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Black Soldier Fly (Linnaeus 1758)

Fig. 1. Hermetia illucens (Linnaeus, 1758). Image from Samuel Ewing. Permission obtained and
accredited under copyright.

Overview
Table of Contents

Overview

Distribution

Locality

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Habitat

Significance of Hermetia illucens

Environmental importance

Economic importance

Morphology and Identification

General anatomy

Larval and pupae 

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Adult characteristics and diagnostic features

Sexual dimorphism

Biology

Diet and Feeding 

Mating behaviour and oviposition

Development and life cycle

Rearing and breeding in captivity 

Predation

Defensive Adaptation

Conservation status

Taxonomy and Systematics

Synonyms 

Type information

Taxonomic classification

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Phylogeny

References

Acknowledgement

Contact Me

The Black Soldier Fly (Hermetia illucens) is a cosmopolitan fly

belonging to the Stratiomyidae family (1). It is a relatively large
fly with wasp-like appearance. However, unlike wasps, H.
illucens only possess one pair of wings and lacks a stinger.
Moreover, the species exhibits a vast geographical range from
America to South-east Asia (2) (3) (4). H. illucens is usually
found in outdoor environments near livestock or decaying organic
matter, including animal waste (5). H. illucens larvae are widely
used in manure management, controlling of housefly populations
and the conversion of organic waste into useful products such as
compost (5). The larvae are also sold as important sources of food
for reared amphibians and fish as they are high in calcium content
(6). In forensic entomology, the life history and development of
H. illucens in decaying corpses is used by medico-legal
investigators for post-mortem analysis (7) (8).

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Distribution
Locality
 (Back to top)
Hermetia illucens is a widespread cosmopolitan species found in
various parts of the world (4). The species is hypothesized to
originate from tropical, subtropical and temperate zones of
America though this is widely contested (4) (9) (10). It is believed
that international transport and commerce in the 20th century led
to the introduction of H. illucens in other parts of the world.
However, others hypothesize that the species is native to
Palearctic regions of earth (3) (4) (11). In Europe, H. illucens was
first recorded in Malta and has since been discovered and
recorded in other parts of Europe and Asia (4); (11); (12). The
widespread distribution of H. illucens is shown in the map below
(4):

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Habitat
Hermetia illucens are commonly found resting on garden plants,
tree trunks and walls and windows in urban residential areas (13).
The species also inhabit areas with livestock and decaying
organic matter (13).

Significance of Hermetia illucens

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Environmental importance
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Unlike the common housefly, Hermetia illucens is not considered

a pest as it is not attracted to food or houses (14). In fact, their
larvae play pivotal roles in both environmental and economic
aspects of waste disposal and processing. H. illucens larvae have
been documented to reduce dry manure biomass by 42-56%.
Nitrogen concentration and mass were also reduced by 24% and
62% respectively (15) (16). This drastically reduces the amount of
environmental pollutants. Moreover, food processed by larvae
aerates and dries up organic waste, thereby reducing the
production of gases such as methane, an important greenhouse
gas (5). Other odours that may attract pests are also eliminated as
the larvae feed on waste (5).
Feeding by H. illucens larvae greatly reduces the production and
establishment of a pest housefly, Musca domestica L., near
decaying organic matter (14) (15) (17). This occurs as H. illucens
larvae inhibit the oviposition of M. domestica L. on food waste,
possibly by overcrowding their food resource and producing
short-lived interspecific chemical signals (18). H. illucens also
alters the microflora composition of organic waste, reducing the
growth of harmful bacteria such as Escherichia coli O157:H7 and
Salmonella in manure (19) (20). Consequently, this reduces the
spread of disease by reducing bacteria growth and disease vectors
such as M. domestica L.

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Vid. 1. Housefly larvae out-competed by Hermetia illucens

larvae. Obtained from YouTube under fair use

Economic importance

Economically, Hermetia illucens reduces costs from waste

collection and landfills. It is estimated that landfilling is three
times more expensive than composting (21). By reducing the
volume of waste by 50%, H. illucens drastically decreases the
need and costs of landfill. Notably, organic waste composting by
H. illucens allows for waste recycling and fertilizer production
(20) (22). Harvested H. illucens larvae and pre-pupae can be sold
as fish feed in various forms (see rearing and breeding in captivity
), replacing more expensive conventional fish feed. H. illucens
has also been used in the production of sugar and biodiesel from
dairy manure (23). It is estimated that manure management using
H. illucens could increase revenue by $25 000 per year per cage
house of livestock (5). However, research into the
industrialization of H. illucens is currently ongoing. For example,
research on the use of H. illucens as a composting agent in
campuses is currently conducted in Singapore and America (24)
(25). 
Hermetia illucens larvae or pre-pupae could be used as high fat and protein feed for fishes as many conventional fish feed lack essential amino acids or have low protein content (6) (26) (27) . Larvae and pre-pupae can be fed live, chopped, frozen or ground

(6) (26) (27). Studies have shown that a diet consisting of H. illucens larvae and pre-pupae does not negatively affect fish growth. It may thus serve as a valuable replacement for other more expensive sources of fish feed (28) (29). H. illucens larvae was so successful as a feeder insect that it became the first to be registered under a U.S trademark and is now marketed under the brand name “Phoenix Worms”. It is also sold under different brands such as "Reptiworms" (Canada) and "Calci Worms" (30) (31) (32).

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Fig. 2. Schematic of an industrial composting process employing the use of Hermetia illucens larvae as a
composting organism and harvesting for production of fish feed (Newton et al., 2005). Obtained and
accredited under fair use

Should Hermetia illucens be unavailable for composting, species

such as the yellow soldier fly (Ptecticus trivittatus) are also
known to degrade organic matter (33). However, unlike H.
illucens, P. trivittatus prefers rotten fruit which attracts their
females to lay eggs (34).  are usually found in compost along
P. trivittatus 

with H. illucens and can be distinguished by their smaller size,

yellowish colouration, golden thorax and short yellow antennae
(35). Little research has focused on the composting ability of P.
trivittatus and its usefulness as a composting organism remains
unknown.

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Fig 3. Image of yellow soldier fly (Ptecticus trivittatus). Obtained from Sally G. Miller. Permission
obtained and accredited under copyright.

Morphology and Identification

General anatomy
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Fig. 5. General anatomy of Hermetia illucens adult.

Frontal view. Scale represents 1mm. Image by
Jonathan Tan

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Fig. 6. General anatomy of Hermetia illucens adult.

Lateral view. Scale represents 5mm. Image by
Jonathan Tan

Fig. 4. General anatomy of Hermetia illucens adult.

Dorsal view. Scale represents 1mm. Image by
Jonathan Tan

Larval and pupae 

Larvae of Hermetia illucens are dull white in colour with
golden-yellow setae and can grow up to 27mm in length (13).
They have a small protruding yellowish-brown head with
mouthparts for feeding. H. illucens larvae have light yellow
ocular prominences on the lateral side of their head while the
antennae are low and situated anterolaterally (4). 
As Hermetia illucens larvae develops into pupae, they turn dark
brown in colour. First instar larvae and pupae of H. illucens have

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not been imaged before. Thus, imaging of key features of H.

illucens larvae will help to distinguish them from other fly larvae
such as Musca domestica L. Moreover, images could also be used
for identification. Mouthparts are also imaged to show
morphological differences. Expertise description of larval and
pupal images are extremely welcomed.
First instar larvae Pupae
Dorsal view

Fig. 7. First instar larvae of Fig. 8. Pupae of Hermetia illucens.

Hermetia illucens. Dorsal view. Dorsal view. Scale represents
Scale represents 1mm. Image by 1mm. Image by Jonathan Tan
Jonathan Tan
Ventral view

Fig. 9. First instar larvae of

Fig. 10. Pupae of Hermetia
Hermetia illucens. Ventral view.
illucens. Ventral view. Scale
Scale represents 1mm. Image by
represents 1mm. Image by
Jonathan Tan
Jonathan Tan
Lateral view

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Fig. 12. Pupae of Hermetia

Fig. 11. First instar illucens. Lateral view. Scale
larvae of Hermetia represents 1mm. Image by
Jonathan Tan
illucens. Lateral
view. Scale
represents 1mm.
Image by Jonathan
Tan

Mouthparts

Fig.13. First instar larvae of Fig. 14. Pupae of Hermetia

Hermetia illucens. View of illucens. View of mouthparts.
mouthparts. Scale represents 1mm. Scale represents 1mm. Image by
Image by Jonathan Tan Jonathan Tan

Adult characteristics and diagnostic features

Trait Property Image
Anterior-posterior Typically large See General Anatomy
length flies, 15- 20mm
in length. Black
in colour. (36)
>

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Antennae Antennae are

elongated and at
least twice the
length of head in
lateral view (4)
Fig. 15. Lateral view of Hermetia
illucens head. Circles represent
numbered segments of antennae.
Note the length of the antennae
compared to the head. Image by
Jonathan Tan
First two abdominal Abdomen Fig. 16. Dorsal view of Hermetia
terga  consisting of 5 illucens abdomen. Circles
represent the transclucent patches
visible segments on abdominal terga 1 and 2 as well
and is black in as reddish-brown hue of the last
abdominal tergum. Image by
colour. Jonathan Tan.

Posterior sides of
tergum one and
two contain two
translucent white
oblong spots.

Last terga has a

reddish-brown
hue. (4)
Legs Legs are black
with white tarsi
and basal halves
of hind tibiae (4)

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Fig. 17. Lateral view of Hermetia

illucens legs. Circles represent the
white tarsi. Note the whitish
portion of the basal half of hind
tibia. Image by Jonathan Tan.

Sexual dimorphism
Hermetia illucens exhibit various sexually dimorphic traits such
as body size, features on the frons and head, abdominal spots and
the shape of their abdominal terminalia (4). The most common
way of distinguishing them are the number of whitish hairs on the
face and shape of abdominal terminalia (4).
Trait Female Male
Face

Whitish hairs on
the face
Fig. 18. Frontal view of Hermetia Fig. 19. Frontal view of Hermetia
Females have illucens female. Note the larger illucens male. Note that whitish
number of whitish hairs on the hairs are more sparse on the
much more greater and lower parts of the face greater and lower parts of the face.
whitish hairs on compared to the male individual.
Scale represents 1mm. Image by
Scale represents 1mm. Image by
Jonathan Tan.
the face such that Jonathan Tan.

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only the four

frontal and facial
tubercles are
dark
Abdominal
terminalia 

Female:

Female Fig. 20. Abdominal terminalia of Fig. 21. Genitalia of Hermetia

abdominal Hermetia illucens female. Scale illucens male. Scale represents
represents 1mm. Image by 1mm. Image by Jonathan Tan.
terminalia is Jonathan Tan.
long. The cerci
has two
segments and is
elongated.

The subgenital
plate points
distally and is
elongated.
Genital furca are
subtriangular
with a big
median aperture.

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It also possess
broad,
leaf-shaped
projections. 

Male:

Male gentalia are

relatively shorter
than the female
terminalia. The
genitalia are
proximally
emarginate with
a synsternum
possessing two
postlateral lobes
on each side.
Presence of
claspers on the
male

Biology

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Diet and Feeding 

 (Back to top) 
Hermetia illucens mainly feed during their larval stages. H.
illucens larvae feed on a large variety of organic matter, including
plant material, decaying animal matter and manure (5) (37) (38)
(39). They are capable of converting large amounts of waste
biomass into stored protein (≥40%) and fat (≥30%). Hence H.
illucens larvae serve as highly proteinaceous animal feed with
high energy content (5) (6). In contrast, H. illucens adults do not
feed on solid material but drink liquids (36) (40). Thus most
nutrient stores are accumulated during larval phases, reducing the
need for feeding as an adult (5) (36). 

Vid. 2: Time lapse of Hermetia illucens larvae finishing a banana.

Obtained from YouTube under fair use

Fig. 22. Image of Hermetia illucens adult feeding on sugar syrup. Image by Jonathan Tan.

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Mating behaviour and oviposition

Male Hermetia illucens exhibit territorial lekking behaviour (41).
At forest edges, they were observed to be resting on plant leaves
and waiting for potential female mates (41). Males tend to
aggregate in large numbers at particular sites in the wild when
mating (41). Aggressive behaviour was also reported when other
male H. illucens invaders entered into the territory of the resting
male. This usually occurs in the form of aerial spiralling in which
one fly tries to fend off the other (41). At the end of this aerial
display, the “victor” would return to its original resting place
whilst the other leaves (41). When a male encounters a passing
female, it would fly up towards her and grasp her (41). The pair
then descends in copula. Mating activities usually occur in the
day when sunlight intensity is high and decreases with
diminishing light intensity (42). The presence of sunlight was
correlated to mating and successful fertilization of H. illucens
eggs (36) (42). However, additional research is required to
determine the correlation between light wavelengths and mating
behaviour.

Vid. 3. Initial courtship of Hermetia illucens individuals without

in copula mating. Obtained from YouTube under fair use

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Fig. 23. Successful mating of Hermetia illucens individuals in copulaImage. Obtained and use with
permission from Foo Maosheng

Once mating has been completed, the female departs in search of

oviposition sites. H. illucens Females tend to lay their eggs on
cracks and crevices such as the flukes of corrugated cardboard
near decaying matter (13) (42) (43). Oviposition is affected by
temperature and humidity. Females tend to deposit eggs at
temperatures higher than 26 °C and at high humidity (≥60%).
80% of the eggs were deposited when environmental humidity
exceeded 60% and this was almost twice the number of eggs laid
at lower humidity (42).

Development and life cycle

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Fig. 24. Hermetia illucens female laying eggs in corrugated cardboard. Image by Gee W. Obtained and
accredited from Wikimedia Creative Commons
A gravid Hermetia illucens female may deposit up to 500
oval-shaped, pale yellow eggs in a single clutch (13). After four
days, larvae will hatch from these eggs (42). The entire
developmental process from birth to adulthood takes
approximately 40−43 days (14) (40) (42). A typical H. illucens
larvae stage consists of 6 instars and last for 14−22 days during
which they feed and accumulate biomass (13) (42). At the end of
the larval stage, H. illucens larvae enter the pre-pupae stage.
During the pre-pupal stage, H. illucens stop feeding and their
mouthparts are modified into a climbing appendage (see Larvae
and Pupae: Mouthparts). Pre-pupae then migrate away from the
larval habitat and organic matter to pupate in a dry habitat using
modified pupal mouth appendages (5). Adults then emerge
approximately 14 days later (40). Pupal time may extend up to a
few months and is highly variable (14). This depends on
environmental conditions, particularly temperature and biotic
conditions such as larval density and accumulated nutrients (40)
(42) (44). Mating starts 2 days after adults emerge (45). The
average life span of adults is approximately 10 days (40).

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Fig. 25. Diagram showing life cycle of Hermetia illucens. Obtained from YouTube under fair use

Fig. 26. First five instars of Hermetia illucens followed by pre-pupae and pupae stages (from left to
right). Image by Jonathan Tan

Rearing and breeding in captivity 

Hermetia illucens are able to process large quantities of organic
matter and are consequently important composting organisms as
manure and waste management tools (5) (46). Several substrates
such as pig manure (5), poultry waste (46), food wastes (24) and
various feed meals (40) have been used as raw material for
composting. Rearing facilities usually consist of cage layered
houses under the animal producing waste (5) (24) (46). Optimum
temperature and humidity ranges are maintained in these facilities

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to ensure high yield of H. illucens larvae production.

Temperatures are kept at 29–31 °C and humidity at 50–70% (24).
Rearing facilities operate by self-collection of H. illucens
pre-pupae when they migrate away from their habitat into
collection bins (5). This occurs as H. illucens larvae climb out of
food bins onto a ramp that leads them into a container where they
pupate (5) (47). Collected H. illucens pupae are then brought into
a greenhouse where they will hatch into adults. The size of screen
cages to effectively rear H. illucens adults vary. Sheppard et al.
(2002) were able to successfully mate H. illucens in a 2 by 2 by
4m screen cage housed in a greenhouse (40). Space is required in
order for the aerial mating process to take place (24). It is also
recommended that adult H. illucens are kept in greenhouses or
exposed to sunlight as sunlight intensity affects mating (42) (46).
Optimal ranges of temperatures and humidity for mating and
oviposition are 24–40°C and 30-90% respectively (36).
Recommended minimum light intensity for mating to occur is 63
µmolm2s-1 and optimal light intensity at over 200 µmolm2s-1
(36). Exposure of the cage to sunlight is recommended though
artificial lighting has also been shown to work. For successful
oviposition, a moist container or corrugated cardboard tray (ie
egg-holding tray) should be provided and placed near decaying
organic matter which attracts gravid females (24).

Vid. 4. Home-made composting bin with Hermetia illucens

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larvae. Obtained from YouTube under fair use

Predation
Not much is known about predators of Hermetia illucens.
However, animals such as birds, frogs and lizards are known
predators of H. illucens larvae.

Defensive Adaptation
 Hermetia illucens are often described as “wasp-like” due to two
translucent patches on their second abdominal tergum, allowing
them to appear to possess a thin slender, wasp-like waist (4) (36).
H. illucens has an elongated antennae and black legs with white
tarsi that are also features of wasp-like biomimicry to help deter
potential predators (4) (36).

Fig. 27. Example of an organ pipe mud dauber wasp Fig. 28. Example of Hermetia illucens adult. Note
(Trypoxylon politum). Image by Stephen Crewell. the similarities in colour and white tarsi of Hermetia
Permission obtained and accredited under copyright. illucens to the wasp. Image by Jonathan Tan

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Conservation status
Hermetia illucens is listed as “extant” and does not face the threat
of extinction due to their widespread and cosmopolitan
distribution (1) (4). 

Taxonomy and Systematics

Synonyms 
(Back to top)

Listed below are some of the synonyms for Hermetia illucens

(Linnaeus, 1758) from the Integrated Taxonomic Information
System (ITIS) (48):
Musca illucens Linnaeus, 1758 
Musca leucopa Linnaeus, 1767
Hermetia rufiventris Fabricus, 1805
Hermetia nigrifacies Bigot, 1879
Hermetia pellucens Macquart, 1834
Hermetia mucens Riley & Howard, 1889
Hermetia illuscens Copello, 1926
Hermetia illucens var. nigritibia Enderlein, 1914

Type information

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The holotype for Hermetia illucens is in the De Geer collection

housed in the Swedish museum of Natural History
(Naturhistoriska Riksmuseet, 4).

Taxonomic classification
The taxonomic classification shown below is referenced to
UniProt Taxonomy (http://www.uniprot.org/taxonomy/343691).
The classification reflects ranks of Hermetia illucens above the
species level and is arranged in descending order from Kingdom
to Species level:
 › Eukaryota
› Opisthokonta
 › Metazoa
 › Eumetazoa
 › Bilateria
 › Protostomia
 › Ecdysozoa
 › Panarthropoda
 › Arthropoda
 › Mandibulata
 › Pancrustacea
 › Hexapoda
 › Insecta
 › Dicondylia
 › Pterygota
 › Neoptera
 › Endopterygota

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 › Diptera
 › Brachycera
 › Stratiomyomorpha
 › Stratiomyidae
 › Hermetiinae
 › Hermetia

Phylogeny
The phylogeny of soldier flies has been well-studied using
molecular and morphological methods in phylogenetic tree
construction (49) (50) (51) (52). Morphological characters include
external and internal morphology of larvae, pupae and adults,
such as wings and genitalia (11) (50) (51) (52). Molecular
characters consist of nuclear protein-coding genes, ribosomal
DNA (18S and 28S) and complete mitochondrial genomes  . (50) (51) (52)

Monophyly of the suborder Brachycera (>100,000 species, 40

families) which contains the family Stratiomyidae of solider flies
is well supported (52). A comprehensive study of the fly tree of
life by Wiegmann et al. (2011) places Stratiomyidae beside sister
clades Xylomyidae and Pantophthalamidae, all of which comprise
the infraorder Stratiomyomorpha or solider flies  . However,
(52)

relationships between the various subfamilies of Stratiomyidae

are less apparent due to less well-supported monophyly and
relationships between certain subfamilies such as Stratiomyinae
and Clitellariinae (50).

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Fig. 29. Molecular phylogenetic tree for Diptera by Wiegmann et al. (2011). Circles on the tree indicate
well-supported nodes with bootstrap values >80%. Nodes with stars indicate improvement of bootstrap
values after postanalysis pruning of non-stable taxa. Coloured squares indicate certain ecological traits
(listed at bottom left) present in at least one species of the family.Obtained and accredited under fair use

Stratiomyidae encompass a family of cosmopolitan flies of

suborder Brachycera and infraorder Stratiomyomorpha  . The (11) (50)

monophyletic family comprises over 2800 species (50).

Monophyly of the subfamily Hermetiinae which contains the
genus Hermetia and species Hermetia illucens is well-supported
both by morphological and molecular data  . (11) (50)

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Fig. 30. Molecular phylogenetic tree showing relationship of various sub-families of Stratiomyidae by
Brammer & Dohlen (2007). The tree is one of the six most-parsimonious trees found with PAUP and
NONA via combined EF-1-alpha and 28S sequences. Branch lengths reflect and are proportional to the
number of base substitutions. Bootstrap values for nodes are indicated below the branches. Box in red
indicates the sub-family Hermetiinae that contains Hermetia illucens. Abbreviations of sub-family
names are as follows: P = Parhadrestiinae, Ch = Chiromyzinae, Pa = Pachygastrinae, B = Beridinae, A =
Antissinae, Cl = Clitellariinae, Cr = Chrysoclorininae, H = Hermetiinae, N = Nemotelinae, S = Sarginae,
St = Stratiomyinae, R = Raphiocerinae. Obtained and accredited under fair use

It is hypothesized that the primary terrestrial lineages of the

suborder Brachyera, including the infraorder Stratiomyomorpha,
diversified in the early Jurassic (160 mya, 52). However, the
common ancestor of the Stratiomyidae family originated more
recently in a radiation event during the early Cretaceous period
(around 129 mya, 50). The major radiation of clades in

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Stratiomyidae, such as Hermetiinae, is estimated to be around

60-80 mya using molecular data (50). The last common ancestor
of H. illucens has been estimated to appear during the early
Cenozoic (around 40 mya, 50).

Fig. 31. Chronogram showing the divergence of dipteran clades and estimated time of divergence by
Wiegmann et al. (2011). Shaded boxes represent uncertainty in phylogenetic analysis and crucial periods
of rapid diversification for diptera. The vertical height of the coloured triangles represents the estimated
numbers of species described in each clade with Stratiomyomorpha indicated in pink. The corresponding
height of the grey scale bar represents 10, 000 species. Timescale represents millions of years before
present. Obtained and accredited under fair use.

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Fig. 32. Chronogram from r8s analysis of both EF-1-alpha and 28S sequences by Brammer & Dohlen
(2007). Timescale represents millions of years before present. Nodes that had their age constrained due
to fossil evidence are represented by filled circles with the corresponding age indicated. Branches with
high posterior probability (>90%) and bootstrap support (>775%) are bolded. The bracket indicates the
Cretaceous period. Sub-family Hermetiinae is highlighted via the red box. Abbreviations of sub-family
names are as follows: P = Parhadrestiinae, Ch = Chiromyzinae, Pa = Pachygastrinae, B = Beridinae, A =
Antissinae, Cl = Clitellariinae, Cr = Chrysoclorininae, H = Hermetiinae, N = Nemotelinae, S = Sarginae,
St = Stratiomyinae, R = Raphiocerinae. Obtained and accredited under fair use

References
(Back to top)

1. Jacinto, V. & Coin, P. eds. (2015). "Hermetia illucens."

Encyclopedia of Life. Retrieved 7 Nov 2015 from
http://eol.org/pages/824054/overview. 
2. Hardouin, J. & Mahoux, G. (2003). Zootechnie d’insectes –

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Elevage et utilisation au bénéfice de l'homme et de certains

animaux. Bureau pour l’Echange et la Distribution de
l’Information sur le Mini-élevage (BEDIM), 164 p.
3. Kim, J. I. (1997). Newly recording two exotic insects species
from Korea. J. Kor. Biota, 2, 223−225.
4. Rozkosný, R. (1983). Subfamily Hermetiinae. In A
Biosystematic Study of the European Stratiomyidae (Diptera):
Volume 2-Clitellariinae, Hermediinae, Pachygasterinae and
Bibliography (Vol. 2). London: Springer Science & Business
Media. pp.431.
5. Newton, G. L., Sheppard, D. C., Watson, D. W., Burtle, G. J.,
Dove, C. R., Tomberlin, J. K., & Thelen, E. E. (2005). The black
soldier fly, Hermetia illucens, as a manure management/resource
recovery tool. In Symposium on the state of the science of
Animal Manure and Waste Management. pp. 5−7.
6. Kroeckel, S., Harjes, A. G., Roth, I., Katz, H., Wuertz, S.,
Susenbeth, A., & Schulz, C. (2012). When a turbot catches a fly:
Evaluation of a pre-pupae meal of the Black Soldier Fly
(Hermetia illucens) as fish meal substitute—Growth performance
and chitin degradation in juvenile turbot (Psetta maxima).
Aquaculture, 364, 345−352.
7. Lord, W. D., Lee Goff, M., Adkins, T. R., & Haskell, N. H.
(1994). The black soldier fly Hermetia illucens (Diptera:
Stratiomyidae) as a potential measure of human postmortem
interval: observations and case histories. Journal of Forensic
Sciences, 39, 215−215.
8. Pujol‐Luz, J. R., Francez, P. A. D. C., Ururahy‐Rodrigues, A., &
Constantino, R. (2008). The Black Soldier‐fly, Hermetia illucens
(Diptera, Stratiomyidae), Used to Estimate the Postmortem
Interval in a Case in Amapá State, Brazil*.Journal of Forensic
Sciences, 53(2), 476-478.

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taxo4254

9. Diener, S., Zurbrügg, C., Roa Gutiérrez, F., Nguyen, D. H.,

Morel, A., Koottatep, T., & Tockner, K. (2011). Black soldier fly
larvae for organic waste treatment – prospects and constraints.
WasteSafe 2011 – 2nd Int. Conf. on Solid Waste Management in
the Developing Countries, Khulna, 13-15 February 2011,
Bangladesh: ResearchGate.
10. Benelli, G., Canale, A., Raspi, A., & Fornaciari, G. (2014). The
death scenario of an Italian Renaissance princess can shed light
on a zoological dilemma: did the black soldier fly reach Europe
with Columbus?. Journal of Archaeological Science, 49,
203-205.
11. Woodley, N. E., & Thompson, F. C. (2001). A world catalog of
the Stratiomyidae (Insecta: Diptera) (Vol. 11). California: North
American Dipterists' Society. pp 473.
12. Roháček, J., & Hora, M. (2013). A northernmost European
record of the alien black soldier fly Hermetia illucens (Linnaeus,
1758)(Diptera: Stratiomyidae)/Nejsevernější evropský výskyt
nepůvodní bráněnky Hermetia illucens (Linnaeus, 1758)(Diptera:
Stratiomyidae). Casopis slezskeho zemskeho muzea (A), 62(2),
101-106.
13. Diclaro II, J. W. ; Kaufman, P. E. (2010). Black soldier fly
Hermetia illucens Linnaeus (Insecta: Diptera: Stratiomyidae).
EENY-461, Entomology and Nematology Department, Florida
Cooperative Extension Service, Institute of Food and
Agricultural Sciences, University of Florida.
14. Furman, D. P., Young, R. D., & Catts, P. E. (1959). Hermetia
illucens (Linnaeus) as a factor in the natural control of Musca
domestica Linnaeus.Journal of Economic Entomology, 52(5),
917–921.
15. Sheppard, C. (1983). House fly and lesser fly control utilizing the
black soldier fly in manure management systems for caged laying

page 33 / 41
taxo4254

hens. Environmental Entomology, 12(5), 1439–1442.

16. Sheppard, D. C., Newton, G. L., Thompson, S., Davis, J.,
Gascho, G., & Bramwell, K. (1998). 1998 Annual Report of
Sustainable Agriculture Research and Education: Using soldier
flies as a manure management tool for volume reduction, house
fly control and reduction, house fly control and feedstuff
production. Retrieved 7 Nov 2015 from
https://www.cals.ncsu.edu/waste_mgt/smithfield_projects/phase2
report05/cd,web%20files/A2.pdf
17. Axtell, R. C., & Arends, J. J. (1990). Ecology and management
of arthropod pests of poultry. Annual review of entomology,
35(1), 101–126.
18. Bradley, S. W., & Sheppard, D. C. (1984). House fly oviposition
inhibition by larvae of Hermetia illucens, the black soldier fly.
Journal of chemical ecology, 10(6), 853–859.
19. Erickson, M. C., Islam, M., Sheppard, C., Liao, J., & Doyle, M.
P. (2004). Reduction of Escherichia coli O157: H7 and
Salmonella enterica serovar enteritidis in chicken manure by
larvae of the black soldier fly. Journal of Food Protection®,
67(4), 685–690.
20. Liu, Q., Tomberlin, J. K., Brady, J. A., Sanford, M. R., & Yu, Z.
(2008). Black soldier fly (Diptera: Stratiomyidae) larvae reduce
Escherichia coli in dairy manure. Environmental entomology,
37(6), 1525–1530.
21. Kunzler, C., & Roe, R. (1995). Food service composting projects
on the rise: Commercial & institutional organics. Biocycle, 36(4),
64–71.
22. Diener, S., Zubrügg, C., Trockner, K. (2009). Conversion of
organic material by black soldier fly larvae: establishing optimal
feeding rates. Waste Management & Research, 27, 603–610.
23. Li, Q., Zheng, L., Qiu, N., Cai, H., Tomberlin, J. K., & Yu, Z.

page 34 / 41
taxo4254

(2011). Bioconversion of dairy manure by black soldier fly

(Diptera: Stratiomyidae) for biodiesel and sugar production.
Waste Management, 31(6), 1316-1320.
24. Barry, T. (2004). Evaluation of the Economic, Social, and
Biological Feasibility of Bioconverting Food Wastes with the
Black Soldier Fly (Hermetia illucens). University of North Texas,
Texas, United States of America.
25. Chiam, A (2014). Composting Idea Take Flight in USP and
UTown. National University of Singapore. Retrieved 7 Nov 2015
from
http://www.usp.nus.edu.sg/highlights/2014/project-herme.html.
26. Regost, C., Arzel, J., & Kaushik, S. J. (1999). Partial or total
replacement of fish meal by corn gluten meal in diet for turbot
(Psetta maxima). Aquaculture,180(1), 99–117.
27. Slawski, H., Adem, H., Tressel, R.-P., Wysujack, K.,
Kotzamanis, & Y., Schulz, C. (2011). Austausch von Fischmehl
durch Rapsproteinkonzentrat in Futtermitteln für Steinbutt (Psetta
maxima L.). Zuchtungskunde, 83(6), 451–460.
28. Bondari, K., & Sheppard, D. C. (1981). Soldier fly larvae as feed
in commercial fish production. Aquaculture, 24, 103–109.
29. Bondari, K., & Sheppard, D. C. (1987). Soldier fly, Hermetia
illucens L., larvae as feed for channel catfish, Ictalurus punctatus
(Rafinesque), and blue tilapia, Oreochromis aureus
(Steindachner). Aquaculture Research, 18(3), 209–220.
30. Calci-Worms (2015). Retrieved November 9, 2015 from
http://www.silkwormstore.co.uk/calci-worms---explained.html.
31. Phoenix Worm Store (2015). Retrieved 9 Nov 2015 from
http://www.phoenixworm.com/.
32. Welcome to ReptiWorms! - ReptiWorms (2015). Retrieved 9
Nov 2015 from http://www.reptiworms.com/index.php?index.
33. Bentley. (2013). “Yellow Soldier Flies” – Revisited. Retrieved

page 35 / 41
taxo4254

November 20, 2015, from

http://www.redwormcomposting.com/bsfl/yellow-soldier-flies-re
visited/
34. Black vs. yellow soldier flies. (2014). Retrieved November 20,
2015, from
http://www.waldeneffect.org/blog/Black_vs._yellow_soldier_flie
s/
35. Sepulveda, T. (2013). Composting Using Black Soldier Flies
(BSF) and Their Effect On Earth Worm Bins | Ed That Matters.
Retrieved November 20, 2015, from
http://www.edthatmatters.com/composing-using-black-soldier-fli
es-bsf-and-their-effect-on-earth-worm-bins/
36. Sheppard, D. C., Tomberlin, J. K., Joyce, J. A., Kiser, B. C., &
Sumner, S. M. (2002). Rearing methods for the black soldier fly
(Diptera: Stratiomyidae) in a colony.Journal of Medical
Entomology, 39(4), 695−698.
37. St‐Hilaire, S., Cranfill, K., McGuire, M. A., Mosley, E. E.,
Tomberlin, J. K., Newton, L., Sealey, W., Sheppard, C., &
Irving, S. (2007). Fish Offal Recycling by the Black Soldier Fly
Produces a Foodstuff High in Omega‐3 Fatty Acids. Journal of
the World Aquaculture Society, 38(2), 309–313.
38. Hem, S., Toure, S., Sagbla, C., & Legendre, M. (2008).
Bioconversion of palm kernel meal for aquaculture: Experiences
from the forest region (Republic of Guinea). African Journal of
Biotechnology, 7(8).
39. Martínez-Sánchez, A., Magaña, C., Saloña, & M. & Rojo, S.
(2011). First record of Hermetia illucens (Diptera: Stratiomyidae)
on human corpses in Iberian Peninsula. Forensic Science
International, 206(1), 76–78.
40. Tomberlin, J. K., Sheppard, D. C., & Joyce, J. A. (2002).
Selected life-history traits of black soldier flies (Diptera:

page 36 / 41
taxo4254

Stratiomyidae) reared on three artificial diets. Annals of the

Entomological Society of America, 95(3), 379–386.
41. Tomberlin, J. K., & Sheppard, D. C. (2001). Lekking behavior of
the black soldier fly (Diptera: Stratiomyidae). Florida
Entomologist, 84(4), 729.
42. Tomberlin, J. K., & Sheppard, D. C. (2002). Factors influencing
mating and oviposition of black soldier flies (Diptera:
Stratiomyidae) in a colony. Journal of Entomological Science,
37(4), 345–352.
43. Booth, D. C., & Sheppard, C. (1984). Oviposition of the black
soldier fly, Hermetia illucens (Diptera: Stratiomyidae): eggs,
masses, timing, and site characteristics. Environmental
entomology, 13(2), 421–423.
44. Holmes, L. (2010). Role of Abiotic Factors on the Development
and Life History of the Black Soldier Fly, Hermetia illucens (L.)
(Diptera: Stratiomyidae) (Unpublished master’s thesis).
University of Windsor, Ontario, Canada.
45. Tomberlin, J. K. (2001). Biological, behavioral, and toxicological
studies on the black soldier fly (Diptera: Stratiomyidae)
(Unpublished doctoral dissertation). University of Georgia,
Athens, United States of America.
46. Sheppard, D. C., Newton, G. L., Thompson, S. A., & Savage, S.
(1994). A value added manure management system using the
black soldier fly. Bioresource Technology, 50(3), 275–279.
47. Tran, G., Gnaedinger, C., & Mélin, C. (2015) Black soldier fly
larvae (Hermetia illucens). Feedipedia, a programme by INRA,
CIRAD, AFZ and FAO. Retrieved 7 Nov 2015 from
http://www.feedipedia.org/node/16388.
48. Hermetia illucens (2015). Integrated Taxonomic Information
System on-line database. Retrieved 7 Nov 2015 from
http://www.itis.gov.

page 37 / 41
taxo4254

49. James, M. T. (1936). The Stratiomyidae of Colorado and Utah.

Journal of the Kansas Entomological Society, 9(1), 33-36.
50. Brammer, C. A., & von Dohlen, C. D. (2007). Evolutionary
history of Stratiomyidae (Insecta: Diptera): The molecular
phylogeny of a diverse family of flies. Molecular Phylogenetics
and Evolution, 43(2), 660–673.
51. Wiegmann, B. M., Yeates, D. K., Thorne, J. L., & Kishino, H.
(2003). Time flies, a new molecular time-scale for brachyceran
fly evolution without a clock.Systematic Biology, 52(6),
745–756.
52. Wiegmann, B. M., Trautwein, M. D., Winkler, I. S., Barr, N. B.,
Kim, J. W., Lambkin, C., Bertone, M. A., Cassel, B. K., Bayless,
K. M., Heimberg, A. M., Wheeler, B. M., Peterson, K. J., Pape,
T., Sinclar, B. J., Skevington, J. H., Blagoderov, V., Caravas, J.,
Kutty, S. N., Schmidt-Ott, U., Kampmeier, G. E., Thompson, F.
C., Grimaldi, D. A., Beckenbach, A. T., Courtney, G. W.,
Friedrich, M., Meier, R, & Yeates, D. K. (2011). Episodic
radiations in the fly tree of life. Proceedings of the National
Academy of Sciences, 108(14), 5690–5695.

Acknowledgement
I would like to express my heartfelt gratitude to the following
individuals, without which the website would not have been made
possible:

Mr. Ang Yuchen who gave invaluable advice and guidance in

imaging.

Mr. Foo Maosheng who had allowed me to take specimens from

page 38 / 41
 taxo4254

his Black Soldier Fly culture for imaging and for sharing with me
his own images and expertise in imaging.

Mr. James Hii who offered help and guidance in editing the
images.

Mr. Theodore Lee who helped me in adult Black Soldier Fly

Photography.

Mr Samuel Ewing who kindly allowed me to use his image of

Hermetia illucens.

Mdm. Sally G. Miller  who kindly allowed me to use her image of Ptecticus.

Mr. Stephen Cresswell who kindly allowed me to use his image

of Trypoxylon politum.

Contact Me
Jonathan Tan is contactable at jonathan.twt@gmail.com
1. Jacinto, V. & Coin, P. eds. (2015). "Hermetia illucens." Encyclopedia of Life. Retrieved 7 Nov 2015 from
http://eol.org/pages/824054/overview. 
2. Hardouin, J. & Mahoux, G. (2003). Zootechnie d’insectes – Elevage et utilisation au bénéfice de l'homme et de certains animaux.
Bureau pour l’Echange et la Distribution de l’Information sur le Mini-élevage (BEDIM), 164 p. 
3. Kim, J. I. (1997). Newly recording two exotic insects species from Korea. J. Kor. Biota, 2, 223−225. 
4. Rozkosný, R. (1983). Subfamily Hermetiinae. In A Biosystematic Study of the European Stratiomyidae (Diptera): Volume
2-Clitellariinae, Hermediinae, Pachygasterinae and Bibliography (Vol. 2). London: Springer Science & Business Media.
pp.431.
5. Newton, G. L., Sheppard, D. C., Watson, D. W., Burtle, G. J., Dove, C. R., Tomberlin, J. K., & Thelen, E. E. (2005). The black

page 39 / 41
 taxo4254

soldier fly, Hermetia illucens, as a manure management/resource recovery tool. In Symposium on the state of the science of
Animal Manure and Waste Management. pp. 5−7.
6. Kroeckel, S., Harjes, A. G., Roth, I., Katz, H., Wuertz, S., Susenbeth, A., & Schulz, C. (2012). When a turbot catches a fly:
Evaluation of a pre-pupae meal of the Black Soldier Fly (Hermetia illucens) as fish meal substitute—Growth performance and
chitin degradation in juvenile turbot (Psetta maxima). Aquaculture, 364, 345−352.
7. Lord, W. D., Lee Goff, M., Adkins, T. R., & Haskell, N. H. (1994). The black soldier fly Hermetia illucens (Diptera:
Stratiomyidae) as a potential measure of human postmortem interval: observations and case histories. Journal of Forensic
Sciences, 39, 215−215.
8. Pujol‐Luz, J. R., Francez, P. A. D. C., Ururahy‐Rodrigues, A., & Constantino, R. (2008). The Black Soldier‐fly, Hermetia
illucens (Diptera, Stratiomyidae), Used to Estimate the Postmortem Interval in a Case in Amapá State, Brazil*.Journal of
Forensic Sciences, 53(2), 476-478.
9. Diener, S., Zurbrügg, C., Roa Gutiérrez, F., Nguyen, D. H., Morel, A., Koottatep, T., & Tockner, K. (2011). Black soldier fly
larvae for organic waste treatment – prospects and constraints. WasteSafe 2011 – 2nd Int. Conf. on Solid Waste Management
in the Developing Countries, Khulna, 13-15 February 2011, Bangladesh: ResearchGate.
10. Woodley, N. E., & Thompson, F. C. (2001). A world catalog of the Stratiomyidae (Insecta: Diptera) (Vol. 11). California:
North American Dipterists' Society. pp 473.
11. Roháček, J., & Hora, M. (2013). A northernmost European record of the alien black soldier fly Hermetia illucens (Linnaeus,
1758)(Diptera: Stratiomyidae)/Nejsevernější evropský výskyt nepůvodní bráněnky Hermetia illucens (Linnaeus,
1758)(Diptera: Stratiomyidae). Casopis slezskeho zemskeho muzea (A), 62(2), 101-106.
12. Diclaro II, J. W. ; Kaufman, P. E. (2010). Black soldier fly Hermetia illucens Linnaeus (Insecta: Diptera: Stratiomyidae).
EENY-461, Entomology and Nematology Department, Florida Cooperative Extension Service, Institute of Food and
Agricultural Sciences, University of Florida.
13. Sheppard, D. C., Tomberlin, J. K., Joyce, J. A., Kiser, B. C., & Sumner, S. M. (2002). Rearing methods for the black soldier fly
(Diptera: Stratiomyidae) in a colony.Journal of Medical Entomology, 39(4), 695−698.
14. Martínez-Sánchez, A., Magaña, C., Saloña, & M. & Rojo, S. (2011). First record of Hermetia illucens (Diptera: Stratiomyidae)
on human corpses in Iberian Peninsula. Forensic Science International, 206(1), 76–78.
15. Hem, S., Toure, S., Sagbla, C., & Legendre, M. (2008). Bioconversion of palm kernel meal for aquaculture: Experiences from
the forest region (Republic of Guinea). African Journal of Biotechnology, 7(8).
16. St‐Hilaire, S., Cranfill, K., McGuire, M. A., Mosley, E. E., Tomberlin, J. K., Newton, L., Sealey, W., Sheppard, C., & Irving, S.
(2007). Fish Offal Recycling by the Black Soldier Fly Produces a Foodstuff High in Omega‐3 Fatty Acids. Journal of the
World Aquaculture Society, 38(2), 309–313.
17. Tomberlin, J. K., Sheppard, D. C., & Joyce, J. A. (2002). Selected life-history traits of black soldier flies (Diptera:
Stratiomyidae) reared on three artificial diets. Annals of the Entomological Society of America, 95(3), 379–386. 
18. Tomberlin, J. K., & Sheppard, D. C. (2001). Lekking behavior of the black soldier fly (Diptera: Stratiomyidae). Florida
Entomologist, 84(4), 729.
19. Booth, D. C., & Sheppard, C. (1984). Oviposition of the black soldier fly, Hermetia illucens (Diptera: Stratiomyidae): eggs,
masses, timing, and site characteristics. Environmental entomology, 13(2), 421–423.
20. Tomberlin, J. K., & Sheppard, D. C. (2002). Factors influencing mating and oviposition of black soldier flies (Diptera:
Stratiomyidae) in a colony. Journal of Entomological Science, 37(4), 345–352.
21. Furman, D. P., Young, R. D., & Catts, P. E. (1959). Hermetia illucens (Linnaeus) as a factor in the natural control of Musca
domestica Linnaeus.Journal of Economic Entomology, 52(5), 917–921.
22. Holmes, L. (2010). Role of Abiotic Factors on the Development and Life History of the Black Soldier Fly, Hermetia illucens
(L.) (Diptera: Stratiomyidae) (Unpublished master’s thesis). University of Windsor, Ontario, Canada. 
23. Tomberlin, J. K. (2001). Biological, behavioral, and toxicological studies on the black soldier fly (Diptera: Stratiomyidae)
(Unpublished doctoral dissertation). University of Georgia, Athens, United States of America. 
24. Sheppard, D. C., Newton, G. L., Thompson, S. A., & Savage, S. (1994). A value added manure management system using
the black soldier fly. Bioresource Technology, 50(3), 275–279.
25. Barry, T. (2004). Evaluation of the Economic, Social, and Biological Feasibility of Bioconverting Food Wastes with the Black
Soldier Fly (Hermetia illucens). University of North Texas, Texas, United States of America.
26. Tran, G., Gnaedinger, C., & Mélin, C. (2015) Black soldier fly larvae (Hermetia illucens). Feedipedia, a programme by

page 40 / 41
 taxo4254

INRA, CIRAD, AFZ and FAO. Retrieved 7 Nov 2015 from http://www.feedipedia.org/node/16388.

27. Regost, C., Arzel, J., & Kaushik, S. J. (1999). Partial or total replacement of fish meal by corn gluten meal in diet for turbot
(Psetta maxima). Aquaculture,180(1), 99–117.
28. Slawski, H., Adem, H., Tressel, R.-P., Wysujack, K., Kotzamanis, & Y., Schulz, C. (2011). Austausch von Fischmehl
durch Rapsproteinkonzentrat in Futtermitteln für Steinbutt (Psetta maxima L.). Zuchtungskunde, 83(6), 451–460.
29. Bondari, K., & Sheppard, D. C. (1981). Soldier fly larvae as feed in commercial fish production. Aquaculture, 24, 103–109.
30. Bondari, K., & Sheppard, D. C. (1987). Soldier fly, Hermetia illucens L., larvae as feed for channel catfish, Ictalurus punctatus
(Rafinesque), and blue tilapia, Oreochromis aureus (Steindachner). Aquaculture Research, 18(3), 209–220.
31. Phoenix Worm Store (2015). Retrieved 9 Nov 2015 from http://www.phoenixworm.com/
32. Welcome to ReptiWorms! - ReptiWorms (2015). Retrieved 9 Nov 2015 from http://www.reptiworms.com/index.php?index
33. Calci-Worms (2015). Retrieved November 9, 2015.
34. Sheppard, D. C., Newton, G. L., Thompson, S., Davis, J., Gascho, G., & Bramwell, K. (1998). 1998 Annual Report of
Sustainable Agriculture Research and Education: Using soldier flies as a manure management tool for volume reduction, house
fly control and reduction, house fly control and feedstuff production. Retrieved 7 Nov 2015 from
https://www.cals.ncsu.edu/waste_mgt/smithfield_projects/phase2report05/cd,web%20files/A2.pdf 
35. Sheppard, C. (1983). House fly and lesser fly control utilizing the black soldier fly in manure management systems for
caged laying hens. Environmental Entomology, 12(5), 1439–1442.
36. Axtell, R. C., & Arends, J. J. (1990). Ecology and management of arthropod pests of poultry. Annual review of entomology,
35(1), 101–126.
37. Bradley, S. W., & Sheppard, D. C. (1984). House fly oviposition inhibition by larvae of Hermetia illucens, the black soldier fly.
Journal of chemical ecology, 10(6), 853–859.
38. Erickson, M. C., Islam, M., Sheppard, C., Liao, J., & Doyle, M. P. (2004). Reduction of Escherichia coli O157: H7 and
Salmonella enterica serovar enteritidis in chicken manure by larvae of the black soldier fly. Journal of Food Protection®,
67(4), 685–690. 
39. Liu, Q., Tomberlin, J. K., Brady, J. A., Sanford, M. R., & Yu, Z. (2008). Black soldier fly (Diptera: Stratiomyidae) larvae
reduce Escherichia coli in dairy manure. Environmental entomology, 37(6), 1525–1530.
40. Kunzler, C., & Roe, R. (1995). Food service composting projects on the rise: Commercial & institutional organics. Biocycle,
36(4), 64–71.
41. Diener, S., Zubrügg, C., Trockner, K. (2009). Conversion of organic material by black soldier fly larvae: establishing optimal
feeding rates. Waste Management & Research, 27, 603–610. 
42. Chiam, A (2014). Composting Idea Take Flight in USP and UTown. National University of Singapore. Retrieved 7 Nov
2015 from http://www.usp.nus.edu.sg/highlights/2014/project-herme.html. 
43. Hermetia illucens (2015). Integrated Taxonomic Information System on-line database. Retrieved 7 Nov 2015
from http://www.itis.gov. 
44. James, M. T. (1936). The Stratiomyidae of Colorado and Utah. Journal of the Kansas Entomological Society, 9(1), 33-36.
45. Brammer, C. A., & von Dohlen, C. D. (2007). Evolutionary history of Stratiomyidae (Insecta: Diptera): The molecular
phylogeny of a diverse family of flies. Molecular Phylogenetics and Evolution, 43(2), 660–673.
46. Wiegmann, B. M., Yeates, D. K., Thorne, J. L., & Kishino, H. (2003). Time flies, a new molecular time-scale for brachyceran
fly evolution without a clock.Systematic Biology, 52(6), 745–756.
47. Wiegmann, B. M., Trautwein, M. D., Winkler, I. S., Barr, N. B., Kim, J. W., Lambkin, C., Bertone, M. A., Cassel, B. K.,
Bayless, K. M., Heimberg, A. M., Wheeler, B. M., Peterson, K. J., Pape, T., Sinclar, B. J., Skevington, J. H., Blagoderov, V.,
Caravas, J., Kutty, S. N., Schmidt-Ott, U., Kampmeier, G. E., Thompson, F. C., Grimaldi, D. A., Beckenbach, A. T., Courtney,
G. W., Friedrich, M., Meier, R, & Yeates, D. K. (2011). Episodic radiations in the fly tree of life. Proceedings of the National
Academy of Sciences, 108(14), 5690–569

"Calciworms" (United Kingdom)

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