Cited by 2Animal Behavior and ReproductionInsect Utilization and EffectsInsect and Arachnid Ecology and Behavior

Beyond the Black Box: Reproductive Strategies of the Black Soldier Fly as a Model for Bridging Evolutionary Biology and Applied Entomology

Open access
Lemke, Noah B., Puniamoorthy, Nalini·2026·Evolutionary Applications

Abstract

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.

AI snapshot

Written from the paper’s full text.

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.

Integrity signals

Registry

Crossref · 10.1111/eva.70249

Cites

199 of 199 references verified0 citation errors · 0 not checkable

Statistics

Nothing to recompute.

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
  • 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. 8

    Several 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. 12

    Increased 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. 5

    Female 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. 4

    BSF 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 checked
    • cites Thinn and Kainoh 2022 source
    • 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
  • May combine two claims.

    Premisecitation
    § 2.1.1   |   Latency and Sexual Maturation · p. 4

    Neurobehavioural 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. 1

    BSF 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. 5

    Male 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. 6

    Visual 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. 5

    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.

    “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. 6

    BSF'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. 7

    BSF 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. 6

    Males 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. 7

    The 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 checked
    • cites Manas, Labrousse, and Bressac 2025 source
    • cites Manas, Venon, et al. 2025 source
  • May combine two claims.

    Findingcitation
    § 2.2.1   |   Copulatory Mechanics, Duration and Sperm Transfer · p. 7

    Initial 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. 6

    Some 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. 6

    The 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. 6

    Some 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. 6

    Lekking 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. 6

    More 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 checked
    • cites Barbosa 2011 source
    • cites Alcock 1990 source
  • May combine two claims.

    Findingcitation
    § 2.3.1   |   Oviposition · p. 8

    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.

    “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. 8

    Delaying 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. 8

    For 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. 8

    Oviposition 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. 7

    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.

    “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. 9

    Seminal 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 checked
    • cites Harjoko et al. 2023 source
    • cites Manas 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
  • May combine two claims.

    Findingcitation
    § BOX 4 |    Reproductive skews in BSF. · p. 7

    Mating 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. 7

    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.

    “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. 8

    Fecundity 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. 8

    The 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. 7

    The 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. 8

    These 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. 7

    A 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. 10

    Intentional 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. 10

    The 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 checked
    • cites Zeh and Zeh 1997

      Couldn’t be matched to a paper in Crossref, so it can’t be checked.

    • cites Laturney et al. 2018 source
    • cites Mack et al. 2003 source
  • May combine two claims.

    Findingcitation
    § 3.1   |   Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10

    Sperm 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 checked
    • cites Malawey et al. 2019 source
    • cites Malawey et al. 2020 source
    • cites Zhang and Puniamoorthy 2025 source

      Split into 2 atoms — 0 confirmed, 0 partially supported, 0 disputed, 0 misattributed, 2 unverifiable. Expand the claim to see per-atom verdicts.

      Experimental — accuracy not yet measured
  • Classified as probably not a substantial claim. May combine two claims.

    Interpretationreasoning
    § 3   |   Implications for Evolutionary Biology · p. 10

    This 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. 10

    Recent 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. 10

    Phenotypic 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. 9

    In 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. 11

    Continuous 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. 11

    BSF 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. 11

    Another 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. 12

    Adult 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. 12

    Providing 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. 12

    Spatial 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. 12

    There 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. 12

    BSF 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. 6

    In 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. 7

    The 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 checked
    • cites Giunti et al. 2018 source
    • cites Laudani et al. 2024 source
    • cites Kortsmit et al. 2025 source
  • May combine two claims.

    Findingcitation
    § 2.2.1   |   Copulatory Mechanics, Duration and Sperm Transfer · p. 7

    Sperm 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. 7

    After 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 checked
    • cites Giunti et al. 2018 source
    • cites Laudani et al. 2024 source
  • Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.

    Findingcitation
    § BOX 3 |    Debates on lekking in BSF. · p. 6

    There 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. 7

    Overall 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. 8

    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.

    “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. 8

    Examination 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. 8

    Male 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 checked
    • cites Chiabotto et al. 2024 source
    • cites Laudani et al. 2024 source
    • cites Jensen et al. 2025 source
  • Probably 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. 8

    Observations 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 claim
  • May combine two claims.

    Findingcitation
    § BOX 6 |    (Continued) · p. 9

    Older 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. 8

    Off-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. 9

    BSF 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. 10

    Laboratory 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. 9

    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.

    “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 checked
    • cites Pitnick et al. 1995 source
    • cites Vahed 1998 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Findingcitation
    § 3.1   |   Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10

    Insects 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. 10

    In 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. 10

    BSF 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 checked
    • cites Jensen et al. 2025 source
    • cites Chiabotto et al. 2024 source
  • May combine two claims.

    Findingnone given
    § BOX 6 |    (Continued) · p. 9

    Female 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 claim
  • May combine two claims.

    Findingcitation
    § 3.1   |   Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10

    Males 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. 10

    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.

    “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. 2

    Adult 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. 2

    The 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. 2

    BSF 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. 4

    The 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. 6

    Acoustic 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 claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § BOX 2 |    Debates on mate discrimination in BSF. · p. 6

    BSF 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. 2

    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.

    “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 claim
  • May combine two claims.

    Premisereasoning
    § BOX 1 |    What makes BSF a model organism in the classic sense? · p. 3

    In 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 claim
  • May combine two claims.

    Interpretationreasoning
    § WILEY · p. 3

    BSF 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 claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § 2.1.1   |   Latency and Sexual Maturation · p. 4

    Protandry 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. 3

    Female BSF carry between 300 and 900 eggs.

    “Females carry between 300 and 900 eggs”
    Quote in the paperNot a substantial claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Interpretationcitation
    § 2.1.2 | Navigation and Lekking · p. 5

    In 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. 2

    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.

    “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 claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § 2.1.1   |   Latency and Sexual Maturation · p. 4

    Protandry 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. 5

    Temporal 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. 6

    Lekking 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. 6

    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.

    “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. 6

    The 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 checked
    • cites Emlen and Oring 1977 source
    • cites Parker 1978 source
    • cites Thornhill and Alcock 1983 source
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § BOX 3 |    Debates on lekking in BSF. · p. 6

    Each 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. 7

    Mating '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. 7

    Although 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. 8

    These 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 claim
  • May combine two claims.

    Interpretationreasoning
    § BOX 4 |    Reproductive skews in BSF. · p. 7

    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.

    “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 claim
  • Probably not a substantial claim: may not pass S3 (load-bearing).

    Premisecitation
    § 3.1   |   Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10

    Males 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. 10

    Females 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 claim
  • May combine two claims.

    Interpretationcitation
    § 3   |   Implications for Evolutionary Biology · p. 9

    This 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. 9

    Multiple 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. 10

    Male 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 checked
    • cites Pitnick and García-González 2002 source
    • cites Fowler and Partridge 1989 source
    • cites Morrow and Gage 2001 source
  • May combine two claims.

    Premisecitation
    § 3.1   |   Cryptic Female Choice, Sperm Competition and Sexual Conflict · p. 10

    Often, 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. 11

    Male 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. 11

    Domesticated 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. 11

    Selective 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. 11

    Mating 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 claim
  • May combine two claims.

    Interpretationcitation
    § 4.3   |   Sex Ratios and Habitat Design · p. 12

    Cage 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. 14

    BSF 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 claim
  • May combine two claims.

    Premisenone given
    § 4.4   |   Chemical Ecology and Oviposition Control · p. 12

    Some 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
104 of 104
  • 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”
11 of 11

Cites

Flagged first, then by citations

Research works printed but not linked: 23

  1. Athanassiou C. G., 2024, A Decade of Advances in Black Soldier Fly Research: From Genetics to Sustainability
  2. Briceño R. D., 1999, Aggressive Behavior in Medflies (Ceratitis capitata) and Its Modification by Mass Rearing (Diptera: Tephritidae), Journal of the Kansas Entomological Society, 72, 17
  3. Dortmans B., 2017, Swiss Fedearl Institute of Aquatic Science and Technology (EAWAG): Department of Sanitation, Water and Solid Waste for Development (Sandec)
  4. Ferdousi L., 2021, Impact of Daylight Duration on Pupation and Adult Emergence of Black Soldier Fly (Hermetia illucens), International Journal of Entomology Research, 6, 142
  5. Freitas Spindola A., 2019, Morphological and Physiological Reproductive Aspects of Black Soldier Fly: Applications for Optimal Industrial Scale Production
  6. Generalovic T. N., 2023, Population Genomics and Domestication of the Black Soldier Fly (Hermetia illucens L.)
  7. 10.1002/efd2.70110 · DOI
  8. Generalovic T. N. C.Sandrock B. J.Roberts et al.2023.“Cryptic Diversity and Signatures of Domestication in the Black Soldier Fly (Hermetia illucens).”bioRxiv. Accessed August 11 2025.https://doi.org/10.1101/2023.10.21.563413v2. · DOI
  9. Herberstein M. E., 2017, Advances in the Study of Behavior, 237
  10. James M. T., 1935, The Genus Hermetia in the United States (Diptera: Stratiomyidae), Bulletin of the Brooklyn Entomological Society, 30, 165
  11. Jiggins C. D.2024.“Genetic Diversity and Improvement of the Black Soldier Fly.”Theatre.https://doi.org/10.1163/23524588‐20241013. · DOI
  12. Kohler R. E., 1994, Lords of the Fly: Drosophila Genetics and the Experimental Life
  13. Michiels N. K., 1989, Differences in Male and Female Activity Patterns in the Dragonfly Sympetrum Danae (Sulzer) and Their Relation to Mate‐Finding (Anisoptera: Libellulidae), Odonatologica, 18, 349
  14. Miller W. E., 2005, Reproductive Bulk in Capital‐Breeding Lepidoptera, Journal of the Lepidopterists' Society, 59, 228
  15. Park K.‐H., 2017, Oviposition Activity of Black Soldier Fly (Hermetia illucens) Under Artificial Illumination, International Journal of Industrial Entomology, 35, 100
  16. Sandrock C., 2021, Book of Abstracts of the 72nd Annual Meeting of the European Federation of Animal Sciences. Davos, Switzerland, 295
  17. Sibonje J. N., 2024, Determination of the Gross Nutrient Composition of Larvae and Substrate During Black Soldier Fly Treatment of Faecal and Kitchen Waste
  18. Sripontan Y., 2017, Egg‐Trapping of Black Soldier Fly, Hermetia illucens (L.) (Diptera: Stratiomyidae) With Various Wastes and the Effects of Environmental Factors on Egg‐Laying, Khon Kaen Agricultural Journal, 45, 179
  19. Tekaat L., 2024, Maternal Oviposition Instincts vs. Their Influence on Developmental Outcomes: Life‐History Analyses of the Black Soldier Fly
  20. Tomberlin J. K., 2025, BugBook: Black Soldier Fly as a Model to Assess Behaviour of Insects Mass Produced as Food and Feed
  21. Wilson E. O., 1999, The Diversity of Life
  22. Yeates D. K., 2005, The Evolutionary Biology of Flies
  23. Zeh J. A., 1997, Last‐Male Sperm Precedence Breaks Down When Females Mate With Three Males, Proceedings of the Royal Society of London. Series B: Biological Sciences, 257, 287

Cited by

Sorted by citations

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
OpenAlex
W4413865651
Cited by
2
References
165
Citations by year
2026: 2
FWCI
16.20
Type
journal article
Language
en
Open access
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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Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.

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Classified by jev-1.13-20260917 (gold-guideline-v0.2) on 2026-10-11.

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