Comadia redtenbacheri (Lepidoptera: Cossidae) and Aegiale hesperiaris (Lepidoptera: Hesperiidae), two important edible insects of Agave salmiana (Asparagales: Asparagaceae): a review

Abstract

This review focuses on the characteristics of the life-cycle, and the nutritional properties, preservation, and marketing of Comadia redtenbacheri (Hammerschmidt) (red agave worm) and Aegiale hesperiaris (Walker) (white maguey worm) as alimentary options with nutritional benefits for humans. These insects are widely consumed in Mexico, the highest-ranking country worldwide in terms of insect diversity, and a place where entomophagy has been practiced for thousands of years, with the red and white maguey worms standing out as two insect pests of the Agave salmiana Otto ex Salm-Dyck. These insects are consumed in their larval stage in various culinary dishes by local people and foreign tourists due to their exquisite, exotic flavor, high protein and fat content, easy digestion, and high content of essential minerals for food metabolism. These properties make them a food source that offers consumers significant health benefits, but their production is seasonal and knowledge of their life-cycle scarce, so production under greenhouse conditions is unfeasible to date, though studies are ongoing to generate mass reproduction strategies under sustainable greenhouse conditions that would promote the incorporation of these insects into functional foods. For maguey worms to become a viable food option, consumers must be informed of the benefits of adding them to their diet and learn how best to integrate them into their foods. Legislation is also required to regulate their production, consumption, and health benefits.

Full text

1 23
International Journal of Tropical
Insect Science
e-ISSN 1742-7592
Int J Trop Insect Sci
DOI 10.1007/s42690-020-00396-1
Comadia redtenbacheri (Lepidoptera:
Cossidae) and Aegiale hesperiaris
(Lepidoptera: Hesperiidae), two important
edible insects of Agave salmiana
(Asparagales: Asparagaceae): a review
Aracely Molina-Vega, Edna María
Hernández-Domínguez, Matilde Villa-
García, et al.

1 23
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MINI-REVIEW
Comadia redtenbacheri (Lepidoptera: Cossidae) and Aegiale
hesperiaris (Lepidoptera: Hesperiidae), two important edible insects
of Agave salmiana (Asparagales: Asparagaceae): a review
Aracely Molina-Vega
1
&Edna María Hernández-Domínguez
2
&Matilde Villa-García
2
&Jorge Álvarez-Cervantes
2
Received: 29 May 2020 /Accepted: 26 November 2020
#African Association of Insect Scientists 2021
Abstract
This review focuses on the characteristics of the life-cycle, and the nutritional properties, preservation, and marketing of Comadia
redtenbacheri (Hammerschmidt) (red agave worm) and Aegiale hesperiaris (Walker) (white maguey worm) as alimentary
options with nutritional benefits for humans. These insects are widely consumed in Mexico, the highest-ranking country
worldwide in terms of insect diversity, and a place where entomophagy has been practiced for thousands of years, with the
red and white maguey worms standing out as two insect pests of the Agave salmiana Otto ex Salm-Dyck. These insects are
consumed in their larval stage in various culinary dishes by local people and foreign tourists due to their exquisite, exotic flavor,
high protein and fat content, easy digestion, and high content of essential minerals for food metabolism. These properties make
them a food source that offers consumers significant health benefits, but their production is seasonal and knowledge of their life-
cycle scarce, so production under greenhouse conditions is unfeasible to date, though studies are ongoing to generate mass
reproduction strategies under sustainable greenhouse conditions that would promote the incorporation of these insects into
functional foods. For maguey worms to become a viable food option, consumers must be informed of the benefits of adding
them to their diet and learn how best to integrate them into their foods. Legislation is also required to regulate their production,
consumption, and health benefits.
Keywords Lepidoptera .Larvae .Life-cycle .Agave .Edible insects
Introduction
Insect consumption in Mexico dates back to pre-Hispanic cul-
tures such as the Zapotec, Mixtec, and Mayan, which gathered
them for their exquisite, unique flavor and used them to elab-
orate various foods. Their use from ancient times to the pres-
ent places Mexico as the nation with the greatest number of
species consumed (549), followed by China with 200 (Van
Huis et al. 2013). In recent years, the taste for consuming
insects has increased in Mexico due to the broad variety of
native edible species, especially in the states of Oaxaca,
Guerrero, Chiapas, Campeche, Puebla, Hidalgo, and
Tlaxcala, where entomophagy is a common practice (Hurd
et al. 2019). The species consumed include Chorthippus
brunneus (Thunberg) (Orthoptera: Acrididae), Tenebrio
molitor (Linnaeus) (Coleoptera: Tenebrionidae), Atta
mexicana (Smith) (Hymenoptera: Formicidae), Liometopum
apiculatum (Mayr) (Hymenoptera: Formicidae), and the lar-
vae of Aegiale hesperiaris (Walker) (white maguey worm)
and Comadia redtenbacheri (Hammerschmidt) (red agave
worm) (Melgar-Lalanne et al. 2019). These insects can be
consumed in their natural state due to their pleasant flavor,
but cooking (toasted, roasted, or as a condiment) enriches that
flavor. These delicacies have begun to appear on restaurant
menus and in ethnic festivals and markets, where they have
gained popularity among middle-and upper-class urban tour-
ists, and enjoy demand in foreign markets that recognize their
commercial and food value (Miranda-Perkins et al. 2013).
Given these attributes, A. hesperiaris and C. redtenbacheri
*Jorge Álvarez-Cervantes
jorge_ac85@upp.edu.mx
1
Maestría en Biotecnología, Universidad Politécnica de Pachuca,
Carretera Pachuca-Cd. Sahagún km 20, Ex Hacienda de Santa
Bárbara, CP-43830 Zempoala, Hidalgo, Mexico
2
Cuerpo Académico Manejo de Sistemas Agrobiotecnológicos
Sustentables, Universidad Politécnica de Pachuca, Carretera
Pachuca-Cd. Sahagún km 20, Ex Hacienda de Santa Bárbara,
CP-43830 Zempoala, Hidalgo, Mexico
International Journal of Tropical Insect Science
https://doi.org/10.1007/s42690-020-00396-1
Author's personal copy

have the potential to be commercialized in European markets
under the denomination “new foods”that the European Union
and its council established in recent regulations (EU,
2015/2283) and which include the category of insects, or parts
of insects, as foods with nutritional benefits (Garofalo et al.
2017).
Specifically, these two insects have gastronomic impor-
tance due to their ancestral use and exquisite flavor. They
are consumed as main dishes or as garnishes, and can be
prepared as snacks, powders, or condiments, fried or grilled
to be eaten with nixtamalized tortillas, or incorporated into
sauces. Their salty taste and crispy texture place them in the
flavor range that consumers deem “exotic”(Ramos-Elorduy
2006; Ramos-Elorduy et al. 2011; Melgar-Lalanne et al. 2019;
Van Huis et al. 2013;Pomaetal.2017). But these larvae are
also –famously–added to the traditional Mexican alcoholic
beverage, mezcal (Van Huis et al. 2013;Pomaetal.2017).
They are processed to produce worm salt, a condiment made
from dried, roasted and ground maguey worms mixed with
common salt and chili powder (typically chili de árbol or
guajillo). The result is a fine powder with a concentrated fla-
vor in yellow and red tones that is considered a unique ingre-
dient worldwide (Ramos-Rostro et al. 2016).
The objectives of this review, therefore, are to establish the
main morphological characteristics, life-cycle features, nutri-
tional importance, and value of agave worms, and to describe
the aspects of food safety, preservation, and commercializa-
tion to enrich our knowledge of these larvae as a food source
with nutritional benefits for humans.
Comadia redtenbacheri
Commonly known as the red magueyworm in its larval stage,
this insect makes its gallery inside the stem of agaves where it
completes its life-cycle. The main host species are the
pulquero maguey Agave salmiana Otto ex Salm-Dyck,
A. mapisaga Trel., and A. atrovirens Karw. ex Salm
(Camacho et al. 2003; Llanderal-Cázares et al. 2010;
Delgado-Tejeda et al. 2017). In this stage, it is used as a food
in traditional gastronomy and is exploited as a source of sea-
sonal income for people in communities in arid and semi-arid
regions of several states in central Mexico, especially the
states of Hidalgo, Puebla, Tlaxcala, Querétaro, San Luis
Potosí, Jalisco, Oaxaca, Chiapas, and even Mexico City
(Granados 1993), where wild populations are gathered inten-
sively (Miranda-Perkins et al. 2013).
Comadia redtenbacheri is a ditrysia lepidoptera that be-
longs to the family Cossidae, sub-family Chilecomadiinae,
also called gusano rojo de maguey, chinicuil, or chilocuil in
Spanish, from the Nahuatl words chilo =pepperandocuilin =
worm; hence, the “chili worm”. During infestation of agave
plants, the female moth deposits masses of soft-textured eggs
at the base of the plant’s leaves, usually around 118 eggs that
will incubate for 30–35 days. Observations show that females
that have not mated may deposit infertile eggs. She will die
just a few hours after depositing her eggs (Camacho et al.
2003; Llanderal-Cázares et al. 2007). The morphology of the
eggs shows a surface ornamented with polygons of near con-
stant size, whose color changes from white to brown due to a
secretion that the female generates during deposition which
may provide strategic adherence, a physical barrier to parasit-
oids and predators, or absorb water and reduce evaporation
(Castro-Torres and Llanderal-Cázares 2016). Hatching pro-
duces larvaethat settle in the rhizome of the agave,where they
complete their development (Granados 1993). The first instar
larva measuresaround 2 mm in length and is colorless, though
it acquires a reddish color as it matures that becomes intense in
the final instar. It also begins to secrete a characteristic odor
that remains on any surface that comes into contact with it.
The head is partially-retracted into the thorax, and the body is
slightly flattened dorsoventrally. This worm has six pairs of
well-developed stemmata arranged in the shape of a ‘2’.Its
body is covered with primary and secondary setae. The final
segment has a chitinous projection shaped like a ‘horn’or
thorn. Dark in color, it protrudes from the center of the tenth
abdominal tergite. The thoracic legs are small and composed
of coxa, femur, tibia, tarsus, and a simple tarsal nail. The
abdominal legs are very small and present in segments III-
VI and X. The square brackets are uni-ordinal and arranged
in transversal bands, except in segment X, where they are
absent (Fig. 1) (Llanderal-Cázares et al. 2017).
The red coloration of the larvae is similar to that of other
cossids, such as Cossus cossus, and other species of the genus
Comadia. It is the only species of the Cossidae family whose
larvae develop in agaves instead of tree trunks or branches
(Vergara et al. 2012; Castro-Torres and Llanderal-Cázares
2016). The larvae leave the rhizome between September and
October and dig into the soil to a depth of 3 cm, where they
remain while weaving cocoons for pupation with silk threads
and particles of the substrate (Llanderal-Cázares et al. 2017).
They complete their life-cycle there, passing through the pupa
stage when their light brown color darkens progressively until
turning almost black when the adult larva is about to emerge.
Two rows of thick spines line the dorsal region and pleura,
inclined towards the pupa’s rear. They are used to break out of
the cocoon. This worm can be distinguished from other lepi-
doptera associated with maguey plants by the underground
location of the larvae and the type of cocoon (Fig. 2)
(Castro-Torres and Llanderal-Cázares 2016). The body of
the adult larva is light brown with two whitish marks that form
an inverted ‘V’onthe rear wings. They are veryevident when
the insect is at rest. It also has a dense covering of spatulated
scales on the surface of the wings with phylliform ones on the
thorax and abdomen. Males have bipectinate antennae, while
those of females are slightly serrated. Females are generally
larger, though size varies greatly (Fig. 3)(Castro-Torresand
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Llanderal-Cázares 2016; Cardenas-Aquino et al. 2018).
Females may have eight ovaries –typical of most
lepidoptera–of the polytrophic type; another basic character-
istic of all members of this Order. Their reproductive system is
of the ditrysia type, marked by the union between the
copulatrix bursa and the oviduct through the ductus seminalis
(Kristensen 2003; Ramírez-Cruz and Llanderal-Cázares
2015). Males characteristically transfer one spermatophore
during copulation, which has led to the assumption that be-
cause of their short life-cycle these insects copulate only once
(Miranda-Perkins and Llanderal-Cázares 2013;Ramírez-Cruz
and Llanderal-Cázares 2015; Llanderal-Cázares et al. 2017).
During colonization of the agave, these insects form colo-
nies of 40–60 worms in the center of the stem near the union
with the fleshy leaves. Normally, all larvae are in the same
stage of development and of similar size because they come
from the same oviposition (Hernández-Livera et al. 2005;
Espinoza-García et al. 2018). Gathering them requires remov-
ing the leaves and the entire stem, so it kills the plant. In fact,
harvesting the red grubs often means destroying the whole
Fig. 1 aMorphology of the larva;
bparts of the thorax; ccrochets of
false legs; dspiracles
Fig. 2 Stages of C. redtenbacheri:ared worm larva; brecently-emerged pupa; cpupa on day 52; dpupa on day 107; epupa on day 137; fpupa on day 187
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plant because they usually dwell in the neck of the stem and
the base of the leaves. If they are not removed, however, the
damage they cause also ends up killing the plant (Fig. 4)
(Esparza-Frausto et al. 2008; Espinoza-García et al. 2018).
Aegiale hesperiaris
This insect is also called the maguey worm, butter worm,
maguey moth, and peca (in Spanish) (CONABIO 2019). Its
larvae develop mainly in the months of April to July in the
lower leaves of such agave species as Agave salmiana, Agave
mapisaga and Agave tequilana. They pierce the abaxial side
at the base of the agave leaves and enter the tissues there to
form their gallery (Jaimes-Rodríguez et al. 2019;Vargas-
Zuñiga et al. 2019). An evident change of coloration, wither-
ing, and slight deformation (Fig. 5) of the plant occur because
this worm feeds exclusively on maguey shoots that provide it
with all the nutrients (proteins, fats, carbohydrates, fiber, vita-
mins, macro- and micronutrients) it requires for adequate
physiological and productive development. Males can reach
a length of 7.5 cm, but females may be larger (Vargas-Zuñiga
et al. 2019).
The State of Mexico, together with Hidalgo, Tlaxcala,
Querétaro, San Luis Potosí, Oaxaca, Jalisco, and Puebla, reg-
ister the highest incidences of the wild form of these larvae.
They are white except for the head and extremities, which are
brown. These worms are extracted for both local consumption
and export at attractive prices, an activity that complements
the income of the families that collect them (Ramos-Rostro
et al. 2013).
Few studies of the life-cycle of A. hesperiaris exist, but the
descriptions available suggest that the female deposits up to
14 eggs near the base of the agave leaves in October to
November. They form white, conically-shaped clusters of 2–
5eggs(Fig.6) that measure 3 mm wide by 2 mm high and
hatch after 15–40 days, producing larvae that are collected
between May and July. Harvesters identify infected agave
plants by wilting and yellow coloration, cut them open, and
determine the depth at which the gallery is located. The worms
are then extracted using a hook. Generally, 1–3 are found per
leaf (Fig. 7) (Jaimes-Rodríguez et al. 2019; Vargas-Zuñiga
et al. 2019). Mature larvae can measure up to 7 cm long and
1.5 cm wide after passing through 5 larval stages. Pupae may
be up to 50 mm long by 15 mm wide. Upon reaching the final
instar stage, the larvae perforate the shoot and seal the hole
with silk where the butterfly will later emerge.
Fig. 3 Pupa and moth of
C. redtenbacheri:afemale pupa;
bfemale moth; cmale pupa; d
male moth
Fig. 4 Agave plant with the presence or infestation of C. redtenbacheri
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Metamorphosis develops inside the gallery, where the lar-
vae attach to the cremaster with their heads positionedtowards
the opercle to generate the pupa, which matures and is trans-
formed into the butterfly between August and September. The
wings of mature butterflies ofboth sexes are dark in color with
light spots (Fig. 8). The estimated life expectancy in this stage
of their life-cycle is 20 days (Higgins and Riley 1980;Vargas-
Zuñiga et al. 2019; Jaimes-Rodríguez et al. 2019). Female
butterflies are larger because they carry an agglomeration of
perhaps 50 eggs adhered and compacted by a sticky substance
that allows them to conserve moisture and protects them from
parasites and pathogens (Vargas-Zuñiga et al. 2019;Jaimes-
Rodríguez et al. 2019).
Nutritional importance and value of agave
worms
These worms are used in traditional gastronomy to prepare
diverse dishes that are now attractive to both native and for-
eign diners (Fig. 9). Since they are consumed, it is necessary to
conduct analyses of their chemical composition in order to
determine the proportions of macro- and micro-molecules
they contain and that contribute to their nutritional quality
(Ramos-Rostro et al. 2012)., which varies in insects in relation
to diet, stage of development, sex, species, growth environ-
ment, and the methods used to determine their chemical com-
ponents (Van Huis and Oonincx 2017). Researchers generally
agree, however, that they are extremely rich in protein (35–
65% dry basis), fat (13–33%), and vitamins (B1,B2, B6, C, D,
E, K), and contain iron, zinc, calcium, copper, phosphorus,
magnesium, and manganese (Rumpold and Schlüter 2013b;
Schluter et al. 2017;deCastroetal.2018; Kim et al. 2019).
On average, the protein content of edible insects ranges from
35 to 60% in dry weight or 10–25% in fresh weight (Melo
et al. 2011; Schluter et al. 2017). These values are higher than
those of such sources of vegetable protein as cereals, soy-
beans, and lentils (Bukkens 1997). In the larval stage,
A. hesperiaris and C. redtenbacheri can provide high-quality
proteins, fatty acids, calories, and polyunsaturated fatty acids
(PUFA), which represent the second-largest portion of nutri-
ent composition in their bodies (Rumpold and Schlüter
(2013a)).
The fatty acids of insects are generally comparable to those
of poultry and fish in terms of their degree of unsaturation, but
Fig. 5 A. salmiana with
symptoms of white worm
infestation: aaleafofA. salmiana
with symptoms of white worm
infestation; bonset of the gallery
with waste substances from
A. hesperiaris
Fig. 6 Agave salmiana Otto ex Salm-Dyck (Asparagales: Asparagaceae)
with A. hesperiaris eggs
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Fig. 7 Maguey white worm
gathered from Agave salmiana in
the municipality of Singuilucan,
Hidalgo: amaguey worm hosted
in an agave leaf; bmaguey white
worm in a gallery
Fig. 8 Stages of the life-cycle of
A. hesperiaris:awhite maguey
worm in the eighth instar; blower
part of the larva of A. hesperiaris;
clarva in the latent stage about to
enter the pupa stage; dpupa of
A. hesperiaris;epupa in meta-
morphosis with presence in the
upper part of the head and wings;
fbutterfly of A. hesperiaris in a
state of putrefaction after attack
by fungi and bacteria in the
gallery
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contain more PUFA. The mean average fat content is 41.7%,
while the caloric count is 6776kcal/kg; levels which show that
red and white agave worms have a high energy content. In
fact, they contribute 370 kcal/kg, which is 50% more than
soya, 63% more than beef, 70% more than fish, 84% more
than vegetables, and 100% more than chicken. Regarding the
nutritional and mineral composition of these two insects, both
have a high percentage of protein and fat (Ramos-Rostro et al.
2012; Rumpold and Schlüter 2013a). Their high sodium and
potassium content (6649,5 g/100 g, and 4149,8 g/100 g)
(Table 1) can help lower blood pressure and prevent arterial
hypertension, cardiopathies, and strokes in consumers when
compared to foods like beef, fish, beans, peas, and potatoes,
which range from 0.14–1.694 g/100 g. Sodium and potassium
are responsible for maintaining the body’s electrolyte balance,
which is important because an imbalance affects bone metab-
olism by modifying urinary pH and the excretion of calcium
and phosphorus through the urine (Ramos-Rostro et al. 2012).
Food safety of agave worms
In terms of food safety, studies of the external bacterial com-
munity and cultivable microbiome of C. redtenbacheri report
a greater bacterial diversity in larvae obtained from vendors
than in those collected directly from agave plants. Those stud-
ies have detected Paenibacillus sp., Bacillus safensis,
Pseudomonas sp., Bacillus pseudomycoides,
Corynebacterium variabile,Enterococcus sp., Gordonia sp.,
Acinetobacter calcoaceticus,Arthrobacter sp., Micrococcus
sp., and Bacillus cereus. It is still unclear, however, whether
these bacteria represent a potential health risk to people who
ingest the larvae of C. redtenbacheri, so it is important to
gather information on the species and strains involved to es-
tablish more accurately their potential health risks for con-
sumers (Hernández-Flores et al. 2015). Some of the bacteria
found in the larvae are known to generate compounds that are
Fig. 9 Forms of consumption of
maguey worms: afried white
maguey worm dish accompanied
by guacamole; bred worms in
salsa; cred worms in pulque; d
roasted red worm
Table 1 Nutritional composition of A. hesperiaris and C. redtenbacheri
Component A. hesperiaris C. redtenbacheri
g/100 g wet base
*
Humidity 77,15 58,3
Ash 1,05 0,87
g/100 g dry base
*
Protein 37,79 31,23
Fat 34,94 58,54
Fiber 4,22 1,85
Nitrogen-free extract 18,44 6,31
μg/g
**
Calcium 223,7 126,9
Magnesium 763,5 473,6
Sodium 72,1 175,1
Potassium 6649,5 4194,8
Zinc 39,3 43,2
Copper 11,4 7,3
Iron 0,5 0,5
Manganese 16,7 10,6
Phosphorus 0,57% 0,33%
Source: (Ramos-Rostro et al. 2012
*
; Rumpold and Schlüter 2013a
**
)
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of interest to the food industry and agriculture; for example,
Paenibacillus sp., which is important for humans, animals,
and plants. This species can be found in soils and is often
associated with plant roots where it promotes growth and pro-
duces antimicrobial peptides and enzymes that could be uti-
lized for bioremediation or to produce valuable chemical sub-
stances (Grady et al. 2016). B. safensis canbeconsidereda
safe industrial microorganism (producer of industrial enzymes
and secondary metabolites) since there is no evidence of path-
ogenicity (Lateef et al. 2015). Regarding B. pseudomycoides,
research has found that it generates a type II antibiotic, an
antimicrobial peptide capable of attacking gram-positive bac-
teria (Basi-Chipalu et al. 2015). Specific studies of the micro-
biological safety of insects as food sources in the literature are,
unfortunately, scarce (Belluco et al. 2013).
Preservation
Traditionally in Mexico, maguey worms are washed after col-
lection and consumed either fresh, fried, or roasted. Today, in
contrast, they may be frozen or dehydrated until used. The
dehydrated form does not require refrigeration to retain nutri-
tional value and good condition due to the antibiotic sub-
stances they generate while alive, which provide protection
and inhibit decomposition during dry storage (Ramos-Rostro
et al. 2012; Van Huis 2013;deCastroetal.2018; Mlcek et al.
2014).
These processes are similar to those utilized with other
insects; for instance, the microbiological content of fresh,
processed, and stored edible insects like the mealworm
(Tenebrio molitor)andcricket(Acheta domesticus and
Brachytrupes sp.) have been evaluated recently. Results indi-
cate that Enterobacteriaceae and spore-forming bacteria can
be isolated from fresh insects, but generally do not belong to
pathogenic species. Boiling these insects for 5 min is an ef-
fective method of eliminating Enterobacteriaceae but not
spore-forming bacteria, so refrigerated storage (5–7 °C) is
suggested. This temperature will also prevent the decomposi-
tion of boiled insects (stable for more than 2 weeks), but is not
efficient in preventing decomposition. Roasting alone did not
kill all the Enterobacteriaceae, so boiling for a few minutes is
suggested before roasting. These studies have also shown that
lactic acid fermentation can inactivate Enterobacteriaceae
and maintain spore-forming bacteria stable at acceptable
levels where they cannot germinate and grow (Klunder et al.
2012). The hygienic handling of insects and adequate storage
conditions are two issues that must be addressed to avoid
potential risks for human health after ingestion (Belluco
et al. 2013). This requires research to obtain information on
consumption that complies with existing norms of food safety,
which now authorize insects as an alimentary alternative.
Commercialization
Maguey worms are used as an ingredient in traditional gas-
tronomy and constitute a source of seasonal economic income
for people in arid and semi-arid regions of several states in
north-central Mexico, as mentioned above (Granados 1993),
where wild populations are gathered intensively (Miranda-
Perkins et al. 2013). In the state of Hidalgo,
C. redtenbacheri is widely-known and has an order of impor-
tance of 27.48% over A. hesperiaris (4.09%).
Commercialization of the larvae is concentrated in the rainy
season when the worms are most abundant. They can be sold
fresh at this time of year and may be ingested live almost
immediately after gathering, or be kept in refrigeration, frozen,
or processed in various forms –dehydrated, fried, boiled, for-
mulated, seasoned, or immersed in alcoholic beverages–for
consumption at any time. In the mezcal industry, a red worm
is actually placed inside the bottle as a distinguishing feature
and attraction that generates added value for the product
(Llanderal-Cázares et al. 2010). These insects can be seen
for sale in several venues: stands along streets or highways,
collection centers, local markets, town squares, and stores in
towns and local communities, but they are also marketed in
other states and exported by companies that sell them in cans
(Contreras-Frias 2013) in tourist areas where they are con-
sumed in home kitchens and restaurants.
Prices vary, depending on such factors as climatic condi-
tions, season, age of the maguey, predators (rodents, birds),
and the quality and evolution of the eggs. In addition, Ramos-
Elorduy (2006) mention a whole range of elements that affect
the commercialization of these insects subject to the rules of
supply and demand: scant knowledge of the biology of the
larvae and the care they require, parasitism of the eggs, crop
abandonment, modernization of agriculture, exposure to dis-
eases due to mismanagement, scarce maguey plantations, al-
ternative hosts, low purchase prices paid to collectors, high
demand, predatory practices, lack of control of hoarders, the
presence of intermediaries, and the absence of price
regulations.
In Mexico, the sale of maguey worms occurs at the local,
regional, and state levels. They may be sold individually (by
piece) or in containers (cans, paper cones, glass cups, plastic
tubs, bags) of different volumes (250 mL, 500 mL, 1 L) or
weight (gr or Kg). In markets, the cost of a liter of
C. redtenbacheri ranges from $41.37–68.95 USD, while a
can similar to the one used for sardines costs $22.98 USD,
and the unit cost is $0.05 USD (Pino-Moreno et al. 2020). Of
course, these prices vary depending on the region, year, com-
munity, and season (low vs. high). The prices of A. hesperiaris
and C. redtenbacheri in the high season range from $28.19–
32.89 and $23.49 USD/L respectively, while out of season the
white worm may cost as much as $46.99–56.39 USD/L and
the red worm perhaps $32.89 USD/L (Miranda et al. 2011).
Int J Trop Insect Sci
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The availability of these worms in restaurants depends on
price, which can rise to prohibitive levels due to the marked
variability in the insects’temporal or spatial distribution,
which limit their availability (Van Huis et al. 2013).
In contrast to the central regions of Mexico where the col-
lection of edible insects represents an ancestral cultural legacy
with identitary importance and a strong sense of belonging, in
places like the north-central area of Zacatecas this is a rela-
tively recent activity that began just 28 years ago. In the mu-
nicipality of Los Pinos, for example, agricultural productivity
is low, and maguey plants represent the most important natu-
ral resource in terms of abundance and variety of uses (De
Luna-Valadez et al. 2013). The lack of agricultural profitabil-
ity and income opportunities have led some people to turn to
gathering maguey as a way to increase their incomes. De
Luna-Valadez et al. (2013) report that 95.1% of collectors
are men and women who have integrated gradually into this
activity, and that 72% of people work individually, while the
rest operate as families. In Los Pinos, collection is carried out
mainly for commercial ends because the culture of local con-
sumption is limited with only 25% of people consuming
worms in small quantities.
Regarding the rate of use of plant material,
C. redtenbacheri demands a larger number of magueys: 54
plants/day compared to 50 plants/day for A. hesperiaris. As in
the central regions of Mexico, the price of maguey worms
varies from year-to-year and community-to-community, but
average prices for A. hesperiaris and C. redtenbacheri are
$15.74 and $28.19 USD/Kg, respectively. Local collectors
sell their harvests to buyers who take the product to markets
in Mexico City, neighboring states, and even to worm-
producing states in central Mexico (De Luna-Valadez et al.
2013). It is clear that the region of origin largely determines
the price of these worms, and that collectors there receive a
better return on their product than those farther north. This
disparity indicates the need to generate marketing strategies
that favor collectors, regardless of region or state, and to es-
tablish fair prices that reflect the product’s nutritional attri-
butes, flavor, and gastronomic exoticism.
The tradition and preparation of these insects have been
kept alive in rural communities over many generations down
to the present (Ramos-Elorduy et al. 2001; Shockley et al.
2018; Hurd et al. 2019). Thanks to their cost and value as
“traditional dishes”, demand for these insects has increased
considerably in recent times. Indeed, sales and consumption
of red and white agave worms in the larval stage are so high
that gathering them has become a significant seasonal eco-
nomic activity in the rainy season; one that is attractive for
many agave producers (Mendoza-Mendoza et al. 2016).
Potential export markets for these larvae could include the
United States and Asia; the former due to the presence of the
many Mexicans and other Latin Americans who are familiar
with maguey worms; the latter because their populations tend
to be more open to new, “rare”products and beliefs. In fact,
people in various cities in China, Japan, and India know
C. redtenbacheri from mezcal exports and believe it has aph-
rodisiac properties apart from its exquisite flavor (Contreras-
Frias 2013).
Without doubt, commerce in maguey worms is attractive
both nationally and internationally, but growing demand has
negatively impacted the host plants because, as mentioned
above, the larvae are collected from wild populations that
are never recovered, re-planted, or tended to. This is generat-
ing a real threat of extinction of the maguey that would mean
the end of their parasites as well (Llanderal-Cázares et al.
2010). Implementing alternative production methods like but-
terfly farms, producing magueys in vitro, and cultivating lar-
vae in greenhouses could help save the maguey and permit the
production of A. hesperiaris and C. redtenbacheri on a com-
mercial scale (Contreras-Frias 2013). Currently, research is
underway on establishing rural production units, and
C. redtenbacheri is being studied in laboratories in terms of
its potential for intensive reproduction. Llanderal-Cázares
et al. (2010), for example, tested various conditions of infes-
tation, greenhouse production, irrigation methods, and soil
moisture using larvae at different densities and distinct instars.
Their results seem promising as the highest establishment rate
was generated using spaced irrigation conditions and low lar-
va densities. They also reported that weekly irrigation gener-
ates higher grammages in the larvae. Without doubt, these
experimental data will be of great value for producing the
red agave worm in greenhouse conditions.
Mexican companies have already developed various prod-
ucts based on these insects: maguey worm salt, agave worm
powder, and red maguey worms in various presentations.
They recognize these larvae as exceptional ingredients with
high culinary and nutritional value, and are promoting the
diversification of consumption by emphasizing the value of
traditions, supporting producers, and supporting the sustain-
ability of these species and the environment (Van Huis 2013;
Van Huis et al. 2013; Melgar-Lalanne et al. 2019). These
companies have also begun to construct the value chain for
maguey worms with a view to commercializing them on a
wider scale. However, the lack of studies and detailed
information on linkages that might benefit stakeholders and
the environment impede defining these parameters. Porter
(1991) has proposed an analytical model to identify and break
down the primary and secondary activities of the value chain.
This model can be used to continue defining and refining the
value chain of maguey worms for their production and com-
mercialization. In this context, field observations and testimo-
nies from producers and sellers identify five key activities –
supply, collection, transformation, marketing, and
consumption–together with 4 secondary activities: purchas-
ing, general administration, logistics, and marketing
strategies.
Int J Trop Insect Sci
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Today, there is an urgent need to monitor, study, and gen-
erate information on the exploitation of agave worms. In terms
of economic aspects, there is almost no statistical data on the
national and international commercialization of
C. redtenbacheri and A. hesperiaris, and the few figures that
do exist are mostly descriptive and lack technical scientific
rigor. For all these reasons, it is important to begin laying
economic foundations that will promote the collection and
production of these larvae as an agricultural alternative with
the potential to benefit numerous communities in central
Mexico and other states that produce edible insects, in addi-
tion to rescuing the maguey plant itself, which is now under
threat.
Conclusions
C. retenbancheri and A. hesperiaris are two insects with po-
tential uses in the food industry due to their nutritional value,
consumer acceptance, and exquisite flavor, but this potential is
limited, primarily by their seasonal availability. In addition,
the number of insects that can becollected is decreasing due to
overexploitation of maguey plants and the activity of butterfly
or egg predators that interfere with the insects’life-cycle. A
solution for this could include: 1) comprehensive management
of magueys in protected areas; 2) the selection of specific
maguey species for infestation; and 3) implementing cultiva-
tion under greenhouse conditions to facilitate studies of the
different stages of development of the larvae and their inter-
action with the host plants. Determining the reproductive
stages of maguey worms in greater detail will allow us to
establish mass reproduction of insect populations and year-
round production. It is also necessary to investigate how sus-
tainable technologies can improve the harvesting, post-har-
vesting, and processing of these insects to generate a
bioeconomic model that would ensure the adequate incorpo-
ration of these insects into food products enriched in proteins,
fats, fibers, vitamins, and minerals, while providing consumer
satisfaction. Finally, achieving the recognition of the maguey
worms C. retenbancheri and A. hesperiaris as alimentary al-
ternatives requires informing consumers of the benefits of
their consumption and how they can be integrated into their
diet, but this will also mean enacting legislation to regulate
production, consumption, and health benefits.
Acknowledgments We thank María del Carmen Ávila Ramírez, M.Sc.,
for her work on the physiology of Comadia redtenbacheri, and Carmelita
Ramos Tecolmalman (from Tlachiquera del Tinacal los Tuzos, munici-
pality of Singuilucan, Hidalgo) for sharing their experiences and taking
samples and photos of C. redtenbacheri and A. hesperiaris for the elab-
oration of this review. We also thank Mexico’s Consejo Nacional de
Ciencia y Tecnología (CONACYT) for the scholarship awarded to the
student Aracely Molina Vega (No.858301) to support her M.Sc. studies
in Biotechnology at the Universidad Politecnica de Pachuca.
Compliance with ethical standards
Conflict of interest The authors declare that they have no conflict of
interest, and that the images included are their property. All authors col-
laborated in the search for information and in the writing of this review,
and approved their participation and the order in which they are listed.
Finally, together they assume all responsibility for this publication.
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