopping in insects
Alcock, J. (1987) · Journal of Natural History · doi:10.1080/00222938700771041 · Cited by 129 · cited 2× · see citing works
“However, lekking itself is no longer viewed within a strict definition (Alcock 1987).”
Zhang, J., Huang, L., He, J., Tomberlin, J. K., Li, J., Lei, C., Sun, M., Liu, Z., & Yu, Z. (2010) · Journal of Insect Science · doi:10.1673/031.010.20201 · Cited by 124 · cited 1× · see citing works
“…the temperature and humidity constraints of a tropical fly (Lemke, Smith, Smink, et al. 2025), factors such as like light spectra and intensity (Zhang et al. 2010), specific substrate volatiles that attract females for oviposition (Zheng et al. 2013) and even the physical design of mating…”
Pheromonal communication involved in courtship behavior in Diptera
Wicker-Thomas, C. (2007) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2007.07.003 · Cited by 121 · see citing works
Lambkin, C. L., Sinclair, B. J., Pape, T., et al. (2012) · Systematic Entomology · doi:10.1111/j.1365-3113.2012.00652.x · Cited by 118 · cited 1× · see citing works
“…(including mosquitoes, gnats, midges, black flies, drain flies, sand flies and crane flies) and Brachycera flies (including hover flies, Tsetse flies, blow flies, bot flies, Tachnid flies, true fruit flies, stalk-eyed flies and vinegar flies) (Yeates and Wiegmann 2005; Lambkin et al. 2013).”
Barrett, M., Chia, S. Y., Fischer, B., & Tomberlin, J. K. (2022) · Journal of Insects as Food and Feed · doi:10.3920/jiff2022.0041 · Cited by 115 · see citing works
Optimization of black soldier fly (Hermetia illucens) artificial reproduction
Hoc, B., Noël, G., Carpentier, J., Francis, F., & Caparros Megido, R. (2019) · PLOS ONE · doi:10.1371/journal.pone.0216160 · Cited by 113 · cited 1× · see citing works
“Fecundity and fertility (i.e., the proportion of larvae that hatch) can trade-off at industrial scale (Hoc et al. 2019) and recent study suggests that larval nutrition can induce phenotypic plasticity in both female fecundity as well as fertility (Zhang and Puniamoorthy 2025) (see Box 6).”
Zheng, L., Crippen, T. L., Holmes, L., Singh, B., Pimsler, M. L., Benbow, M. E., Tarone, A. M., Dowd, S., Yu, Z., Vanlaerhoven, S. L., et al. (2013) · Scientific Reports · doi:10.1038/srep02563 · Cited by 110 · cited 3× · see citing works
“It is these microbes (i.e., colonizing the surface of eggs and larvae, as well as living in the gut and substrates of the larval media) (Heussler et al. 2023) which release VOCs that have evolved to be attractive to mature females, driving detection and oviposition behaviour (Zheng et al. 2013).”
A high-quality, chromosome-level genome assembly of the Black Soldier Fly ( Hermetia illucens L.)
Generalovic, T. N., McCarthy, S., Warren, I. A., Wood, J. M. D., Torrance, J., Sims, Y., Quail, M. A., Howe, K., Pipan, M., Durbin, R. M., et al. (2021) · G3 Genes Genomes Genetics · doi:10.1093/g3journal/jkab085 · Cited by 109 · see citing works
My favorite molecule. The sex‐peptide
Kubli, E. (1992) · BioEssays · doi:10.1002/bies.950141111 · Cited by 107 · cited 1× · see citing works
“In addition, males may transfer seminal fluid proteins (e.g., sex peptides) that influence female re-mating latency or stimulate reproductive development (Kubli 1992).”
Lalander, C., Ermolaev, E., Wiklicky, V., & Vinnerås, B. (2020) · Science of The Total Environment · doi:10.1016/j.scitotenv.2020.138968 · Cited by 106 · cited 1× · see citing works
“Throughout their development, BSF larvae produce high amounts of heat (Li et al. 2023) and gaseous waste products (e.g., carbon dioxide (CO 2 ), ammonia (NH 3 ), etc.) (Coudron et al. 2024) as they digest their substrate, which must all be properly ventilated (Lalander et al. 2020).”
Laturney, M., & Billeter, J. (2016) · Nature Communications · doi:10.1038/ncomms12322 · Cited by 104 · see citing works
Lekking Behavior of the Black Soldier Fly (Diptera: Stratiomyidae)
Tomberlin, J. K., & Sheppard, D. C. (2001) · The Florida Entomologist · doi:10.2307/3496413 · Cited by 102 · cited 9× · see citing works
“The mating system of BSF is described as ‘lekking’ (Tomberlin and Sheppard 2001) or ‘lek-like’ (Birrell 2018), in which males aggregate at spatially distinct sites which females visit strictly to mate (Alcock 1990).”
InfluentialSame-sex sexual behavior in insects and arachnids: prevalence, causes, and consequences
Scharf, I., & Martin, O. Y. (2013) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-013-1610-x · Cited by 98 · cited 1× · see citing works
“Interestingly, the examination of same-sex sexual (SSS) behaviours in 111 other insects and arthropods revealed that SSS is more common between males than females and is highly associated with captivity, high-rearing densities and the presence of female pheromones (Scharf and Martin 2013).”
Insect Cuticular Hydrocarbons as Dynamic Traits in Sexual Communication
Ingleby, F. C. (2015) · Insects · doi:10.3390/insects6030732 · Cited by 85 · see citing works
Oonincx, D. G. A. B., Volk, N., Diehl, J. J. E., van Loon, J. J. A., & Belušič, G. (2016) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2016.10.006 · Cited by 84 · cited 1× · see citing works
“These honest signals putatively are detected visually thanks to high photoreceptive sensitivity of the ommatidia in BSF compound eyes (Oonincx et al. 2016).”
Bertinetti, C., Samayoa, A. C., & Hwang, S. (2019) · Journal of Insect Science · doi:10.1093/jisesa/iez001 · Cited by 81 · cited 1× · see citing works
“Their apparent preference for complex sugars against a white background (Romano et al. 2020), suggesting they are pollinators of white, nectar-producing flowers (Bertinetti et al. 2019).”
Female resistance and harmonic convergence influence male mating success in Aedes aegypti
Aldersley, A., & Cator, L. J. (2019) · Scientific Reports · doi:10.1038/s41598-019-38599-3 · Cited by 79 · see citing works
Oh, the places they’ll go: Female sperm storage and sperm precedence in Drosophila melanogaster
Schnakenberg, S. L., Siegal, M. L., & Bloch Qazi, M. C. (2012) · Spermatogenesis · doi:10.4161/spmg.21655 · Cited by 76 · cited 1× · see citing works
“Last-male sperm precedence may arise—not just males remove or displace their rivals' sperm—but if the female herself digests/ ejects sperm from her genital tract prior to a subsequent mating (Luck et al. 2007; Schnakenberg et al. 2012).”
Buzatto, B. A., & Machado, G. (2008) · Behavioral Ecology and Sociobiology · doi:10.1007/s00265-008-0638-9 · Cited by 72 · cited 1× · see citing works
“Conversely, when females become rare, the mating system can shift to either resource-defence polygyny or female-defence polygyny once males start to monopolise female resources or the females themselves (Buzatto and Machado 2008), respectively.”
Sexual conflict as fuel for evolution
Chapman, T., & Partridge, L. (1996) · Nature · doi:10.1038/381189a0 · Cited by 71 · cited 1× · see citing works
“…2019; Plesnar-Bielak and Łukasiewicz 2021) because interactions be-tween males and females can impose fitness costs on one sex while benefiting the other, potentially driving coevolutionary dynamics between the prevalence of male manipulation and female resistance (Chapman and Partridge 1996).”
Chimeno, C., Hausmann, A., Schmidt, S., Raupach, M. J., Doczkal, D., BARANOV, V. A., Hübner, J., Höcherl, A., Albrecht, R., Jaschhof, M., et al. (2022) · Insects · doi:10.3390/insects13010082 · Cited by 71 · cited 2× · see citing works
“However, this makes BSF relatively large compared to other insects (considering most insect diversity is housed within dark taxa which are generally tiny/miniscule Diptera and Hymenoptera (Chimeno et al. 2022)).”
Ståhls, G., Meier, R., Sandrock, C., Hauser, M., Šašić Zorić, L., Laiho, E., Aracil, A., Doderović, J., Badenhorst, R., Unadirekkul, P., et al. (2020) · BMC Evolutionary Biology · doi:10.1186/s12862-020-01627-2 · Cited by 70 · cited 5× · see citing works
“…diversity resides within Neotropics strains, although industrial strains often harbour a narrowed gene pool due to breeding stocks having shared provenance, founding events from colony establishments and repeated genetic bottlenecks (Ståhls et al. 2020; Sandrock et al. 2021; Kaya et al. 2021).”
InfluentialSperm competition experiments between lines of crickets producing different sperm lengths
Morrow, E. H., & Gage, M. J. G. (2001) · Proceedings of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rspb.2001.1807 · Cited by 61 · see citing works
Liedo, P., Salgado, S., Oropeza, A., et al. (2007) · Florida Entomologist · doi:10.1653/0015-4040(2007)90[33:impoms]2.0.co;2 · Cited by 60 · see citing works
Giunti, G., Campolo, O., Laudani, F., & Palmeri, V. (2018) · Entomologia Generalis · doi:10.1127/entomologia/2018/0657 · Cited by 60 · cited 6× · see citing works
“…adaptations that challenge earlier assumptions of BSF as a monogamous or behaviourally uniform species (Nakamura et al. 2016; Park et al. 2017; Giunti et al. 2018; Short lifespan : In captivity, BSF take ~5 days to hatch from eggs, ~2–3 weeks to develop through 6 larval instars, ~2 weeks to…”
InfluentialComputer vision and deep learning in insects for food and feed production: A review
Nawoya, S., Ssemakula, F., Akol, R. N., Geissmann, Q., Karstoft, H., Bjerge, K., Mwikirize, C., Katumba, A., & Gebreyesus, G. (2023) · Computers and Electronics in Agriculture · doi:10.1016/j.compag.2023.108503 · Cited by 60 · cited 1× · see citing works
“Coupled with their dark bodies, this enables BSF to be more easily studied via videography and computer vision applications (Nawoya et al. 2024; Smetana et al. 2025) because less specialised hardware is needed to capture high contrast images of adult behaviour.”
Cold Hardiness of the Black Soldier Fly (Diptera: Stratiomyidae)
Spranghers, T., Noyez, A., Schildermans, K., & CLERCQ, P. D. (2017) · Journal of Economic Entomology · doi:10.1093/jee/tox142 · Cited by 59 · cited 1× · see citing works
“In the wild, BSF breed year-round in the tropics (Tomberlin and Sheppard 2001), but this is restricted to fewer generations in temperate regions where presumably they must overwinter (Spranghers et al. 2017).”
Why do males emerge before females? Sexual size dimorphism drives sexual bimaturism in insects
Teder, T., Kaasik, A., Taits, K., & Tammaru, T. (2021) · Biological Reviews · doi:10.1111/brv.12762 · Cited by 59 · cited 3× · see citing works
“This strategy is common in butterflies and other insects, occurring in ~36% of investigated insect species (Teder et al. 2021).”
Optimization of Hermetia illucens (L.) egg laying under different nutrition and light conditions
Macavei, L. I., Benassi, G., STOIAN, V. H., & Maistrello, L. (2020) · PLoS ONE · doi:10.1371/journal.pone.0232144 · Cited by 58 · cited 1× · see citing works
“Freitas Spindola 2019; Macavei et al. 2020; Malawey et al. 2020; Surendra et al. 2020; Awal et al. 2022).”
Rhode, C., Badenhorst, R., Hull, K. L., Greenwood, M. P., Bester‐van der Merwe, A. E., Andere, A. A., Picard, C. J., & Richards, C. (2020) · Animal Genetics · doi:10.1111/age.12961 · Cited by 58 · cited 4× · see citing works
“For instance, although some studies document inbreeding depression and reduced progeny in captive populations (Rhode et al. 2020; Laudani et al. 2024), another shows the opposite, with increased production traits as a result of inbreeding (Cai et al. 2022).”
InfluentialBehaviour of two fly species reared for livestock feed: optimising production and insect welfare
Kortsmit, Y., van der Bruggen, M., Wertheim, B., Dicke, M., Beukeboom, L. W., & van Loon, J. J. A. (2022) · Journal of Insects as Food and Feed · doi:10.3920/jiff2021.0214 · Cited by 58 · cited 1× · see citing works
“Past reviews treated either the fundamental (Lemke et al. 2023) or the applied (Barrett, Chia, et al. 2023; Kortsmit et al. 2023; Meneguz et al. 2023) aspects of BSF biology in isolation.”
An ordination of life histories using morphological proxies: capital vs. income breeding in insects
Davis, R. B., Javoiš, J., Kaasik, A., Õunap, E., & Tammaru, T. (2016) · Ecology · doi:10.1002/ecy.1435 · Cited by 57 · cited 1× · see citing works
“Although classically described as capital breeders, providing BSF adults supplemental nutrition pushes them slightly towards the middle of the capital-income continuum (Davis et al. 2016).”
How does breeding system variation modulate sexual antagonism?
Bedhomme, S., Bernasconi, G., Koene, J. M., Lankinen, Å., Arathi, H. S., Michiels, N. K., & Anthes, N. (2009) · Biology Letters · doi:10.1098/rsbl.2009.0401 · Cited by 54 · cited 1× · see citing works
“The mating system of BSF may be subject to sexual antagonism or sexual conflict (Bedhomme et al. 2009; Candolin 2019; Plesnar-Bielak and Łukasiewicz 2021) because interactions be-tween males and females can impose fitness costs on one sex while benefiting the other, potentially driving…”
Field Evaluation of a Push-Pull System to Reduce Malaria Transmission
Menger, D. J., Omusula, P., Holdinga, M., Homan, T., Carreira, A. S., Vandendaele, P., Derycke, J.-L., Mweresa, C. K., Mukabana, W. R., van Loon, J. J. A., et al. (2015) · PLOS ONE · doi:10.1371/journal.pone.0123415 · Cited by 53 · cited 1× · see citing works
“…VOCs elicit stronger responses than individual components alone (Thomas et al. 2024), this means that a precise blend can be engineered as part of a push–pull strategy (Menger et al. 2015; Cui et al. 2022) to more precisely direct egg-laying both away from cage materials and towards the trap.”
Last male sperm precedence is modulated by female remating rate in <i>Drosophila melanogaster</i>
Laturney, M., van Eijk, R., & Billeter, J.-C. (2018) · Evolution Letters · doi:10.1002/evl3.50 · Cited by 53 · cited 1× · see citing works
“However, the degree to which sperm precedence follows ‘last male wins’ patterns, as seen in other Diptera, is unknown, nor how this pattern may subsequently break down with high amounts of remating (Zeh and Zeh 1997; Laturney et al. 2018) or as flies senesce (Mack et al. 2003).”
Female age and sperm competition: last-male precedence declines as female age increases
Mack, P. D., Priest, N. K., & Promislow, D. E. L. (2003) · Proceedings of the Royal Society of London. Series B: Biological Sciences · doi:10.1098/rspb.2002.2214 · Cited by 49 · see citing works
Julita, U., Lusianti F, L., Eka Putra, R., & Dana Perma, A. (2020) · Journal of Entomology · doi:10.3923/je.2020.117.127 · Cited by 49 · cited 2× · see citing works
“Reports of male–male mating interactions led to the hypothesis that BSF males are largely indiscriminate (Julita et al. 2020; Jones and Tomberlin 2021; Chiabotto et al. 2024; Kortsmit et al. 2025).”
Acoustic Pattern Recognition and Courtship Songs: Insights from Insects
Baker, C. A., Clemens, J., & Murthy, M. (2019) · Annual Review of Neuroscience · doi:10.1146/annurev-neuro-080317-061839 · Cited by 47 · cited 1× · see citing works
“These visual displays are accompanied by vibratory ‘songs’ produced by wing-buzzing (Giunti et al. 2018; Laudani et al. 2024; Kortsmit et al. 2025), similar to many other groups of insects (e.g., crickets grasshoppers) (Baker et al. 2019).”
Lemke, N. B., Dickerson, A. J., & Tomberlin, J. K. (2022) · BioEssays · doi:10.1002/bies.202200162 · Cited by 46 · see citing works
Self-citationEwusie, E. A., Kwapong, P., Ofosu-Budu, G. K., Sandrock, C., Akumah, A. M., Nartey, E. K., Tetegaga, C., & Agyakwah, S. K. (2019) · Scientific African · doi:10.1016/j.sciaf.2019.e00134 · Cited by 45 · cited 1× · see citing works
“Most accounts of wild-trapped females are typically associated with anthropogenic waste (Sripontan et al. 2017; Nyakeri et al. 2017; Ewusie et al. 2019; Ferdousi et al. 2022; Purkayastha and Sarkar 2023; Sable and Chavan 2024; Yaseen et al. 2025).”
Patterns of Genetic Diversity and Mating Systems in a Mass-Reared Black Soldier Fly Colony
Hoffmann, L., Hull, K. L., Bierman, A., Badenhorst, R., Bester-van der Merwe, A. E., & Rhode, C. (2021) · Insects · doi:10.3390/insects12060480 · Cited by 45 · cited 3× · see citing works
“…BSF to mate up to four (Chiabotto et al. 2024; Laudani et al. 2024), five (Jensen et al. 2025) or six times (Manas, Venon, et al. 2025) and females up to twice (Chiabotto et al. 2024) but higher female polyandry, up to five times, has been supported by parentage assignments (Hoffmann et al. 2021).”
Zeni, V., Benelli, G., Campolo, O., Giunti, G., Palmeri, V., Maggi, F., Rizzo, R., Verde, G. L., Lucchi, A., & Canale, A. (2021) · Molecules · doi:10.3390/molecules26195898 · Cited by 43 · cited 2× · see citing works
“In addition to directing oviposition, aromatherapy using both plant essential oils (e.g., α-copaene derived from sweet orange, grapefruit, guava, papaya and mango; methyl eugenol; raspberry keytone; α-Pinene; Zingerone) (Zeni et al. 2021) and synthetic compounds (e.g., Trimedlure) (Shelly et al. 1993) has been shown t…”
Jones, B. M., & Tomberlin, J. K. (2021) · Journal of Insects as Food and Feed · doi:10.3920/jiff2020.0001 · Cited by 41 · cited 6× · see citing works
“Indeed BSF of multiple size-classes successfully mate, although with dramatically increased variability (Jones and Tomberlin 2021).”
InfluentialJang, E. B., McInnis, D. O., Kurashima, R., et al. (1999) · Agricultural and Forest Entomology · doi:10.1046/j.1461-9563.1999.00025.x · Cited by 40 · cited 2× · see citing works
“…could be a physiological ‘switch’ that is triggered post-mating for females (Lemke, Li, Dickerson, et al. 2025), as has been found in tephritids (Jang et al. 1999), because unmated/virgin female BSF showed no clear behavioural preferences for these chemical cues, whilst gravid females exhibited…”
Cuticular hydrocarbons of the screwworm,Cochliomyia hominivorox (Diptera: Calliphoridae)
Pomonis, J. G. (1989) · Journal of Chemical Ecology · doi:10.1007/bf01012083 · Cited by 40 · cited 1× · see citing works
“However, the New World Screwworm Fly Cochliomyia hom-inivorax is a counter-example which, like BSF, lacks sexual dimorphic CHCs altogether (Pomonis 1989).”
Black soldier fly (Diptera: Stratiomyidae) larval heat generation and management
Li, C., Addeo, N. F., Rusch, T. W., Tarone, A. M., & Tomberlin, J. K. (2023) · Insect Science · doi:10.1111/1744-7917.13198 · Cited by 39 · see citing works
Papanastasiou, S. A., Diamantidis, A. D., Nakas, C. T., Carey, J. R., & Papadopoulos, N. T. (2011) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2011.07.004 · Cited by 38 · cited 1× · see citing works
“For instance, older, less competitive males may interfere with females mating with younger mates, as observed in tephritid flies (Papanastasiou et al. 2011), which in these species has necessitated habitat design that features distinct areas for each sex (Meza et al. 2025).”
Artificial selection of insects to bioconvert pre-consumer organic wastes. A review
Fowles, T. M., & Nansen, C. (2019) · Agronomy for Sustainable Development · doi:10.1007/s13593-019-0577-z · Cited by 38 · cited 1× · see citing works
“However, the precise tolerance of BSF to inbreeding and the optimal outcrossing regime remain unresolved (Fowles and Nansen 2019).”
Addeo, N. F., Li, C., Rusch, T. W., Dickerson, A. J., Tarone, A. M., Bovera, F., & Tomberlin, J. K. (2021) · Journal of Insects as Food and Feed · doi:10.3920/jiff2021.0076 · Cited by 38 · see citing works
Sexual Selection on Leks: A Fruit Fly Primer
Shelly, T. E. (2018) · Journal of Insect Science · doi:10.1093/jisesa/iey048 · Cited by 36 · cited 1× · see citing works
“Indeed, in lek-based systems such as in true fruit flies (Tephritidae) where males display in groups, reproductive success is often determined by position within the lek, e.g., a central position or on specific trees (Shelly 2018).”
Copulation duration in the soldier fly: the roles of cryptic male choice and sperm competition risk
Barbosa, F. (2011) · Behavioral Ecology · doi:10.1093/beheco/arr137 · Cited by 35 · cited 2× · see citing works
“Retention duration, sperm viability and selective sperm use suggest potential for cryptic female choice as described in other stratiomyids (Barbosa 2009, 2011, 2012, 2015).”
Pélisson, P., Bel‐Venner, M., Rey, B., Burgevin, L., Martineau, F., Fourel, F., Lécuyer, C., Menu, F., & Venner, S. (2011) · Functional Ecology · doi:10.1111/j.1365-2435.2011.01925.x · Cited by 34 · cited 1× · see citing works
“The opposite is true as well, with capital breeding and strict proovigeny (i.e., the emergence of a female's entire potential life-time complement of eggs) being associated with stable and predictable larval conditions (Pélisson et al. 2012).”
Meneguz, M., Miranda, C. D., Cammack, J. A., & Tomberlin, J. K. (2022) · Journal of Insects as Food and Feed · doi:10.3920/jiff2022.0055 · Cited by 33 · see citing works
Butterworth, N. J., White, T. E., Byrne, P. G., & Wallman, J. F. (2021) · Journal of Evolutionary Biology · doi:10.1111/jeb.13759 · Cited by 32 · cited 1× · see citing works
“For instance, BSF exhibit sexual dimorphism in wing interference patterns created by differences in structural colouration (Butterworth et al. 2021; Rebora et al. 2024).”
Laursen, S. F., Flint, C. A., Bahrndorff, S., Tomberlin, J. K., & Kristensen, T. N. (2024) · Waste Management · doi:10.1016/j.wasman.2024.04.010 · Cited by 32 · cited 1× · see citing works
“This direct link between larval nutrition, adult size and egg number leads to significant natural variation in both fecundity (number of eggs) and fertility (hatching success) (Laursen et al. 2024; Zhang and Puniamoorthy 2025).”
Cryptic female choice by female control of oviposition timing in a soldier fly
Barbosa, F. (2009) · Behavioral Ecology · doi:10.1093/beheco/arp083 · Cited by 31 · cited 1× · see citing works
“Retention duration, sperm viability and selective sperm use suggest potential for cryptic female choice as described in other stratiomyids (Barbosa 2009, 2011, 2012, 2015).”
Sexual conflict in a changing environment
Plesnar‐Bielak, A., & Łukasiewicz, A. (2021) · Biological reviews/Biological reviews of the Cambridge Philosophical Society · doi:10.1111/brv.12728 · Cited by 31 · see citing works
Malawey, A. S., Mercati, D., Love, C. C., & Tomberlin, J. K. (2019) · Annals of the Entomological Society of America · doi:10.1093/aesa/saz045 · Cited by 30 · see citing works
Klüber, P., Arous, E., Zorn, H., & Rühl, M. (2023) · Life · doi:10.3390/life13020355 · Cited by 29 · see citing works
Shelly, T. E., Whittier, T. S., & Kaneshiro, K. Y. (1993) · Annals of the Entomological Society of America · doi:10.1093/aesa/86.3.341 · Cited by 28 · cited 2× · see citing works
“In addition to directing oviposition, aromatherapy using both plant essential oils (e.g., α-copaene derived from sweet orange, grapefruit, guava, papaya and mango; methyl eugenol; raspberry keytone; α-Pinene; Zingerone) (Zeni et al. 2021) and synthetic compounds (e.g., Trimedlure) (Shelly et al. 1993) has been shown t…”
Malawey, A. S., Zhang, H., McGuane, A. S., Walsh, E. M., Rusch, T. W., Hjelmen, C. E., Delclos, P. J., Rangel, J., Zheng, L., Cai, M., et al. (2021) · Journal of Insects as Food and Feed · doi:10.3920/jiff2020.0017 · Cited by 27 · see citing works
Boafo, H. A., Gbemavo, D. S. J. C., Timpong-Jones, E. C., et al. (2022) · Journal of Insects as Food and Feed · doi:10.3920/jiff2022.0034 · Cited by 27 · see citing works
Silvaraju, S., Zhang, Q., Kittelmann, S., & Puniamoorthy, N. (2024) · Animal Microbiome · doi:10.1186/s42523-024-00340-5 · Cited by 27 · cited 1× · see citing works
“Nevertheless, other pre-zygotic variables, including strain-specific microbi-omes (Silvaraju et al. 2024) or even behavioural variation in visual and acoustic courtship signals, have yet to be explored as barriers to gene flow.”
Self-citationThe unpaved road towards efficient selective breeding in insects for food and feed—A review
Hansen, L. S., Laursen, S. F., Bahrndorff, S., Sørensen, J. G., Sahana, G., Kristensen, T. N., & Nielsen, H. M. (2024) · Entomologia Experimentalis et Applicata · doi:10.1111/eea.13526 · Cited by 25 · see citing works
Alcock, J. (1984) · The Southwestern Naturalist · doi:10.2307/3671000 · Cited by 24 · cited 1× · see citing works
“A central challenge for reproducing insects is locating a mate, leading to the emergence of diverse strategies like pursuing, patrolling, hilltopping, perching and ambushing, just to name a few (Alcock 1984, 1987).”
Performance evaluation of black soldier fly larvae fed on human faeces, food waste and their mixture
Purkayastha, D., & Sarkar, S. (2022) · Journal of Environmental Management · doi:10.1016/j.jenvman.2022.116727 · Cited by 24 · see citing works
Males responding to sperm competition cues have higher fertilization success in a soldier fly
Barbosa, F. (2012) · Behavioral Ecology · doi:10.1093/beheco/ars035 · Cited by 23 · cited 1× · see citing works
“Retention duration, sperm viability and selective sperm use suggest potential for cryptic female choice as described in other stratiomyids (Barbosa 2009, 2011, 2012, 2015).”
Cai, M., Li, L., Zhao, Z., Zhang, K., Li, F., Yu, C., Yuan, R., Zhou, B., Ren, Z., Yu, Z., et al. (2022) · Life · doi:10.3390/life12060873 · Cited by 23 · cited 1× · see citing works
“For instance, although some studies document inbreeding depression and reduced progeny in captive populations (Rhode et al. 2020; Laudani et al. 2024), another shows the opposite, with increased production traits as a result of inbreeding (Cai et al. 2022).”
A window into fly sex: mating increases female but reduces male longevity in black soldier flies
Harjoko, D. N., Hua, Q. Q. H., Toh, E. M. C., Goh, C. Y. J., & Puniamoorthy, N. (2023) · Animal Behaviour · doi:10.1016/j.anbehav.2023.03.007 · Cited by 23 · cited 7× · see citing works
“In BSF, because such physical changes are quite conspicuous, aging can be estimated based on the decreasing opacity of the abdominal ‘windows’ (Harjoko et al. 2023), through which one can observe changes in the stored nutrient reserves (which also include changes in colour).”
InfluentialSelf-citationAlcock, J. (1990) · Psyche: A Journal of Entomology · doi:10.1155/1990/72328 · Cited by 22 · cited 5× · see citing works
“The mating system of BSF is described as ‘lekking’ (Tomberlin and Sheppard 2001) or ‘lek-like’ (Birrell 2018), in which males aggregate at spatially distinct sites which females visit strictly to mate (Alcock 1990).”
InfluentialLekking as collective behaviour
Rathore, A., Isvaran, K., & Guttal, V. (2023) · Philosophical Transactions of the Royal Society B Biological Sciences · doi:10.1098/rstb.2022.0066 · Cited by 21 · see citing works
Munsch‐Masset, P., Labrousse, C., Beaugeard, L., & Bressac, C. (2023) · Entomologia Experimentalis et Applicata · doi:10.1111/eea.13358 · Cited by 19 · see citing works
The respiratory structures in the eggs of higher Diptera
Wigglesworth, V. B., & Beament, J. W. L. (1960) · Journal of Insect Physiology · doi:10.1016/0022-1910(60)90080-9 · Cited by 18 · cited 1× · see citing works
“Indeed, adaptions of eggs to thrive in dryer substrates (e.g., development of chorion respiratory structures (Wigglesworth and Beament 1960), and protective mucilaginous secretions around egg clutches) allowed ancestral Brachycera to transition away from fully aquatic development into amphibious…”
Muraro, T., Lalanne, L., Pelozuelo, L., & Calas-List, D. (2024) · Journal of Insects as Food and Feed · doi:10.1163/23524588-20220175 · Cited by 18 · see citing works
Romano, N., Fischer, H., & Egnew, N. (2020) · Journal of Environmental Biology · doi:10.22438/jeb/41/5(si)/ms_03 · Cited by 17 · see citing works
Ammonia emissions related to black soldier fly larvae during growth on different diets
Coudron, C. L., Berrens, S., Peer, M. V., et al. (2024) · Journal of Insects as Food and Feed · doi:10.1163/23524588-00001049 · Cited by 17 · see citing works
Survival and reproduction when food is scarce: implications for a lekking Hawaiian Drosophila
Jones, T. M., & Widemo, F. (2005) · Ecological Entomology · doi:10.1111/j.0307-6946.2005.00705.x · Cited by 16 · cited 1× · see citing works
“However, adult nutrition is variable too and has negative consequences for income breeders (Jones and Widemo 2005).”
Klüber, P., Arous, E., Jerschow, J. J., Fraatz, M. A., Bakonyi, D., Rühl, M., & Zorn, H. (2023) · Insect Science · doi:10.1111/1744-7917.13287 · Cited by 16 · see citing works
Zaalberg, R. M., Nielsen, H. M., Noer, N. K., Schou, T. M., Jensen, K., Thormose, S. F., Kargo, M., & Slagboom, M. (2024) · Journal of Insects as Food and Feed · doi:10.1163/23524588-00001126 · Cited by 16 · cited 1× · see citing works
“Although larval traits dominate economic models (Zaalberg et al. 2024), adult reproduction is the rate-limiting step in closed-cycle rearing (Boller 1972), leading some industry producers to specialise strictly in BSF egg/neonate production and selling these to other producers who then can…”
Manas, F., Piterois, H., Labrousse, C., Beaugeard, L., Uzbekov, R., & Bressac, C. (2024) · Royal Society Open Science · doi:10.1098/rsos.241205 · Cited by 16 · see citing works
Tomberlin, J. K., Picard, C. J., Jordan, H. R., Preyer, C., Warburton, C., Crowley, P., Zheng, R., Boulanger, F. X., Banks, I., Lefranc, M., et al. (2022) · Journal of Insects as Food and Feed · doi:10.3920/jiff2022.x001 · Cited by 14 · see citing works
Ferdousi, L., Sultana, N. N., Bithi, U. H., Lisa, S. A., Hasan, M. R., & Siddique, M. A. B. (2022) · Proceedings of the National Academy of Sciences India Section B Biological Sciences · doi:10.1007/s40011-021-01340-0 · Cited by 14 · see citing works
Chiabotto, C., Grosso, F., Doretto, A., & Meneguz, M. (2024) · Journal of Insects as Food and Feed · doi:10.1163/23524588-20230165 · Cited by 14 · see citing works
Permana, A. D., Fitri, L. L., & Julita, U. (2020) · Jurnal Biodjati · doi:10.15575/biodjati.v5i2.9049 · Cited by 13 · cited 3× · see citing works
“This timing varies based on biological age (Dickerson et al. 2024) and mating experience (Permana et al. 2020).”
Giannetti, D., Schifani, E., Reggiani, R., Mazzoni, E., Reguzzi, M. C., Castracani, C., Spotti, F. A., Giardina, B., Mori, A., & Grasso, D. A. (2022) · Insects · doi:10.3390/insects13020127 · Cited by 13 · see citing works
CRISPR/Cas9-mediated efficient white genome editing in the black soldier fly Hermetia illucens
Sui, Z., Wu, Q., Geng, J., Xiao, J., & Huang, D. (2024) · Molecular Genetics and Genomics · doi:10.1007/s00438-023-02088-0 · Cited by 13 · see citing works
Impact of age on the reproductive output of <i>Hermetia illucens</i> (Diptera: Stratiomyidae)
Dickerson, A. J., Lemke, N. B., Li, C., & Tomberlin, J. K. (2024) · Journal of Economic Entomology · doi:10.1093/jee/toae107 · Cited by 13 · see citing works
Self-citationZhang, Q.-H., & Puniamoorthy, N. (2025) · Insects · doi:10.3390/insects16020142 · Cited by 13 · see citing works
Self-citationMale and female effects on sperm precedence in the giant sperm species Drosophila bifurca
Luck, N., Dejonghe, B., Fruchard, S., et al. (2006) · Genetica · doi:10.1007/s10709-006-9103-8 · Cited by 12 · cited 1× · see citing works
“Last-male sperm precedence may arise—not just males remove or displace their rivals' sperm—but if the female herself digests/ ejects sperm from her genital tract prior to a subsequent mating (Luck et al. 2007; Schnakenberg et al. 2012).”
Laksanawimol, P., Singsa, S., & Thancharoen, A. (2023) · PeerJ · doi:10.7717/peerj.15701 · Cited by 12 · cited 2× · see citing works
“…experiments have revealed that BSF prefer to oviposit (depending on the choices available to them) in grain/vegetable/fruit wastes (Kotzé and Tomberlin 2020; Laksanawimol et al. 2023; Zim et al. 2023) as well as the medium near aged carrion (Kotzé and Tomberlin 2020) and manure (Zim et al. 2023).”
(2009) · Americanae (AECID Library) · doi:10.15517/rbt.v57i0.21348 · Cited by 12
Jensen, K., Thormose, S. F., Noer, N. K., Schou, T. M., Kargo, M., Gligorescu, A., Nørgaard, J. V., Hansen, L. S., Zaalberg, R. M., Nielsen, H. M., et al. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10205 · Cited by 12 · see citing works
Barrett, M., Godfrey, R. K., Sterner, E. J., & Waddell, E. A. (2022) · Arthropod Structure & Development · doi:10.1016/j.asd.2022.101174 · Cited by 11 · see citing works
Early adaptations of true flies (Diptera) to moist and aquatic continental environments
Peñalver, E., Matamales‐Andreu, R., Nel, A., et al. (2022) · Papers in Palaeontology · doi:10.1002/spp2.1472 · Cited by 11 · see citing works
Coital behavior in dogs. IX. Sequelae to “coitus interruptus” in males and females
Beach, F. A. (1970) · Physiology & Behavior · doi:10.1016/0031-9384(70)90096-x · Cited by 10 · cited 1× · see citing works
“This can be contrasted with a genital lock that might be formed via the swelling of genitalia, which famously occurs in Canids (Carnivora: Canidae) (Beach 1970).”
Fine Structure of Maxillary Palps in Adults of Hermetia illucens (Diptera: Stratiomyidae)
Pezzi, M., Scapoli, C., Bharti, M., et al. (2020) · Journal of Medical Entomology · doi:10.1093/jme/tjaa251 · Cited by 10 · cited 1× · see citing works
“It has also been theorised that BSF males may not be as responsive to these volatiles, especially because female antennae have longer flagellum than males, which would suggest increased function (Pezzi et al. 2021).”
Rebora, M., Piersanti, S., Romani, A., Kovalev, A., Gorb, S., & Salerno, G. (2024) · Scientific Reports · doi:10.1038/s41598-024-70684-0 · Cited by 10 · cited 2× · see citing works
“Males perform aerial displays of wing-fanning, which combined with wing-interference patterns (WIPs) putatively serve as visual signals for conspecifics (Rebora et al. 2024).”
Barbosa, F. (2015) · Springer eBooks · doi:10.1007/978-3-319-17894-3_14 · Cited by 9 · cited 1× · see citing works
“Retention duration, sperm viability and selective sperm use suggest potential for cryptic female choice as described in other stratiomyids (Barbosa 2009, 2011, 2012, 2015).”
Kotzé, Z., & Tomberlin, J. K. (2020) · Journal of Medical Entomology · doi:10.1093/jme/tjaa021 · Cited by 9 · see citing works
Assessing the Welfare of Captive Group-Housed Cockroaches, Gromphadorhina oblongonota
Free, D., & Wolfensohn, S. (2023) · Animals · doi:10.3390/ani13213351 · Cited by 9 · cited 1× · see citing works
“By mimicking natural conditions, the effects of providing plants can potentially reduce stress, as is the case in captive cockroaches (Blattodea) (Free and Wolfensohn 2023), though in practice it may be logistically unfeasible for BSF producers to cultivate/purchase and maintain live plants as well.”
Manas, F., Labrousse, C., & Bressac, C. (2025) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2025.104751 · Cited by 9 · see citing works
Influences of artificial light on mating of black soldier fly (Hermetia illucens)—a review
Awal, M. R., Rahman, M. M., Choudhury, M. A. R., Hasan, M. M., Rahman, M. T., & Mondal, M. F. (2022) · International Journal of Tropical Insect Science · doi:10.1007/s42690-022-00786-7 · Cited by 9 · see citing works
Coudron, C. L., Adamaki-Sotiraki, C., Yakti, W., Pascual, J. J., Wiklicky, V., Sandrock, C., Van Peer, M., Athanassiou, C., Peguero, D. A., Rumbos, C., et al. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10240 · Cited by 9 · see citing works
Reproductive physiology and behaviour
Moore, P. J. (2014) · Oxford University Press eBooks · doi:10.1093/acprof:oso/9780199678020.003.0005 · Cited by 8 · cited 1× · see citing works
“This distinguishes BSF from many other proovigenic insects whose eggs are all mature at the onset of their adult life, as well as many insects whose fecundity is linked to both the number and variation of ovari-oles (Moore 2014).”
THINN, A. A., & KAINOH, Y. (2022) · Japan Agricultural Research Quarterly: JARQ · doi:10.6090/jarq.56.211 · Cited by 8 · cited 3× · see citing works
“However, BSF are not strict capital breeders and supplemental nutrition can benefit adult fitness (Thinn and Kainoh 2022; Klüber et al. 2023; Barrett et al. 2025), though this is not always consistently observed (Lemke et al. 2023; Coudron et al. 2025).”
Lemke, N. B., Rollison, L. N., & Tomberlin, J. K. (2024) · Insects · doi:10.3390/insects15100770 · Cited by 8 · see citing works
Self-citationZim, J., Chkih, H., Bouharroud, R., Sarehane, M., & Lhomme, P. (2023) · Journal of Insects as Food and Feed · doi:10.1163/23524588-20220139 · Cited by 7 · see citing works
Antennal olfactory responses in the black soldier fly Hermetia illucens
Piersanti, S., Rebora, M., Marri, G. C., et al. (2024) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2024.104722 · Cited by 7 · cited 1× · see citing works
“…recordings confirm the olfactory abilities of BSF of both sexes in response to various volatiles with substantial overlap in their responses (Piersanti et al. 2024), suggesting that differences in behavioural responses of each sex to volatiles must then be driven by sex-specific…”
Correlated response to selection for increased body weight on fecundity in <i>Hermetia illucens</i>
Shrestha, K., Junes, P., van den Boer, E., Christianen, I., Jacobse, R., & Schmitt, E. (2025) · Entomologia Experimentalis et Applicata · doi:10.1111/eea.13564 · Cited by 7 · see citing works
Genetic differentiation in domesticated black soldier fly (Hermetia illucens) populations
Silvaraju, S., Loh, R. K., Kittelmann, S., & Puniamoorthy, N. (2025) · BMC Biology · doi:10.1186/s12915-025-02480-9 · Cited by 7 · cited 7× · see citing works
“…breakdown) have not been explicitly tested, but the observed genomic divergence and reduced heterozygosity in domesticated lines raise the possibility that hybrids could suffer reduced fitness, especially under the intense selection regimes typical of industrial rearing (Silvaraju et al. 2026).”
InfluentialSelf-citationDoes Hermetia illucens recognize sibling mates to avoid inbreeding depression?
Laudani, F., Campolo, O., Latella, I., Modafferi, A., Palmeri, V., & Giunti, G. (2024) · Entomologia Generalis · doi:10.1127/entomologia/2024/2746 · Cited by 6 · see citing works
FlyCount: High-Speed Counting of Black Soldier Flies Using Neuromorphic Sensors
James, A., Seth, A., Marcireau, A., et al. (2024) · IEEE Sensors Journal · doi:10.1109/jsen.2024.3504289 · Cited by 6 · see citing works
Exploring the potential of crossbreeding to enhance black soldier fly (Hermetia illucens) production
Meyermans, R., Broeckx, L., Mondelaers, J., Gorssen, W., Frooninckx, L., Janssens, S., Van Miert, S., & Buys, N. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-00001391 · Cited by 6 · cited 6× · see citing works
“This latency likely serves to synchronise their reproductive readiness with female oocyte maturation (Tomberlin et al. 2002; Meyermans et al. 2025) and allows them time to establish positions within lekking aggregations (Lemke, Li, Dickerson, et al. 2025).”
InfluentialRepeatable phenotypic but not genetic response to selection on body size in the black soldier fly
Generalovic, T. N., Zhou, W., Zhao, L. C., Leonard, S., Warren, I. A., Pipan, M., & Jiggins, C. D. (2025) · doi:10.1101/2025.02.25.640052 · Cited by 6 · see citing works
Bruno, D., Manas, F., Bonelli, M., Gold, M., Marzari, M., Roma, D., Valoroso, M. C., Montali, A., Guillaume, J. B., Rebora, M., et al. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-20250002 · Cited by 6 · see citing works
Cammack, J. A., & Tomberlin, J. K. (2023) · Journal of Insects as Food and Feed · doi:10.1163/23524588-20230026 · Cited by 6 · see citing works
Sexual Selection and Sexual Conflict
Candolin, U. (2018) · Elsevier eBooks · doi:10.1016/b978-0-12-409548-9.10886-3 · Cited by 5 · cited 1× · see citing works
“The mating system of BSF may be subject to sexual antagonism or sexual conflict (Bedhomme et al. 2009; Candolin 2019; Plesnar-Bielak and Łukasiewicz 2021) because interactions be-tween males and females can impose fitness costs on one sex while benefiting the other, potentially driving…”
Cui, Z., Si, P., Liu, L., Chen, S., Wang, Y., Li, X., Zhou, J., & Zhou, Q. (2022) · Journal of Applied Entomology · doi:10.1111/jen.13067 · Cited by 5 · cited 1× · see citing works
“…VOCs elicit stronger responses than individual components alone (Thomas et al. 2024), this means that a precise blend can be engineered as part of a push–pull strategy (Menger et al. 2015; Cui et al. 2022) to more precisely direct egg-laying both away from cage materials and towards the trap.”
Zhang, Q., Ng, K. H., Chin, W. S., Tang, Y. J., Lin, J., & Puniamoorthy, N. (2025) · Animal Behaviour · doi:10.1016/j.anbehav.2025.123299 · Cited by 5 · cited 8× · see citing works
“This direct link between larval nutrition, adult size and egg number leads to significant natural variation in both fecundity (number of eggs) and fertility (hatching success) (Laursen et al. 2024; Zhang and Puniamoorthy 2025).”
InfluentialSelf-citationManas, F., Venon, P., Yang, L., Labrousse, C., & Bressac, C. (2025) · Entomologia Experimentalis et Applicata · doi:10.1111/eea.13595 · Cited by 4 · cited 5× · see citing works
“Moreover the duration of the genital lock appears mostly unaffected by adult size but varies with age (Manas, Labrousse, and Bressac 2025; Manas, Venon, et al. 2025), suggesting that BSF might not engage in mate guarding, which in other species can arise as a mating tactic to prevent females from…”
InfluentialMeza, J. S., Ibañez‐Palacios, J., Cardenas‐Enriquez, D. P., Luis‐Alvares, J. H., & Liedo, P. (2025) · Insect Science · doi:10.1111/1744-7917.70105 · Cited by 4 · cited 3× · see citing works
“In other mass-reared Diptera, injuries are often associated with harassment (Meza et al. 2025) and in captive male tephritids typically become hyper-aggressive under artificial selection (Briceño et al. 1999).”
Exploring interkingdom communication: the case of black soldier fly mass-rearing
Klammsteiner, T., Heussler, C. D., Insam, H., Tomberlin, J. K., Schlick-Steiner, B. C., & Steiner, F. M. (2025) · Current Opinion in Insect Science · doi:10.1016/j.cois.2025.101427 · Cited by 4 · see citing works
Barrett, M., Patel, N., McCarry, B., Shellenberger, G. K., Schwartz, E. R., Fiocca, K., & Waddell, E. A. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10295 · Cited by 4 · cited 5× · see citing works
“Adult BSF's access to water and nutrient supplements (e.g., honey, pollen, or sugar water) can extend lifespan, shorten pre-oviposition intervals and increase egg production under several conditions (Thinn and Kainoh 2022; Klüber et al. 2023; Barrett et al. 2025).”
InfluentialSensory morphology, investment and plasticity in the black soldier fly (Hermetia illucens)Preprint
Birrell, N. W. (2019) · doi:10.31237/osf.io/v8zs2 · Cited by 4 · see citing works
The hidden sweet tooth of the black soldier fly ( <i>Hermetia illucens</i> )
Merle, M., Gueddes, T., Gueye, M. M., Foughar, M., Jiogue-Lacdo, J., Maquart, P.-O., Marion-Poll, F., & Filée, J. (2026) · Journal of Experimental Biology · doi:10.1242/jeb.252160 · Cited by 3 · cited 10× · see citing works
“For instance, insects such as Drosophila , crickets and butterflies can bias sperm use through differential storage or ejection of sperm from less-preferred males (Eberhard 1991).”
InfluentialStage-Specific Microbiota Transitions Throughout Black Soldier Fly Ontogeny
Klammsteiner, T., Heussler, C. D., Stonig, K. T., Insam, H., Schlick-Steiner, B. C., & Steiner, F. M. (2026) · Microbial Ecology · doi:10.1007/s00248-025-02691-1 · Cited by 3 · see citing works
Lemke, N. B., Li, C., Dickerson, A. J., Salazar, D. A., Rollinson, L. N., Mendoza, J. E., Miranda, C. D., Crawford, S., & Tomberlin, J. K. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10275 · Cited by 3 · see citing works
Self-citationBugBook: Data analysis methods in studies of insects for food and feed
Smetana, S. M., Coudron, C. L., Deruytter, D., Francis, A., Pascual, J. J., Klammsteiner, T., Lemke, N. B., Sandrock, C., & Zanoli, R. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10209 · Cited by 3 · see citing works
Self-citationSable, A. V., & Chavan, K. (2024) · UTTAR PRADESH JOURNAL OF ZOOLOGY · doi:10.56557/upjoz/2024/v45i184418 · Cited by 2 · cited 1× · see citing works
“Most accounts of wild-trapped females are typically associated with anthropogenic waste (Sripontan et al. 2017; Nyakeri et al. 2017; Ewusie et al. 2019; Ferdousi et al. 2022; Purkayastha and Sarkar 2023; Sable and Chavan 2024; Yaseen et al. 2025).”
Parentage assignment in black soldier fly (Hermetia illucens) using genotyping-by-sequencing
Dufresne, G., Bolduc, C., Warburton, C., Vandenberg, G. W., Deschamps, M., & Alnahhas, N. (2025) · Frontiers in Genetics · doi:10.3389/fgene.2025.1541812 · Cited by 2 · cited 1× · see citing works
“Recent work has begun to characterise BSF genetics (i.e., via a chromosome assembly (Generalovic et al. 2021), parentage assemblies (Hoffmann et al. 2021; Dufresne et al. 2025), quantification of inbreeding in captivity (Rhode et al. 2020) and more).”
Lemke, N. B., Smith, M. B., Smink, J. A., Jordan, H. R., Picard, C. J., & Tomberlin, J. K. (2025) · doi:10.20944/preprints202512.0942.v1 · Cited by 2 · see citing works
Self-citationInfluence of diet on initial mating behaviour in the black soldier fly (Hermetia illucens)
Kortsmit, Y., Sklavounou, M., Dicke, M., & van Loon, J. J. A. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10342 · Cited by 2 · cited 10× · see citing works
“As the size difference between females and males increases, so does the degree of SBM (Teder et al. 2021; Kortsmit et al. 2025).”
InfluentialA chemically defined oviposition attractant and repellent of black soldier flies (Hermetia illucens)
Thomas, N. K., Karpati, Z., Schmitt, T., & Riabinina, O. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10312 · Cited by 2 · see citing works
Thomas, N. K., Karpati, Z., Schmitt, T., & Riabinina, O. (2024) · doi:10.1101/2024.06.04.597456 · Cited by 2
Rollinson, L. N., Lemke, N. B., Crawford, S., Woolley, J. B., & Tomberlin, J. K. (2026) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10414 · Cited by 2 · see citing works
Self-citationBarrett, M., Godfrey, R. K., Sterner, E. J., et al. (2022) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2022.01.25.477588 · Cited by 2 · see citing works
Yaseen, Y., Ullah, A., Khan, I. R., Murad, W., Taj, R., Ahmad, M., Alrefaei, A. F., Ali, S., & Hamayun, M. (2024) · Polish Journal of Environmental Studies · doi:10.15244/pjoes/187594 · Cited by 1 · see citing works
Dearlove, E., Van Gestel, C. A. M., Loureiro, S., Svendsen, C., Lloyd, M., Mugo-Kamiri, L., Petersen, J. M., Bessette, E., Edwards, S., Lim, F. S., et al. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10203 · Cited by 1 · see citing works
Rollinson, L. N., Lemke, N. B., Crawford, S., Woolley, J. B., & Tomberlin, J. K. (2025) · doi:10.1101/2025.08.07.666420 · Cited by 1 · cited 3× · see citing works
“However, a recent study shows that male BSF genitalia are hypo-allometric (Rollinson et al. 2025), allowing males to mate with females across a wide range of body sizes and increase their pool of available mates (Eberhard et al. 1998), and thus is unlikely to be a physical barrier to reproduction.”
InfluentialSelf-citationGarcía‐Castillo, P., Peralta, C., Hernández‐Pelegrín, L., & Herrero, S. (2025) · eFood · doi:10.1002/efd2.70110 · Cited by 1 · see citing works
Muchina, P., Kinyua, J. K., Khamis, F. M., Tanga, C. M., Satti, M. A., Gebreyesus, G., Sahana, G., & Cai, Z. (2026) · Genomics · doi:10.1016/j.ygeno.2026.111206 · Cited by 1 · see citing works
Lemke, N. B., Li, C., & De Smet, J. (2026) · Journal of Economic Entomology · doi:10.1093/jee/toag117 · Cited by 1 · see citing works
Self-citationZhang, Y., Henawy, A. R., Rehman, K. U., van Huis, A., Cai, M., Zheng, L., Huang, F., Ding, X., Lei, H., & Zhang, J. (2026) · Insect Science · doi:10.1111/1744-7917.70064 · Cited by 1 · see citing works
Male and female effects on sperm precedence in the giant sperm species Drosophila bifurca.
Luck, N., Dejonghe, B., Fruchard, S., Huguenin, S., & Joly, D. (2007) · Zenodo · doi:10.5281/zenodo.10755761 · Cited by 1 · see citing works
Cui, Z. (2022) · Zenodo (CERN European Organization for Nuclear Research) · doi:10.5281/zenodo.6497038 · Cited by 1 · see citing works
Julita, U., Permana, A. D., Supriatna, A., Hambali, A., & Fitri, L. L. (2025) · Journal of the Entomological Research Society · doi:10.51963/jers.v27i1.2684 · Cited by 1 · see citing works
Microbial influences on black soldier fly reproduction: A focus on egg surface colonization
Heussler, C. D., Klammsteiner, T., Stonig, K. T., Insam, H., Schlick-Steiner, B. C., & Steiner, F. M. (2023) · doi:10.21203/rs.3.rs-3314156/v1 · Cited by 1 · see citing works
Biomimicry in love-cage design: boosting black soldier fly mass production
Grosso, F., Lattarulo, A., Meneguz, M., & Padula, C. (2025) · Journal of Insects as Food and Feed · doi:10.1163/23524588-bja10248
Lemke, N. B., Li, C., & De Smet, J. (2025) · doi:10.1101/2025.08.26.672371
Self-citationMuchina, P., Kinyua, J. D. M., Khamis, F. M., Tanga, C. M., Satti, M. A., Gebreyesus, G., Sahana, G., & Cai, Z. (2025) · SSRN Electronic Journal · doi:10.2139/ssrn.5867966
Nitrogen Journey in Plants: from Uptake to Metabolism to Stress response and Microbe Interaction
Zayed, O., Hewedy, O. A., Abdelmoteleb, A., et al. (2023) · doi:10.20944/preprints202308.1575.v1
Dearlove, E., Gestel, C. A. M. V., Loureiro, S., et al. (2026) · BugBook · doi:10.1163/9789004760790_014
Antennal Olfactory Responses in the Black Soldier Fly Hermetia Illucens
Piersanti, S., Rebora, M., Carboni-Marri, G., et al. (2024) · doi:10.2139/ssrn.4855898
Silvaraju, S., Zhang, Q.-H., Kittelmann, S., & Puniamoorthy, N. (2024) · doi:10.21203/rs.3.rs-4661186/v1
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Cited by
Sorted by citations
McKenzie, M. J., Liu, Z., Tran, A. K. N., Botero, J., Edmonds, R. L., Hermansen, C., Fabisik, F., Fletcher, K. M., & Najar‐Rodriguez, A. (2026) · Waste Management & Research The Journal for a Sustainable Circular Economy · doi:10.1177/0734242x261468120
Piersanti, S., Petroni, G., Marri, G. C., et al. (2026) · Journal of Insect Physiology · doi:10.1016/j.jinsphys.2026.105059
- Authors
- Lemke, N. B., & Puniamoorthy, N.
- Affiliations
- KU Leuven (BE)
- National University of Singapore (SG)
- Corresponding author
- Lemke, Noah B.
- Puniamoorthy, Nalini
- Venue
- Evolutionary Applications
- Publisher
- Wiley
- Published
- 2026-05
- Volume
- 19
- Issue
- 5
- Pages
- e70249-e70249
- ISSN
- 1752-4563 (ISSN-L)
- 1752-4571
- PMC
- PMC13156471
- OpenAlex
- W4413865651
- Semantic Scholar
- f5bcb5eceab5b8e1b1abee2b9f6b5ef61daa506d
- Cited by
- 2
- References
- 165
- Citations by year
- 2026: 2
- FWCI
- 16.20
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Licence
- Locations
- Evolutionary Applications (publishedVersion, cc-by)
- Preprints.org (acceptedVersion, cc-by)
- PubMed (publishedVersion)
- PubMed Central (submittedVersion, cc-by)
- Funders
- Fulbright Association
- National Research Foundation Singapore
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Animal Behavior and Reproduction
- Life Sciences › Agricultural and Biological Sciences › Insect Science › Insect Utilization and Effects
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Genetics › Insect and Arachnid Ecology and Behavior
- Keywords
- Hermetia illucens, reproductive biology, sexual selection, lek mating system, protandry, sperm storage, sperm precedence, mating behavior, oviposition, insect biotechnology, chemical ecology, environmental effects on reproduction
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