ESI-VOL.84 (3) 2022 - Anshika Singh 743-753
ESI-VOL.84 (3) 2022 - Anshika Singh 743-753
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ABSTRACT
Endeavours to recycle organic waste by utilizing black soldier fly (BSF) Hermetia illucens (L.) into waste
management and high nutrient biomass development have indeed picked up momentum recently. But
there is not much data on their appropriate rearing conditions. Very few studies delineating the fecundity
and reproduction capacities of BSFs concluded that the presence of natural sunlight or artificial light with
intensity of 110-200 μ mol m2s-1 and above triggers mating and oviposition (about 85-90%) at successfully
higher rates along with ideal temperature (26 to 40°C) and relative humidity (40-70%) conditions. Optimum
food moisture (50-80%) also plays a vital role in enhancing the consumption rate of waste and therefore the
treatment efficiency of the larvae. Significant development of the BSF larvae and the treatment efficiencies
were also observed to be governed by the pH of waste and the optimum range was defined to be of 6.0 to
9.0. The type of organic waste also equally influences the development, fecundity, and the lifespan of flies.
The present review highlights the significant research that has been conducted with respect to lifecycle of
BSFs, under the set of different light combinations (LED and fluorescent lights), temperatures and organic
waste composites (protein rich and fat rich substrates). Conclusively, it was inferred that ameliorations
in rearing conditions such as investigation of suitable light source, modifications in egg collection and
hatching structures and knowledge of biology of flies can further boost the reproductive capability of fly
thereby promoting successful insect rearing and mass production.
Key words: Hermetia illucens, mating, environmental factors, oviposition, lifecycle, mass rearing, organic
waste, development time, survival rate, adult weight, life-history traits
Form the past few decades, black soldier fly (BSF) BSFs. In general, insects are ubiquitously distributed
Hermetia illucens (L.) larvae have played a vital role in in environment based on their degree of tolerance,
treatment of organic waste such as food waste, abattoir adaptivity and physiology (Dixon et al., 2009). Certain
waste, animal, and human faeces, etc. embracing the set of environmental conditions such as temperature,
prospects of circular economy worldwide. The larvae relative humidity, moisture, sunlight, and aeration
are also regarded as the impeccable source of proteins, are crucial factors for successful copulations among
lipids and carbohydrates supplementing the animal feed individuals of opposite sex, egg laying activities and
crisis (Tomberlin et al., 2009). Apart from this, bioactive successful mass production (Jarosik et al., 2004; Singh
compounds, degrading enzymes and antimicrobial and Kumari, 2019). One such study stated that regions
peptides secretions within the species offers a with wider temperature profile is highly favourable for
tremendous outlook from industrial point of view. The insect development encompassing higher population
larval extracts have now become of great interest as comparative to the regions with smaller temperature
they make the insect mass rearing economically sound profiles (Addo-Bediako et al., 2000; Deutsch et al.,
because of their inherent distinctive properties and pave 2008).
the ways for development of antimicrobial compounds
(Muller et al., 2017). Promoting the excellent breeding Black soldier flies [BSF-Hermetia illucens (L.)] has
activity among species to recover huge quantity of eggs the capability to thrive even under hostile conditions
and thus the larvae are the most troublesome task to (Diener et al., 2011a) and has a typical life cycle
manage. Therefore, rearing the species demands the of 1- 2 months. The basic anatomy of adult fly and
utmost care and improved technical skill especially the larvae is shown in Fig. 1. The larval stage is the
dealing with the oviposition and mating events in the only feeding stage wherein they feed voraciously on
744 Indian Journal of Entomology 84(3) 2022 Review
its utilization for organic waste treatment and other unnecessary moisture levels (more than the optimum
societal benefits. levels) in the feeding substrates can impact feeding rate
and might result in the development of thick and clumpy
1. Environmental factors material causing trouble in further handling (Diener et
Mating among opposite sexes of BSFs is strongly al., 2011b). Another group of researchers have reasoned
under the control of surrounding temperature, substrate that 80% moisture content in the feed is ideal for BSF
properties and moisture content. Not just this, the natural development (Cheng et al., 2017; Dortmans et al.,
conditions (relative humidity, sunlight) contribute 2017). Similarly, Barragan-Fonseca and her co-workers
towards ideal egg laying and incubation events and the defined that the moisture level in between 52-70% are
larval development. Besides that, substrate selection the most suitable conditions for proper larval growth
also equally contributes to the successful eggs laying and development (Barragan-Fonseca et al., 2017).
and hatching activities since the females prefers to lay Comparable results were likewise referenced in the
eggs near to the food source with strong putrescence investigation of De Smet et al. (2018) where a moisture
(Tomberlin and Sheppard, 2001; 2002). level underneath 40% was unfit for development of the
flies at various stages therefore influencing the fecundity
Ideal thermal conditions (temperature) and the rate or the mating in the files.
relative humidity: Earlier studies have concluded
that about 99% mating and oviposition occurs in the Optimum light exposure and intensities: In
temperature range of 27.5 to 37.5°C combined with general, different parts of an insect’s eye have
60% relative humidity (Sheppard et al., 2002; Holmes, different spectral sensitivity and the spectral sensitivity
2010). In a similar setting, 50-90% relative humidity functioning may differ among species. The ommatidia
has been defined as the ideal condition for enhancement are the structural unit of insect’s compound eyes
of BSF rearing at research centres in temperate regions and have photoreceptor cells arranged in different
according to different group of others (Diener et al., fashion. Most of the insect species have the light
2009). So also, Tomberlin et al. (2009) found significant sensitivity lying in the range <300 to >700 nm. They
development of males and females at 27°C± 2 however don’t see the light past 700 nm, a light source with
at higher temperatures (30-36°C) smaller males and corresponding wavelength between 450-700 nm is
females were observed in the study and life expectancy ideal for reproductive activities in adult flies (Briscoe
was also decreased. On the other hand, it was noticed and Chittka, 2001; Zhang et al., 2010). The visual
that temperature and humidity can have genuine impacts pigment or photoreceptors in insect eyes is composed
on egg eclosion and development at different stages of chromophore and opsin protein which interacts
if not looked after ideally (Park, 2016). Holmes et al. with light sensitivity of shorter or longer wavelengths
(2012) additionally reasoned that less than 25% relative (Peitsch et al., 1992; Stavenga, 1992; Cronin et al.,
humidity can have higher parching rates and higher 2000). Similarly, BSFs have photoreceptors belonging
mortality of species. In the context of eggs hatching, to a specific class which is highly sensitive to UV light
the lower humidity levels cause moisture loss from egg (367 nm) and blue light (440nm). Ventral part of eye was
membrane leading to desiccation. A humidity level as maximally sensitive to blue light whereas dorsal retina
low as 25% results in higher desiccation and mortality was sensitive to blue green (504 nm) and UV light has
rates whereas at 70% and above, adults live longer, additional peak in both ventral (40%) and dorsal retina
and the eggs retains the proper health conditions. More (20%) (Fig. 2) (Oonincx et al., 2016).
precisely, relative humidity between 70-90% provides
the absolute conditions for mating and oviposition For successful mating in BSF, direct daylight is
activities mediated by the adults in combination with assumed to be significant in the common habitat and
27°C temperature of the surrounding air (Holmes et al., this is the explanation that huge mating doesn’t happen
2012; Park, 2016). Chia et al. (2018) also stated in his in winter seasons however creating a similar warm
study, that presence of optimum temperature conditions environment may promote high mating activities.
improves the fertility and fecundity rate of adult flies. Studies considering the indoor investigations on BSF
require artificial light source. About 85% mating events
Substrate moisture levels: The development and happen within the sight of common daylight with an
endurance of the species is exceptionally affected intensity of 110 µmol m2s-1 however the mating rates
by the moisture of the feed (Cheng et al., 2017). decreases below that (Park, 2016). Previously, Briscoe
Numerous researchers have peculiarly identified that the and Chittka (2001) also stated that a 500-watt quartz
746 Indian Journal of Entomology 84(3) 2022 Review
iodide lamp with an intensity as high as 135 µmol is preferred by the BSF unlike the other insects such as
m2s-1 was able to promote the mating and oviposition mealworms (Tomberlin et al., 2002). Similarly, the high
at rates similar to those observed in the presence of protein or highly fibrous food often lowers the larval or
natural sunlight. Similarly, in another study, artificial prepupal growths, the mating behaviour, and the eggs
light source was accounted to have effect on mating resilience (Tschirner and Shimon, 2015; De Smet et al.,
wherein the mating and oviposition of the species 2018). Regardless of that, the higher larval development
was associated with the utilization of Quartz-iodide was likewise seen in the neonates fed with plant-based
light (61% efficiency) with the same intensities as the substrate when contrasted with animal-based diet
counterfeit light source as compared to natural sunlight since they harbour microorganisms delivering plant
and rare earth lamps (Zhang et al., 2010). This specific processing enzymes (Liu et al., 2008; Tomberlin et al.,
strategy can be extremely worthwhile and compelling 2009; Manyara, 2018). In one such study of Chia et al.
for raising the species outside their local natural (2018), considering the mating and eggs production, it
surroundings, where daylight is the principle affecting was found that heavier prepupae were obtained in the
source. Besides that, Tomberlin and Sheppard (2001) treatment where the larval were fed with Brewer’s spent
have also reported that lekking behaviour among male grain supplemented with Brewer’s yeast (nutritionally
and females is restricted by the type of habitats, for balanced diet).
instance, males aggregate near bushy areas establishing
the territory which serve as the attractant for females and Danieli et al. (2019) also suggested that a mix of
initiating the mating. Notwithstanding, to have better different by-products of food such as alfalfa, barley,
comprehension of science of species affected by light and wheat by products are remarkable source of feed for
and other factors as discussed above, extra research is enhanced larval development enhancing the nutritional
required to significantly promote the mass scale rearing accumulation and survivorship of the BSFs. He also
of the insect (BSF). added, diet should be fat enriched instead of protein or
carbohydrate rich substrates because the latter two can
Effect of substrate: The substrate (diet) composition negatively affect the rearing and overall development.
has been extensively studied worldwide to raise BSF In addition, Ewusie and his co-workers, also determined
however due to their diversity to eat any kind of waste that the type of substrate affects the trapping size of
(food waste, poultry waste, animal faeces, human egg clutches of BSF in combination with the other
faeces, animal waste, etc.) such problems are not usually environmental factors. In their study, among the piggery,
encountered (Boykin, 2019). But still, the substrate sheep, and poultry waste, the piggery waste was the
composition is equally influential for the healthy profile most influential (Ewusie et al., 2019).
(nutritional aspects) and development of life history
traits of BSF (Sheppard et al., 2002; Tomberlin et al., pH: A concern for pH effect on BSF development
2002; Boykin, 2019) because it affects the feeding and survival has been also raised by Green and Popa
rate and conversion, the gut loading and digestibility (2012). Larvae were found to regulate a pH of 9.0
(Fig. 3). For instance, larvae prefer to consume food occurring in leachates or other substrates (Green and
with high fat content (Nguyen et al., 2015). Moist diet Popa, 2012). Similarly, the increased larvae growth
Black soldier fly Hermetia illucens (L.): Ideal environmental conditions and rearing strategies 747
Anshika Singh et al.
and heavier pupal mass can be positively correlated and substrate moisture) on different life history traits
with substrates having pH in between 6.0 to 10.0 of BSF in which the baiting material used for trapping
however; a greatly reduced development is seen at pH adult flies included a mix of manures of chicken and
below 2.0 (Ma et al., 2018). In one such recent study, rabbits, food and household wastes. The development
in which the researchers examined the effect of pH and was monitored for different array of temperatures (10
the feeding system on BSF larvae found that initially to 42 °C) combined with relative humidity between
for a week pH significantly affected the larval weight 70 to 72% and 12:12: Light: Dark photoperiod. The
however at the end it was same at all pH between 6 to population growth rate was highest in treatments as
9 under the environmental conditions of 29.3± 1.4°C compared to controls (without Brewer’s yeast). The
temperature and 70.0± 5.0% relative humidity. In fact, authors concluded that the number of successful
the larval feeding activity also modifies the pH of the mating and fecundity rate can also be correlated with
feed in between 8.9 to 9.4. But, larval and pupal weight, pupal mass of insects. The temperatures below 15°C
pupation rate, sex ratio survival and mortality rates were and above 40°C was found to negatively impact the
majorly influenced by the feeding system (Meneguz et survival of the fly at all stages resulting in extremely
al., 2018). high mortality rates. The findings of Chia et al. (2018)
have been shown in Table 1. Likewise, Shomu et al.
2. Rearing (2019a; 2019b) also published work closely related
Effect of temperatures and substrate types: to the study of Chia et al. wherein they examined the
Upkeep of appropriate ecological conditions has always effect of temperature and substrate type (brewer’s spent
been the chief significant thing to effectively manage the grain and cow dung) on the growth, development, and
mass rearing activities of BSF. In the similar context, survival of BSFs and mating rates in adults. The findings
researchers are trying their best to introduce best suitable stated that the development was faster for the larvae fed
conditions to manage the species to offer secondary with brewer’s spent grains as compared to cow dung
solution for the public and the concerned people i.e., and the optimum range of temperature suitable for BSF
the waste management and its valorisation. One of was 25-30°C. Srikanth and Sharanabasappa reported
the more recent study examined the effect of different that kitchen waste was significantly superior obseved
ecological factor (i.e., temperature, relative humidity, with maximum larval (0.22g/ larva) and pupal weight
Table 1. Effect of temperature on life history traits of H. illucens (Chia et al., 2018)
S. Light history traits Temperature Days Survival rate (%)
No.
1. Egg eclosion 15°C 14 (both D1 and D2) Highest at 35°C (75% and 30°C
(80%) for both D1 and D2
35°C 2.60(both D1 and D2) -
2. Larval development 15°C 13.14 (both D1 and D2) -
30°C 62.4 (both D1 and D2) -
3. Prepupal development 15°C 83-86 (both D1 and D2) -
(Failed at 40°C) 30°C 8-10 (both D1 and D2) -
4. Larval to adult 15°C 184 (D1); 181(D2) -
development 30°C 28 (D1); 31(D2) -
5. Oviposition period 20°C 16 (both D1 and D2) -
35°C 5 (both D1 and D2) -
6. Larval survival rate - - 92% (35°C); 90% (30°C); 28%
(40°C) for both D1 and D2
7. Prepupal survival rate - - 83% (25°C) and 82% (30°C) for
D1; 79% (35%) and 77% (30°C)
for D2
8. Pupal survival rate - - 77% (30°C) and 5% (37°C) for
D1; 75% (30°C) and 20% (37°C)
9. Adult survival rate Decreased with increase in temperature from 15°C to 37°C with increased
fecundity at higher temperatures
D1: BSGs with brewer’s yeast; D2: BSGs without brewer’s yeast (control)
748 Indian Journal of Entomology 84(3) 2022 Review
(0.20g/ pupa). The Fecundity was observed to be 698 Diener et al. (2011b), Gobbi et al. (2013), Nguyen et
eggs/ female when reared up to 14 day, significantly al. (2013), Li (2014), Oonincx et al. (2015a, b) and
the maximum amount of feed consumed was 18100g Srikanth and Sharanabasappa, (2021) have affirmed
in 20days, feed conversion ratio (97.37) was maximum the BSFs against various natural substrate to assess
in mixed vegetable waste. diverse life stages. Table 2 has compiled the studies on
life history traits of BSF fed against variety of organic
The selection of substrate should be wise as it effects diets having different nutritional contents which was
both the physiological and morphological development originally combined in the studies of Barragan-Fonseca
(gonads development, sex ratio, mortality, duration of et al. (2017) from the studies executed up until now. The
different stages) of both neonates and the adult flies. abiotic conditions were as follows: temperature- 27±
Gobbi et al. (2013) testified the insect’s development 2°C; relative humidity-70± 10%; food moisture- 66±
against three different diets i.e., hen feed, fish meal and a 4%. Interestingly, a conclusion was also drawn that the
mixture of both. The larvae fed with fish meal and mixed availability of abundant food positively affects the larval
diet showed the prominent growth and development growth and development however, the waste reduction
than the larvae fed with pure meat meal (Gobbi et efficiency is greatly reduced (Liu et al., 2008; Diener
al., 2013). Notably, till date various examinations et al., 2009; Banks, 2014).
have been set up testing the effect of various sorts of
organic waste on life history attributes of BSF and Harden and Tomberlin (2016) also deeply
their performances, for example, larval growth and investigated the synergistic effect of temperature and
development, pupal development, adult weights and diet on BSF development. A mix of grains (corn meal,
lengths, longevity, and the endurance. Similar in case, wheat bran and alfa alfa), beef and pork meet were used
the investigations of Tomberlin and Sheppard (2001), as the diet source and the temperatures conditions were
Table 2. Life history traits and performance of H. illucens larvae fed with organic waste
S. Life history traits Organic waste
No. (Mean± SD) Chicken feed Meat waste Faeces Vegetable
(or feed with waste
similar nutrient
content i.e.,
Protein~14%;
Fat~4%)
Larval stage
1. Development time (days) 24.6± 6.2 32.5± 8.2 27.5± 3.8 34± 13.5
2. Survival rate (%) 89.4± 9.4 48.2± 8.7 89± 7.5 78.9± 13.2
3. Fresh matter (FM) weight (g) 0.158± 0.02 0.158± 0.0 0.17± 0.03 0.13± 0.03
4. Dry matter (DM) weight (g) .044± 0.0 - 0.031± 0.02 0.028± 0.01
5. Larval dry matter (DM) 36± 1.8 - - 36.3± 2.5
content (%)
Prepupal and pupal stage
6. Pupal development time (days) 14.8± 6.8 16.5± 7.5 17.8± 3 22.9± 1.2
7. Fresh matter (FM) prepupal 0.105± 0.005 0.115± 0.01 0.193± 0.08 0.179± 0.03
weight (g)
8. Dry matter (DM) prepupal 0.037± 0.004 - 0.018; 0.04 0.071± 0.01
weight (g)
9. Fresh matter (FM) pupal 0.150± 0.03 - - -
weight (g)
Adult stage
10. Adult weight (g DM) 0.021± 0.0 - - -
11. Adult weight (g FM) 0.053± 0.01 - 0.046± 0.01 -
12. Adult length (mm) 15.8± 0.0 - - -
13. Adult longevity (days) 9.4± 0.2 - 12.5± 2.1 -
14. Total cycle (days) 40.2± 6.4 - - -
Black soldier fly Hermetia illucens (L.): Ideal environmental conditions and rearing strategies 749
Anshika Singh et al.
between 29-33°C and average relative humidity was natural sunlight however the number of fertilized eggs
71.0± 16.3%. They found that eggs development was obtained were higher under the effect of natural sunlight
not significantly affected with variations in temperature. in comparison to LED illumination. The oviposition was
However, larval length and development was affected considerably increased from 4 to 17 days. Examining
notably with temporal variations and was highest at the sugar solution treatment and water treatment it
32.2 and 27.6°C. Moreover, about 23.1%- and 139.7% was observed that the longevity of adults (3 times in
more-degree hours were required to complete the males and 2 times in females) increased when fed with
larval development fed with pork diet in comparison sugar solution comparative to water alone (Nakamura
to beef and grain-based diet. Comparatively, larval et al., 2016). The findings also concluded the absence
development fed with grain diet was inconsistent when of mating below 69 μmol m-2 s-1 and highest at 200
assessed with the age of field larvae and the pork and μmol m-2 s-1.
beef based larval development were appreciable.
Heussler et al. (2018) scrutinized the impact of
Effect of light: Oonincx et al. (2016) and Nakamura three different artificial light sources (LED green,
et al. (2016), studied LED expand based radiance on fluorescent lamp, and halogen lamp) on the life history
indoor reproduction rates and the oviposition periods. traits (oviposition and half-life) and mass production of
The former study included the photoreceptor spectral BSF where the larvae were reared in a plastic box under
sensitivity of the compound eyes of BSF while the latter the environmental conditions at 27°C temperature and
investigated the oviposition and survival rates under 60% relative humidity respectively. He also encountered
natural sunlight supplemented with LED irradiation. the similar observations for oviposition as in above-
Oonincx et al. (2016) found that the ommatidia of mentioned studies. Three replications were made for
BSF contains photoreceptors cells sensitive to blue, each light source with intensity of 59 μmol m-2 s-1 in
green and UV light having trichromatic visibility; different cages having a light: dark photoperiod of 16
therefore, the LEDs based illumination significantly and 8 hours. Mating and oviposition rates were found
increased the egg clutch yields resulting in higher similar in all the three conditions where the peak was
larval production comparative to fluorescent tubes observed from 4 to 8 days of emergence however
(control) (Figs. 4, 5). Similarly, small cage (27 x 27 x the half-life of both males and females significantly
27 cm) study of Nakamura et al. containing 100 adult reduced. Shorter half-lives of adults were observed
flies (50 male and female each) in each implied similar under halogen light conditions as compared to LED
patterns of oviposition in both LED illumination and illumination lasting from 6 to15 days for males and
Increased
adult
survival
a. BSF compound eye sensitivity for b. Effect of LED illumination over control
LED illumination (fluorescent light)
Fig. 4. Diagrammatic representation of photosensitivity of H. illucens and effect of LED illumination on
adult survival and larval development (Oonincx et al. 2016)
Fig. 4. Diagrammatic representation of photosensitivity of H. illucens and effect Fig 5. Schematic view
of LED illumination on adult survival and larval development (Oonincx et al. longevity and survival
2016)
750 Indian Journal of Entomology 84(3) 2022 Review
Fig. 5. Schematic view of effect of LED illumination on egg production and adult
longevity and survival in H. illucens (Nakamura et al., 2016)
illucens and effect Fig 5. Schematic view of effect of LED illumination on egg production and adult
nt (Oonincx et al. longevity and survival in H. illucens (Nakamura et al., 2016)
Fig. 6.
Fig. 6: Different
Different set
set of
of environmental
environmentalconditions
conditionsideal forfor
ideal BSF development
BSF development
3 to 13 days for females. The authors also concluded, along with similar microclimatic conditions (22-28°C
the shorter half-lives of adults in Halogen light may temperature, 40-60% humidity and 8 hours of artificial
be due to unsuitably higher temperature and excess light exposure) and the substrate (brewer’s grains). In
heat generation. Attiogbe et al. (2019) also applied the all setups over 98% of adult emergence were recorded
similar microclimatic conditions to achieve maximum and maximum amount of egg masses was found in
mating, suitable oviposition and larval development experimental cage made up of wood followed by
in order to get higher efficiency towards mercury corrugated cardboard, plastic structures, and glass
contaminated waste wherein direct association between structures wherein each cage was populated with 50g
larval density and reduced mercury content from waste of pupa at the start of experiment (Boaru et al., 2019).
was observed. Based upon the findings of different studies the range
for different set of environmental conditions ideal for
In the similar context, Boaru et al. (2019) described BSF development is shown in Fig. 6.
that suitable oviposition structures improves the
reproductive process of adults in captivity. The study 3. Cooccurrence of house flies while rearing BSF
included four different types of cage structure for While rearing, co-occurrence of houseflies along
oviposition or egg laying sites i.e., wood cages, glass with BSFs is also very evident and need to be discussed
cage, corrugated cardboards, and plastic material as it impacts the waste conversion efficiency and the
Black soldier fly Hermetia illucens (L.): Ideal environmental conditions and rearing strategies 751
Anshika Singh et al.
insects’ growth. Though the BSF are reported to repel and accuracy of larval age gauges which would serve
the housefly oviposition yet the experimental areas for other important purposes such as sustainable and
established for BSF rearing and mass production environmentally sound management of organic waste
often experiences the presence of house flies ensuing as it outshines the present conventional techniques
competition for the available substrate. In one such in several ways (less time for composing, higher
study (Miranda et al., 2019), no BSF pre pupation efficiencies of waste reduction, carbon sequestration,
occurred in the treatment (pig manure) inoculated with pathogen control).
house fly larvae at initial stages and reached maximum
pupation when reared alone on the fresh pig manure. ACKNOWLEDGEMENTS
In addition, the negative impact on BSF growth was The authors thank the director of CSIR-National
also attributed to the presence of houseflies and age Environmental Engineering Research Institute, Nagpur
of resource. Similarly, Hassan and Dina (2019) also for permission to write this review.
encountered the co-occurrence of a total of 3554
insects including the BSF where the fermented coconut REFERENCES
waste was used as egg laying sites in plastic bins. BSF
Addo-Bediako A, Chown S L, Gaston K J. 2000. Thermal tolerance,
larvae were highest in number in oviposition media but climatic variability and latitude. Proceedings of the Royal Society
Drosophila melanogaster and M. scalaris also have B: Biological Sciences 267: 739-745.
significant counts in competition to BSF. Alvarez L. 2012. The role of black soldier fly, Hermetia illucens (L.)
(Diptera: Stratiomyidae) in sustainable waste management in
CONCLUSIONS northern climates, Electronic Theses and Dissertations 402. https://
scholar.uwindsor.ca/etd/402 .
Black soldier flies have been performing
Attiogbe F K, Ayim N Y K, Martey J. 2019. Effectiveness of black
outstandingly well to deal with the concerns of soldier fly larvae in composting mercury contaminated organic
waste management and feed supplementation. waste. Scientific African. 6, e00205. https://doi.org/10.1016/j.
Correspondingly it demands the thorough knowledge sciaf.2019.e00205.
on specific tolerance limits of BSF towards ecological Banks I J. 2014. To Assess the impact of black soldier fly (Hermetia
illucens) larvae on faecal reduction in pit latrines. PhD thesis.
conditions. It was concluded that BSF growth and
London School of Hygiene and Tropical Medicine. https://doi.
development is directly influenced by temperature and org/10.17037/PUBS.01917781.
the diet composition serving to be the most crucial factor Barragan- Fonseca K B, Dicke M, van Loon J J. 2017. Nutritional value
in environment. The temperature and relative humidity of the black soldier fly (Hermetia illucens L.) and its suitability as
largely control the insect development, daily cycles, and animal feed–a review. Journal of Insects as Food and Feed 3(2):
105-120.
seasonal variations and affects the biology of insects,
their survival, life span, reproduction rate, population Boaru A, Vig A, Ladoşi D, Păpuc T, Struţi D, Georgescu B. 2019. The use
of various oviposition structures for the black soldier fly, Hermetia
growth parameters and the sex ratio. Optimum light illucens L. (Diptera: Stratiomyidae) in improving the reproductive
conditions, humidity also equally contribute to the process in captivity. Animal Biology and Animal Husbandry 11(1).
insect’s physiology, behaviour and morphological traits. Boykin K L. 2019. Assessing the nutritional value of black soldier
Moreover, wood based, and plastic material-based egg fly larvae (Hermetia illucens) used for reptile foods. https://
hatching structures may serve as the better oviposition digitalcommons.lsu.edu/gradschool_theses/4927/
sites and mass egg production in comparison to other Briscoe A, Chittka L. 2001. The evolution of color vision in insects.
Annual. Review of Entomology 46: 471-510.
conventional setups. However, it should be emphasized
that breeding activity is largely influenced by the Cammack J A, Tomberlin J K. 2017. The impact of diet protein and
carbohydrate on select life-history traits of the black soldier fly
specific environmental conditions. Hermetia illucens (L.) (Diptera: Stratiomyidae). Insects 8(2): 56.
Further, intensification of fecundity rate in BSF under Caruso D, Devic E, Wayan S I, Talamond P, Baras E. 2013. Technical
handbook of domestication and production of Diptera black soldier
indoor environment requires appropriate investigation fly (BSF) Hermetia illucens, Stratiomyidae. https://betuco-wp.be/
of suitable light source and intensities which may BSF/BSF%20-Technical%20Handbook.pdf
effectively enhance the mating rates, egg production Cheng J Y, Chiu S L, Lo I M. 2017. Effects of moisture content of food
and thus other life history parameters. In addition, there waste on residue separation, larval growth and larval survival in
is a need to examine how to improve the reproductive black soldier fly bioconversion. Waste Management 67: 315-323.
capacities of the flies for a progressive proficient raising Chia S Y, Tanga C M, Khamis F M, Mohamed S A, Alifu D, Sevgan
at the study sites. Future research ought to build up S, Fiaboe K K, Niassy S, van Loon J J, Dicke M, Ekesi S. 2018.
Threshold temperatures and thermal requirements of black soldier
extra models for distinguishing improvement of every fly Hermetia illucens: Implications for mass production. PloS One
instar, which may help in improving the exactness 13(11). p. e0206097.
752 Indian Journal of Entomology 84(3) 2022 Review
Cronin T W, Jarvilehto M, Weckstrom M, Lall A B. 2000. Tuning scale indoor rearing. PloS One 2018 13(5) e0197896. https://doi.
of photoreceptor spectral sensitivity in fireflies (Coleoptera: org/10.1371/journal.pone.0197896.
Lampyridae). Journal of Comparative Physiology A 186: 1-12
Holmes L. 2010. Role of abiotic factors on the development and life
Danieli P P, Lussiana C, Gasco L, Amici A, Ronchi B. 2019. The effects history of the black soldier fly, Hermetia illucens (L.) (Diptera:
of diet formulation on the yield, proximate composition, and fatty Stratiomyidae. Electronic Theses and Dissertations 285. https://
acid profile of the black soldier fly (Hermetia illucens L.) prepupae scholar.uwindsor.ca/etd/285.
intended for animal feed. Animals 9(4): 178.
Holmes L, Van Laerhoven S, Tomberlin J. 2016. Lower temperature
De Smet J, Wynants E, Cos P, Van Campenhout L. 2018. Microbial threshold of black soldier fly (Diptera: Stratiomyidae) development.
community dynamics during rearing of black soldier fly larvae Journal of Insects as Food and Feed 2(4): 255-262.
(Hermetia illucens) and impact on exploitation potential. Applied
Holmes L A, Vanlaerhoven S L, Tomberlin J K. 2012. Relative
Environmental Microbiology 84(9): e02722-17.
humidity effects on the life history of Hermetia illucens
Deutsch C A, Tewksbury J J, Huey R B, Sheldon K S, Ghalambor C (Diptera:Sstratiomyidae). Environmental Entomology 41 (4):
K, Haak D C, Martin P R. 2008. Impacts of climate warming on 971-978.
terrestrial ectotherms across latitude. Proceedings of the National
Jarosík V, Kratochvíl L, Honék A, Dixon A F A. 2004. General rule for the
Academy of Sciences of the United States of America 105: 6668-
dependence of developmental rate on temperature in ectothermic
6672.
animals. Proceedings of the Royal Society B: Biological Sciences
Diener S, Studt Solano N M, Roa Gutiérrez F, Zurbrügg C, Tockner 271: S219-S221.
K. 2011b. Biological treatment of municipal organic waste using
Leong S Y, Kutty S R M, Malakahmad A, Tan C K. 2016. Feasibility study
black soldier fly larvae. Waste Biomass Valorization 2: 357-363.
of biodiesel production using lipids of Hermetia illucens larva fed
Diener S, Zurbrügg C, Roa Gutiérrez F, Hong Nguyen D, Morel A, with organic waste. Waste Management 47: 84-90.
Koottatep T, Tockner K. 2011a. Black soldier fly larvae for organic
Li C J. 2014. Conversion of spent grains and DDGS by black soldier flies.
waste treatment- prospects and constraints. Khulna, Bangladesh.
MSc thesis, Laboratory of Entomology, Wageningen University,
Proceedings. Waste Safe 2nd international conference on solid waste
Wageningen, the Netherlands.
management in the developing countries 978-984-33-2705-5.
Liu Q, Tomberlin J K, Brady J A, Sanford M R, Yu Z. 2008. Black soldier
Diener S, Zurbrugg C, Tockner K. 2009. Conversion of organic material
fly (Diptera: Stratiomyidae) larvae reduce Escherichia coli in dairy
by black soldier fly larvae: establishing optimal feeding rates. Waste
manure. Environmental Entomology 37: 1525-1530.
Management and Research 27(6): 603-610.
Ma J, Lei Y, Rehman KU, Yu Z, Zhang J, Li W, Li Q, Tomberlin J K,
Dixon A F G, Honˇek A, Keil P, Kotela M A A, Šizling A L, Jarošík
Zheng L. 2018. Dynamic effects of initial pH of substrate on
V. 2009. Relationship between the minimum and maximum
biological growth and metamorphosis of black soldier fly (Diptera:
temperature thresholds for development in insects. Functional
Stratiomyidae). Environmental Entomology 47 (1): 159-165.
Ecology 23: 257-264.
Manyara N E. 2018. Optimization of production of black soldier fly larvae
Dortmans B, Diener S, Verstappen B M, Zurbrügg C. 2017. Black soldier
(Hermetia illucens L) for fish feed formulation. https://greeinsect.
fly biowaste processing: A step-by-step guide. Swiss Federal
ku.dk/phd-projects-edible-insects/evans-nyakeri/hb_phd-thesis-
Institute of Aquatic Science and Technology (Eawag), Dübendorf,
cover/Evans-Thesis.pdf.
Switzerland, 88.
Martínez-Sánchez A, Magaña C, Saloña M Rojo S. 2011. First record of
Ewusie E A, Kwapong P K, Ofosu-Budu G, Sandrock C, Akumah A
Hermetia illucens (Diptera: Stratiomyidae) on human corpses in
M, Nertey E K, Teye-Gaga C, Agyakwah S K. 2019. The black
Iberian Peninsula. Forensic Science International 206(1): e76-e78.
soldier fly, Hermetia illucens (Diptera: Stratiomyidae): Trapping
and rearing of wild colonies in Ghana. Scientific African 5, e00134. Meneguz M, Gasco L, Tomberlin J K .2018. Impact of pH and feeding
system on black soldier fly (Hermetia illucens, L; Diptera:
Gasco L, Dabbou S, Trocino A, Xiccato G, Capucchio M T, Biasato I,
stratiomyidae) larval development. PLoS One 13 (8): e0202591.
Dezzutto D, Birolo M, Meneguz M, Schiavone A. 2019. Effect of
https://doi.org/10.1371/journal.pone.0202591.
dietary supplementation with insect fats on growth performance,
digestive efficiency and health of rabbits. Journal of Animal Science Miranda C D, Cammack J A, Tomberlin J K. 2019. Life-History
and Biotechnology 10: 4. traits of the black soldier fly, Hermetia illucens (L.) (Diptera:
Stratiomyidae), reared on three manure types. Animals 9(5): 281.
Gobbi P, Martinez-Sanchez A, Rojo S. 2013. The effects of larval
https://doi.org/10.3390/ani9050281.
diet on adult life-history traits of the black soldier fly, Hermetia
illucens (Diptera: Stratiomyidae). European Journal of Entomology Müller A, Wolf D, Gutzeit H O. 2017. The black soldier fly, Hermetia
110(3): 461. illucens– a promising source for sustainable production of proteins,
Green T R, Popa R. 2012. Enhanced ammonia content in compost leachate lipids and bioactive substances. Zeitschrift für Naturforschung C.
processed by black soldier fly larvae. Applied Biochemistry and 72(9-10): 351-363.
Biotechnology 166 (6): 1381-1387. Myers H M, Tomberlin J K, Lambert B D, Kattes D. 2014. Development
Harden L M, Tomberlin J K. 2016. Effects of temperature and diet on of black soldier fly (Diptera: Stratiomyidae) larvae fed dairy
black soldier fly, Hermetia illucens (L.) (Diptera: Stratiomyidae) manure. Environmental Entomology 37(1): 11-15.
development. Forensic Science International 266:109-116. Nakamura S, Ichiki R T, Shimoda M, Morioka S. 2016. Small-scale
Hasan H A, Dina F. 2019. Co-occurrence of different insect species in rearing of the black soldier fly, Hermetia illucens (Diptera:
oviposition media of black soldier fly, Hermetia illucens (Diptera: Stratiomyidae), in the laboratory: low-cost and year-round rearing.
Stratiomyidae). Serangga 24(2). Applied Entomology and Zoology 51(1): 161-166.
Heussler C D, Walter A, Oberkofler H, Insam H, Arthofer W, Schlick- Nguyen T T, Tomberlin J K, Vanlaerhoven S. 2013. Influence of resources
Steiner B C, Steiner F M. 2018. Influence of three artificial light on Hermetia illucens (Diptera: Stratiomyidae) larval development.
sources on oviposition and half-life of the black soldier fly, Journal of Medical Entomology 50: 898-906.
Hermetia illucens (Diptera: Stratiomyidae): Improving small- Nguyen T T X, Tomberlin J K, Vanlaerhoven S. 2015. Ability of black
Black soldier fly Hermetia illucens (L.): Ideal environmental conditions and rearing strategies 753
Anshika Singh et al.
soldier fly (Diptera: Stratiomyidae) larvae to recycle food waste. nutritive value of black soldier fly larvae reared on common organic
Environmental Entomology 44 (2): 406-410. waste streams in Kenya. Scientific Reports 9(1): 1-3.
Nowak V, Persijn D, Rittenschober D, Charrondiere U R. 2016. Review Singh A, Kumari K. 2019. An inclusive approach for organic waste
of food composition data for edible insects. Food Chemistry 193: treatment and valorisation using black soldier fly larvae: A review.
39-46. Journal of Environmental Management, 251: 109569. https://doi.
org/10.1016/j.jenvman.2019.109569.
Oonincx D G A B, Van Broekhoven S, Van Huis A, Van Loon J J A. 2015b.
Feed conversion, survival and development, and composition of Spranghers T, Michiels J, Vrancx J, Ovyn A, Eeckhout M, De Clercq P,
four insect species on diets composed of food by products. PLoS De Smet S. 2018. Gut antimicrobial effects and nutritional value
One 10, e0144601. of black soldier fly (Hermetia illucens L.) prepupae for weaned
piglets. Animal Feed Science and Technology 235: 33-42.
Oonincx D G A B, Van Huis A. Van Loon J J A. 2015a. Nutrient utilisation
by black soldier flies fed with chicken, pig, or cow manure. Journal Stavenga D G. 1992. Eye regionalization and spectral tuning of retinal
of Insects as Food and Feed 1: 131-139. pigments in insects. Trends in Neurosciences 15: 213-218.
Oonincx D G A B, Volk N, Diehl J J E, Van Loon J J A, Belušič G. 2016. Tomberlin J K, Adler P H, Myers H M. 2009. Development of the black
Photoreceptor spectral sensitivity of the compound eyes of black soldier fly (Diptera: Stratiomyidae) in relation to temperature.
soldier fly (Hermetia illucens) informing the design of LED-based Environmental Entomology 38(3): 930-934.
illumination to enhance indoor reproduction. Journal of Insect Tomberlin J K, Sheppard D C. 2002. Factors influencing mating and
Physiology 95: 133-139. oviposition of black soldier flies (Diptera: Stratiomyidae) in a
Park H H. 2016. Black soldier fly larvae manual. http://scholarworks. colony. Journal of Entomological Science 37(4): 345-352.
umass.edu.Erişim.tarihi Tomberlin J K, Sheppard D C, Joyce J A. 2002. Selected life-history
Pathak R, Sharma S, Prasad R. 2015. Study on occurrence of black soldier traits of black soldier flies (Diptera: Stratiomyidae) reared on three
fly larvae in composting of kitchen waste. International Journal of artificial diets. Annals of the Entomological Society of America
Research in Biosciences 4 (4): 38-45. 2002 95(3): 379-386.
Peitsch D, Feitz A, Hertel H, de Souza J, Ventura D F, Menzel R. 2001. Tomberlin, J K, Sheppard D C. 2001. Lekking behaviour of the black
The spectral input systems of hymenopteran insects and their soldier fly (Diptera: Stratiomyidae). Florida Entomologist 84:
receptor-based colour vision. Journal of Comparative Physiology 729-730.
A 170: 23-40. Tsagkarakis A E, Arapostathi E, Strouvalis G. 2015. First record of
Renna M, Schiavone A, Gai F, Dabbou S, Lussiana C, Malfatto the black soldier fly, Hermetia illucens, in Greece. Entomologia
V, Prearo M, Capucchio M T, Biasato I, Biasibetti E. 2017. Hellenica 24: 27-30.
Evaluation of the suitability of a partially defatted black soldier fly Tschirner M, Simon A. 2015. Influence of different growing substrates
(Hermetia illucens L.) larvae meal as ingredient for rainbow trout and processing on the nutrient composition of black soldier fly
(Oncorhynchus mykiss Walbaum) diets. Journal of Animal Science larvae destined for animal feed. Journal of Insects as Food and
and Biotechnology 8: 57. Feed 1: 249-259.
Sharanabasappa D, Srikanth B H, Maruthi M S , Pavithra H B. 2019. Ustuner T, Hasbenli A, Rozkosny R. 2003. The first record of Hermetia
Biology of black soldier fly Hermetia illucens (L.) (Diptera: illucens (Linnaeus, 1758) (Diptera, Stratiomyidae) from the near
Stratiomyidae) on muskmelon fruit. Indian Journal of Entomology east. Studia Dipterologica 10 (1): 181-185.
81(1): 153-155.
Van Huis A, Tomberlin J. 2017. The potential of insects as food and
Sheppard C D, Tomberlin J K, Joyce J A, Kiser B C, Sumner S M. 2002. feed. Insects as food and feed: from production to consumption;
Rearing methods for the black soldier fly (Diptera: Stratiomyidae). Van Huis A, Tomberlin J (eds). Wageningen Academic Publishers:
Journal of Medical Entomology 39(4): 695-698. Wageningen, The Netherlands. pp. 25-58.
Srikanth B H,Sharanabasappa D. 2021. Growth performance and Win SS, Ebner J H, Brownell S A, Pagano S S, Cruz-Diloné P, Trabold
bioconversion rate of black soldier fly, Hermetia illucens (L.) T A. 2018. Anaerobic digestion of black solider fly larvae (BSFL)
when reared on organic feed wastes. Indian Journal of Entomology biomass as part of an integrated bio refinery. Renewable Energy
83(2): 155-158. 127: 705-712.
Shumo M, Khamis F M, Tanga C M, Fiaboe K K, Subramanian S, Ekesi Yu G, Chen Y, Yu Z, Cheng P. 2009. Research progress on the larvae
S, Van Huis A, Borgemeister C. 2019. Influence of temperature on and prepupae of black soldier fly Hermetia illucens used as animal
selected life-history traits of black soldier fly (Hermetia illucens) feedstuff. Chinese Bulletin of Entomology 46 (1): 41-45.
reared on two common urban organic waste streams in Kenya.
Zhang J, Huang L, He J, Tomberlin J K, Li J, Lei C, Sun M, Liu Z, Yu Z.
Animals 9(3): 79.
2010. An artificial light source influences mating and oviposition
Shumo M, Osuga I M, Khamis F M, Tanga C M, Fiaboe K K, of black soldier flies, Hermetia illucens. Journal of Insect Science
Subramanian S, Ekesi S, van Huis A, Borgemeister C. 2019. The 10 (1). https://doi.org/10.1673/031.010.20201.