Archaeoentomological study of a pre-Columbian funerary bundle, Huchet and Al 2013

Annales de la Société entomologique de France (N.S.), 2013
Vol. 49, No. 3, 277–290, />
Archaeoentomological study of a pre-Columbian funerary bundle (mortuary cave of Candelaria,
Coahuila, Mexico)
Jean-Bernard Hucheta,b,c*, Grégory Pereirad, Yves Gomye, Thomas Keith Philipsf, Carlos Eduardo Alatorre-Bracamontesg,
Miguel Vásquez-Bolañosg & Josefina Mansillah
Muséum national d’Histoire naturelle, Département Systématique et Evolution (entomologie), UMR 7205 du CNRS – Origine, Structure
et Evolution de la Biodiversité. CP 50, 45 rue Buffon, F-75005 Paris Cedex 05, France; bMuséum national d’Histoire naturelle,
Département Ecologie et Gestion de la Biodiversité, UMR 7209 du CNRS – Archéozoologie, Archéobotanique : sociétés, pratiques et
environnements. CP 56, 55 rue Buffon F-75005 Paris, France; cUMR 5199 du CNRS, PACEA – Anthropologie des populations passées
et présentes, Université Bordeaux 1, Avenue des Facultés, F-33405 Talence Cedex, France; dUMR 8096 “Archéologie des Amériques”,
CNRS Maison de l’Ethnologie et de l’Archéologie, 21 allée de l’Université, F-93023, Nanterre Cedex, France; e2 boulevard Victor Hugo,
F-58000 Nevers, France; fSystematics and Evolution Laboratory, Department of Biology, Western Kentucky University, 1906 College
Heights Blvd. Bowling Green, KY 42101-3576, USA; gEntomología, Centro de Estudios en Zoología, Departamento de Botánica y
Zoología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Km. 15.5 Carr. Nogales, Las
Agujas, Zapopan, Jalisco. CP. 45110, Apdo. Postal 134, México; hDirección de Antropología Física, Museo Nacional de Antropología,
Reforma y Gandhi s/n, 11560 D.F., Colonia Polanco, México
a

(Accepté le 23 mai 2013)

Summary. The archaeoentomological study of insect remains recovered from a pre-Columbian funerary bundle (10th–11th
centuries AD) is presented and illustrated. Among this material, 12 species belonging to 10 families and four distinct orders
have been identified. From the biological data of the different taxa, some hypotheses about the funerary practices of the
hunter–gatherer semi nomads of Northern Mexico are proposed.
Résumé. Etude archéoentomologique d’un ballot funéraire précolombien (grotte sépulcrale de la Candelaria,
Mexique). L’étude archéoentomologique d’un échantillon d’insectes provenant d’un ballot funéraire précolombien (Xe–
XIe siècles A.D.) est présentée et illustrée. Au sein de cet assemblage, douze espèces appartenant à 10 familles et à 4 ordres
distincts d’insectes ont pu être identifiées. A partir des données biologiques des différents taxons en présence, des
hypothèses sur les pratiques funéraires des populations préhistoriques nomades du nord du Mexique sont proposées.
Keywords: funerary archaeoentomology; pre-Colombian America; burial practices; insect remains; Diptera; Coleoptera;

Hymenoptera; Lepidoptera

The climate in northern Mexico, combined with a karstic
environment, allows an exceptional preservation of
organic matter and other perishable materials. For two
millennia prior to the Spanish conquest (1519–1521),
numerous natural caves in northern Mexico were used as
burial sites by hunter–gatherer populations who buried
their dead in these locations over several generations. In
the dry environment of these natural sanctuaries, the
bodies placed in mortuary bundles made of plaited vegetal
materials have been preserved over the centuries, as well
as the fragile remains of necrophagous insects linked to
processes of thanatomorphosis.
The Candelaria funerary cave (Coahuila State,
Mexico) excavated in the early 1950s has provided many
human remains, often remarkably preserved. These populations have been the subject of numerous archaeological
and bioarchaeological studies (Martínez del Río 1953a,
1953b; Romano 1956; Taylor 1968; Mansilla & Pijoan
*Corresponding author. Email:

© 2013 Société entomologique de France

2000). While the presence of insects had been previously
noted, the first archeoentomological analysis based on
some of these remains was performed very recently by
Vergara-Pineda et al. (2009). These last authors provided
the first inventory of entomofauna associated with human
remains at Candelaria, basing their analysis both on a
sample recovered from an adult cranium and from the

funerary bundle of a young child. A second sample set
recovered from the same bundle is presented and illustrated here.
Beyond a strictly taxonomic interest, the study of
invertebrate necrofauna in an archaeological funerary
context also sheds light on the immediate environment
of the site at the moment of its use. From a “paleoforensic” perspective, it may provide a new insight into
the reconstruction and understanding of the funerary
practices of semi nomadic groups from northern
Mexico.


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Geographical setting

The Candelaria funerary cave is located near San Pedro de
las Colonias, in the semidesert zone of the State of
Coahuila, northern Mexico [approximately 25°01′ N,
102°46′ W] (Figure 1). It is a natural cavity that formed
in Cretaceous limestone of the Sierra de la Candelaria, a
small mountain formation that borders the Las Delicias
endorheic basin (Maldonado-Koerdell 1956). The region
is characterized by an arid climate (less than 200 mm of
annual precipitation) and Chihuahuan desert vegetation
(primarily cacti, yucca, and thorn shrubs).
The cavity occurs at 1000 m asl. It is accessible through
a shaft approximately 9 m high and 1 m wide. The funerary
deposits were located in two adjoining rooms where the
floors show a regular inclination (Figure 2).

Archaeological context
The Candelaria funerary cave is an exceptional site in
the archaeology of northern Mexico for the spectacular
state of conservation of its organic remains. The site
offers a particularly rich image of funerary practices
and the material culture of the hunter–gatherers of this

Figure 1.

Figure 2. Candelaria Cave, seen in cross section, with the
location of the funerary bundles (adapted from MaldonadoKoerdell 1956).

desert region. The cave was excavated in 1953 and 1954
by a group of Mexican archaeologists under the
Dirección de Prehistoria of the Instituto Nacional de

Geographic location of the funerary cave of Candelaria in Coahuila State, Mexico.


Annales de la Société entomologique de France (N.S.)
Antropología e Historia (Aveleyra Arroyo de Anda,
Maldonado-Koerdell and Martínez del Río), after evidence of looting activities had been reported. The
deposits have been partially disturbed prior to the beginning of the excavations. While the possibility of postdepositional contamination cannot be totally excluded,
the composition and the biology of the faunal composition presented here leads us to believe that all of these
studied insect remains are linked to processes of postmortem decay, and are therefore contemporaneous with
the deposit.
Despite the perturbations, we can discern precise information regarding the funerary practices. The cavity functioned as a collective burial place during the prehispanic
period, with at least 116 individuals (83 adults and 33
subadults) buried there. The skeletal collection comprises
mainly a deposit of mixed bones and two intact infant

mortuary bundles. Commingling of bones was caused
apparently by natural rock falls inside the cave and the
destruction of originally individual mortuary bundles by
looters. These recurring burials took place over several
generations. The bodies were brought into the cave as
mortuary bundles (Romano 1956) and placed on the
floor in the cave, without making a pit or covering them
with earth (González Arratia 2007). The corpses were
wrapped in a fetal position with pieces of cloth made
from yucca fibers maintained by a network of fine cords
(Weitlaner-Johnson 1977) (Figures 3, 4), and the bundles
were tied with a cord to keep in place the bent legs and
arms. Each bundle was placed on a scaffold comprised of

279

varied perishable materials: mats, trays, frames of wood,
and various objects (bows, digging sticks, etc.) and individually separated by vegetative matter (cactus pads,
yucca leaves, etc.).
Many other objects deposited around the deceased
were also collected (Martínez del Río 1953a, 1953b;
Maldonado-Koerdell 1956; Aveleyra Arroyo de Anda
1964; González Arratia 1999): wooden tools and hunting
weapons, arrows, hafted chert knives, basketry, sandals,
cordage, stone ornaments, shell, bone, etc. The characteristics of some of these objects and the first radiocarbon
dates obtained (Aveleyra Arroyo de Anda 1964; González
Arratia 1999, p. 48–49) suggested that the cave’s occupation occurred around the twelfth and thirteenth centuries
14
AD. New C
dating obtained by one of the authors (J.

Mansilla, in: Pineda et al. 20091) indicates that the funerary deposits are considerably more ancient (AD 940 ± 24
and 1020 ± 28).
Given the excellent conservation of the human skeletal
remains, numerous studies have been conducted (e.g. craniometry: Romano 1956; pathology: Mansilla & Pijoan
2000, 2005; Pineda et al. 2009). However, despite the
exceptional preservation of the organic material, the soft
tissues were only very rarely preserved.
Only three bundles were recovered intact: those of one
adult and two children. A radiological study carried out on
one of the two juvenile bundles by Mansilla & Pijoan
(2000) (Figure 5) determined that the deceased individual
was three years old. The second child died before he or
she reached one year of age.

Figures 3–5. 3, Funerary bundle of the youngest child where the insect remains were recovered. 4, Funerary bundle of the second
child. 5, idem, radiography showing the skeleton in anatomical position: the lower and upper limbs bent (photos Figures 3 and 4 DAF/
INAH; Figure 5 R. Enríquez DAF/INAH).


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Cultural significance of the funerary bundle in ancient
Mexico
Although the precise meaning behind the funerary bundles
created by the hunter–gatherers at Candelaria Cave
remains unknown, it is important to point out that this
practice was not only restricted to this site and culture.
The custom of wrapping a corpse within a cloth or mat

bound with cords occurred widely throughout the
Americas, from the Southwestern USA to the Andes. In
Mexico, this practice has been observed archaeologically
in the northern semi-arid regions where the climate favors
the preservation of the organic remains. While the first aim
of this system of wrapping was probably to facilitate the
transport of the dead to the burial place, the bundle is not
restricted to this single function and, in different cultures,
it also carried a symbolic meaning. Among the Aztec and
other Late Postclassic societies (AD 1200–1521), funerary
bundles were considered to represent the chrysalis of a
butterfly (Taube 2002, p. 308–309). Representations from
pictographic manuscripts (codices) show that these bundles were often decorated with butterfly imagery or with
specific anatomic attributes of the chrysalis. Those destined for flames were themselves symbolized by a butterfly, the symbol of both fire and the souls of the dead
doomed to the celestial beyond (Beutelspacher 1988,
p. 16–20). Regardless, there is no evidence to date that
the inhabitants of Candelaria Cave shared this particular
symbolism, which corresponds to very distinct groups
who did not practice cremation burials. Instead, it appears
that this form of funerary preparation served a more pragmatic purpose, such as, in the present case, facilitating the
transport of the dead to a cave that was difficult to access.

The insect remains
The insect remains studied here2 (Table 1) were recovered
from different damaged parts of the funerary bundle of the

younger child (Figure 3). The collected residues have been
processed to dry sieving (mesh size: 300 µm), examined
under a stereomicroscope and mounted according to standard entomological procedures (glued on card-points and
labeled).

The faunal composition recovered from this context is
hardly surprising since, to a large extent, the different taxa
are those found to most frequently colonize carrion. The
sampling includes 12 species belonging to 10 families and
four distinct orders (Diptera, Coleoptera, Hymenoptera
and Lepidoptera).
Diptera
The remains or “sclerites” attributable to this order (nearly
complete or fragmented puparia) correspond to three
sarco-saprophagous families of cyclorraphous flies:
Muscidae, Fanniidae and Sarcophagidae. These three
families traditionally figure among the most relevant taxa
for estimating the postmortem interval (PMI) in forensic
entomology.

Muscidae
Synthesiomyia nudiseta Van der Wulp 1883
(Figures 6, 7)
This species usually colonizes cadavers a few days after
the first settlement of necrophagous flies (Skidmore 1984).
According to Buxton & Hopkins (1927), S. nudiseta
invades carrion at the active decay stage (the “second
wave” according to the terminology of Easton & Smith
[1970]). When pupation occurs, the mature larvae exude a
frothy and sticky substance which solidifies to form a
protective cocoon upon which sand particles and other
exogenous debris adhere (Siddons & Roy 1942;
Skidmore 1984). Unlike most necrophagous fly larvae,
which migrate some distance from the food source to

Table 1. Summary of insect remains recovered from the funerary bundle sample set.
Order
Diptera
Diptera
Diptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Coleoptera
Hymenoptera
Lepidoptera

Family

Taxa

Number of insect remains

MNI*

Muscidae
Fanniidae
Sarcophagidae
Dermestidae
Histeridae
Histeridae
Histeridae

Staphylinidae
Anobiidae
Trogidae
Formicidae
Tineidae ?

Synthesiomyia nudiseta van der Wulp
Fannia sp.
Genus ? sp. ?
Dermestes (Dermestinus) carnivorus Fabricius
Saprinus (s. str.) alienus J.L. LeConte
Xerosaprinus (s. str.) coerulescens (J.L. LeConte)
Xerosaprinus (s. str.) vitiosus (J.L. LeConte)
Genus ? sp. ?
Niptus n. sp.
Omorgus sp.
Acromyrmex versicolor (Pergande)
Genus ? sp. ?
Total

> 10
10
>3
>9
1
1
1
1
> 10
2

> 10
1
> 59

4
5
3
5
1
1
1
1
8
1
3
1
34

* MNI = minimal number of individuals.


Annales de la Société entomologique de France (N.S.)
pupate, S. nudiseta usually completes the entire biological
cycle in situ (in close contact with the cadaver). In the
Neotropical region, this species among the flies is most
frequently associated with human remains in archaeological contexts (Huchet & Greenberg 2010). In this respect,
the unidentified muscid species associated with sacrificed
human bodies from Pacatnamu (Peru) mentioned by
Faulkner (1986, figure 4) may be attributed to S. nudiseta.
Field experiments conducted by one of the authors (JBH)

on the archaeological site of Huaca de la Luna (Peru)
highlighted that this species can sometimes invade carrion
early on and then replace the pioneer blowflies.
Due to its great dispersal ability, S. nudiseta is now
widely distributed throughout the warmer parts of the
globe, and sometimes very distant from the supposed
area of origin (South America). In the National
Museum of Natural History (Paris), the species is represented by specimens from various origins, including New
Caledonia, Thailand and Japan. The species has been
recently collected in Europe from both Spain (Ebejer &
Gatt 1999) and Malta (Carles-Tolrá 2002). In the field of
forensic entomology, S. nudiseta is frequently used in the
estimation of the PMI (Jirón et al. 1983; Lord et al.
1992; Oliveira-Costa et al. 2001; Calderón-Arguedas
et al. 2005; Kumara et al. 2009). A recent discovery in
a pre-Columbian grave in Peru (Mochica civilization)
(Huchet & Greenberg 2010) provided evidence of complex funeral practices, notably a long exposure of the
corpse prior to burial, as evidenced by the presence of
this species.
The characteristic shape of the anal spiracular slits
(Eldridge & James 1957; Siddons & Roy 1942; Skidmore
1984) combined with the presence of distinct remains of
sandy cocoons recovered from the sample leaves no doubt
about the specific identification of this species.

Fanniidae
Fannia sp.
(Figure 8)
The Fanniidae species usually develop in relatively
advanced cadavers (Nuorteva et al. 1974; Smith 1986;

Lee & Marzuki 1993). The immature stages of the species
belonging to the genus Fannia Robineau-Desvoidy 1830
are morphologically characteristic. The larva and puparia
are flattened dorsoventrally and bear many well-developed
plumose lateral and dorsal processes. The Neotropical
fauna includes 66 species (Carvalho et al. 2003).
Remains of Fannia canicularis (L. 1761) were
recently recovered from an Egyptian mummy of the
Ptolemaic Period (Gerisch 2001). As with the species
previously quoted, the presence of a representative of the
genus Fannia is undoubtedly linked with the postmortem
decay of the young child.

281

Sarcophagidae
Genus ? sp. ?
(Figure 9)
The Sarcophagidae or “flesh flies” includes nearly 2600
species worldwide (Pape et al. 2009). Although probably
more diversified than in any other region of the world,
little is known regarding the Neotropical fauna (Pape
1989). Unlike the Calliphoridae, the Sarcophagidae
females are larviparous and directly deposit first instar
larvae on carrion. The females usually arrive at corpses
slightly later than the pioneer blowflies, attracted by the
first odors of decay.
The identification at the family level is based on the
presence of a puparium fragment (Figure 9). Contrary to
other sarcosaprophagous diptera families, the posterior

spiracles are hidden in a crateriform-shaped cavity located
in the upper part of the last abdominal segment. No
exhaustive work on puparia is available and the only
existing reference concerning the Sarcophagidae (Greene
1925) is incomplete and now obsolete. Ten additional
puparia attributed to this family that were recovered
from the funerary bundle are mentioned by VergaraPineda et al. (2009).
Hymenoptera
Formicidae
The presence of Formicidae on carrion can be linked to
two distinct ecological habits: predation upon eggs,
larvae or pupae of thanatophagous insects, or necrophagy (ants feeding on exudates or decomposing tissues) (Moretti de Carvalho & Ribeiro 2006; Clark &
Blom 1991). When ants are found in association with
human remains, they are sometimes used as part of
forensic investigations (Goff & Win 1997; Martínez
et al. 1997). The presence of a fungus-growing ant
species within the bundle raises many questions.
Different hypotheses on how and why such colonization
occurs are proposed below.
Myrmicinae Lepeletier 1835
Attini Smith 1858
The Attini is a New World endemic tribe primarily
Neotropical in distribution which includes 95% of the
known species, with the other 5% restricted to the
Nearctic region (Fernández 2003, 2006; Mayhé-Nunes
& Jaffe 1998). Attini comprises a monophyletic group
of approximately 230 described species (Schultz &
Brady 2008) arranged in 14 distinct genera (Bolton
et al. 2006). These fungus-growing ants use the organic
detritus they collect as a substrate for their fungal cultivars, the main source of their diet (basidiomycete fungus of the genus Leucocoprinus (Pat.), notably

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Agaricaceae and Lepiotaceae). The Attini tribe includes
highly sophisticated forms including the leaf-cutting
ants (genera Atta Fabricius 1804, and Acromyrmex
Mayr 1865) to less specialized taxa using various
organic material (Fernández 2006; Mackay et al.
2004). Fungi are cultivated in specialized “rooms”
located in the inner part of the ant’s nest.

colonization by ants would have to have taken place
within a short period after the placement of the funerary
bundle in the cave given that freshly-cut leaves and other
plant parts are required for fungi cultivation. Additionally,
unintentional transportation of ants within the traditional
offerings following burial remains a possibility. Lastly, the
presence of ants may be linked with some prehistoric
intent carrying unknown symbolism.

Acromyrmex Mayr 1865
Acromyrmex versicolor (Pergande 1894)
(Figures 10, 11)
The genus Acromyrmex shares many morphological affinities with Atta. The Neotropical region includes 26 species. The workers are polymorphous, bearing numerous
spines (more than three pairs) on the mesosoma (Figure
10). Four species occur in Mexico: Acromyrmex echinatior
Forel 1899 (Chihuahua), A. lundii (Guérin-Méneville 1838)

(without precise locality data), A. octospinosus (Reich
1793) (including two subspecies) (Guerrero and Yucatan)
and A. versicolor (Lower California, Durango and Sonora)
(Rojas 2001; Alatorre-Bracamontes & Vásquez-Bolaños
2010). To a large extent, Acromyrmex species live mainly
in tropical forests but also occur in open environments
(pasture). An exception to the rule, A. versicolor lives in
arid and desert environments, building gigantic nests
including many mounds and entrances. In most cases, the
founding of ant colonies results from the alliance of several
ant queens; this singular behavior is called “pleometrosis”
(Fisher & Cover 2007).
In Mexico and in several other countries of Central
America, the leaf-cutting ants Acromyrmex and Atta,
from their activity of defoliation, represent true pests
of cultivated and sylvan plants. From the significant
plant biomass consumed, the ecological role of
Acromyrmex and Atta take the place of large herbivorous mammals.

Presence of A. versicolor in the funerary bundle
A. versicolor is recorded for the state of Coahuila for the
first time. The presence of this species in a funerary
context raises several questions. Firstly, the Attini are
strictly fungus-eaters and do not include any necrophagous or necrophilous taxa.3 Secondly, the tribe does not
include any troglobite or troglophilous forms. Therefore
their presence in such context is linked with other phenomena we elucidate below.
Acromyrmex ants build large and complex nests, and
hence the hypothesis that an entrance (or an exit) may
have once opened into the cave cannot to be ruled out.
Another important fact to consider is that the funerary

bundle, mainly composed of vegetable fiber cords, may
have attracted these insects. In this hypothesis, the

Lepidoptera
Tineidae?
The presence of this order is attested by the remains of
moth cocoons including various exogenous debris. These
mainly dermatophagous insects in the larval stage are
adapted to consume fabrics, feathers, integuments and
other organic debris. As in the trogid and dermestid beetles, these moths very likely colonized the funerary bundle
attracted by the epidermal remains and also by the fabrics
that they consumed and shredded. Distinct signs of activity attributed to this order have been reported by VergaraPineda et al. (2009).
Coleoptera
Dermestidae
The genus Dermestes L. 1758, which means etymologically “skin-feeder”, includes 92 species and sub-species
worldwide (Háva 2010). Among the representatives of this
genus, many species have a cosmopolitan distribution
(passive transportation via the international trade of
leather, hides, and various stored products). In regards to
their economic importance, their biology has been the
subject of a significant amount of literature (Lepesme
1938; Hinton 1945; Osuji 1975; Woodroffe & Coombs
1979; Delobel & Tran 1993).
Adults are medium-sized beetles (6–10 mm), oblongshaped, the dorsal surface reddish brown to black, and the
elytra sometimes with a pale transverse band and with or
without setose spots. In many species, the abdominal
sternites are covered with short, dense, chalk-white setae
and dark brown patches in the middle and on the sides.
In their natural habitat, both adults and larvae feed on
carrion at different stages of postmortem decay with a

marked predilection for desiccated cadavers (Byrd &
Castner 2009). From their relative frequency on human
corpses, Dermestes beetles have forensic significance in
helping to estimate the PMI of the cadaver (Smith 1986;
Schroeder et al. 2002; Pasquerault et al. 2008; Byrd &
Castner 2009; Kumara et al. 2009). In an archaeological
context, a few species figure among the classic hosts of
the Egyptian mummies (Hope 1834, 1836; Neolitzky
1911; Lesne 1930; Strong 1981; Adams 1990; Taylor
1995).


Annales de la Société entomologique de France (N.S.)
Dermestes L. 1758
Subgenus Dermestinus Zhantiev 1967
Dermestes (Dermestinus) carnivorus Fabricius 1775
(Figures 12, 13)
This species has been identified from several nearly complete adult specimens perfectly preserved (Figures 12, 13)
and from larval remains notably with the terminal abdominal segment bearing typical urogomphi (horn-like protrusions) on the upper surface. Oddly enough, VergaraPineda et al. (2009) identified the dermestid remains as
Dermestes (Dermestinus) maculatus Degeer 1774, from
both the adult head (one single adult) and funerary bundle
(three adults). Given that both species are close morphologically, we are inclined to have some reservations about
the simultaneous occurrence of both taxa in the same
context. D. maculatus can be reliably distinguished from
D. carnivorus by the presence of a small acuminate tooth
located at the inner apical angle of the elytron. Moreover,
only the fourth abdominal sternite bears a small median
stout bristle in males (this characteristic occurs on both the
third and fourth segments of the abdominal sternites in
males of D. carnivorus).

Although Dermestes carnivorus is probably native to
the Neotropical region, the species can currently be found
throughout most zoogeographical regions of the world
(Fauvel 1889; Lepesme 1946). In Mexico in its natural
habitat, this species is frequently reported to be found in
caves where it seems to be particularly attracted by bat
guano (Ryckman 1956; Constantine 1958). Experiments
conducted by Ryckman (1956) in the Ney cave provided
evidence that a D. carnivorus colony could skeletonize a
dead bat within six hours. Jirón & Cartín (1981) mentioned this species as frequently present on mammal
carcasses in Costa Rica. In an archaeological context,
D. carnivorous was recovered from funerary deposits
and offerings in the mortuary cave “La Chagüera”
(Morelos state, Mexico) (Muñiz Vélez 2001). The recurrence of this species in a karstic environment would
suggest that the colonization of the bundle by these
insects took place in situ, and was conducted by a resident population.
Staphylinidae
Aleocharinae Fleming 1821
Staphylinidae or “rove beetles” is an immensely ecologically diverse group including almost 47,000 species
arranged in nearly 3300 genera (Newton et al. 2000).
Navarrete-Heredia & Zaragoza-Caballero (2006) mentioned 1522 described species for Mexico including 725
endemics. The distribution by state shows marked differences as Coahuila appears to be relatively lower with only
19 known species for an area of 151,571 km2

283

(comparatively, Veracruz, half the size of Coahuila
(72,005 km2), includes 672 reported species).
Staphylinidae exhibit a wide diversity in their diet.
However, a few species can be regarded as strictly necrophagous (Fichter 1949). Most of the taxa frequenting

carrion are predacious in both the larval and adult stages
and prey on other arthropods (mainly eggs, maggots and
beetle larvae). According to Gennard (2007), rove beetles
usually colonize a body in the bloat stage of decomposition. Within the Aleocharinae sub-family (notably the
genus Aleochara Gravenhorst 1802), many species in
their larval form are solitary parasitoids of cyclorrhaphous
diptera (Peschke & Fuldner 1977). The first instar larvae
actively seek a puparium to parasitize, then perforate the
cuticle and finally devour its host. Depending on the
species, pupation occurs in situ or in the ground after the
larva leaves the puparium.
In all likelihood, the presence of one species of
Aleocharinae in this context is linked with the presence
of sarcosaprophagous flies within the funerary bundle.
Only an exhaustive and careful examination of the funerary bundle would allow us to reasonably conclude the
presence of perforated puparia due to the parasitoid behavior of these staphylinids.
Histeridae
This relatively small family of beetles includes nearly 4000
species worldwide (Mazur 1997). Most of the species are
considered rare and their specific identification can be very
difficult. Apart from aquatic environments, the Histeridae or
“clown beetles” have colonized most ecosystems and habitats and some groups exhibit noteworthy ecological adaptations (e.g. sub-cortical saproxylic, myrmecophilous,
sabulicolous, and cavernicolous species; Degallier & Gomy
1983; Gomy 2010). Their ethology is poorly known and few
larvae have been observed and described. It is generally
accepted that all species are predacious and that both adults
and larvae of the most widely distributed genera (e.g. Hister
L. 1758, and Saprinus Erichson 1834) are useful auxiliary
agents to fight against fly proliferation (notably Muscidae,
Calliphoridae, and Sarcophagidae). In light of their ecology,

histerid beetles are commonly associated with carrion and
sometimes used in forensic cases for establishing the PMI
(Introna et al. 1998; Byrd & Castner 2001). In a funerary
archaeological context, several histerid species have been
found in association with human remains from distinct geographical, cultural or chronological contexts (Faulkner 1986;
Huchet & Gallis 1996; Muñiz Vélez 2001).
The Histerid remains recovered from the funerary
bundle consist of three species belonging to the
Saprininae: Saprinus (s. str.) alienus J.L. LeConte 1851
(Figure 14), Xerosaprinus (s. str.) vitiosus (J.L. LeConte


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J.-B. Huchet et al.

Figures 6–19. Insect remains recovered from the funerary bundle of the Candelaria cave. 6, Synthesiomyia nudiseta Van der Wulp (Diptera
Muscidae), puparium. 7, idem, close-up of sinuous anal spiracular slits. 8, Fannia sp. (Diptera Fanniidae). 9, Diptera Sarcophagidae, puparium,
last segment in caudal view, the posterior spiracles lacking. 10, Acromyrmex versicolor (Pergande) (Hymenoptera Formicidae), part of the
mesosoma and gaster. 11, idem, pronotum and part of the mesonotum (upside-down view). The pair of the spines belongs to the pronotal area.
12, Dermestes (Dermestinus) carnivorus Fabricius (Coleoptera Dermestidae), dorsal view. 13, idem, lateral view. 14, Saprinus (s. str.) alienus
Le Conte (Coleoptera Histeridae), right elytron. 15, Xerosaprinus (s. str.) vitiosus (Le Conte) (Coleoptera Histeridae). 16, Xerosaprinus
coerulescens (Le Conte) (Coleoptera Histeridae). 17, Niptus n. sp., (Coleoptera Anobiidae Ptininae), lateral view. 18, Omorgus sp. (Coleoptera
Trogidae), abdominal sternite. 19, idem, part of pronotum. (Photos H.-P. Aberlenc, CIRAD-CBGP, Montpellier, France, 2007).


Annales de la Société entomologique de France (N.S.)
1851) (Figure 15) and Xerosaprinus (s. str.) coerulescens
(J.L. LeConte 1851) (Figure 16), detailed below.

Saprininae Blanchard 1845
Saprinus Erichson 1834
Subgenus Saprinus s. str.
Saprinus (Saprinus) alienus J.L. Le Conte 1851
(Figure 14)
Initially described from California, this species occurs in
New Mexico, Texas and Nevada and is also recorded in
Mexico. The only ecological record available regarding
this species is attributed to David S. Verity who collected
many specimens using rotten-meat traps (pitfall traps).
Xerosaprinus Wenzel in Arnett 1962
Subgenus Xerosaprinus s. str.
Xerosaprinus (Xerosaprinus) vitiosus (J. L. Le Conte
1851)
(Figure 15)
Described initially from Colorado, this species occurs in
California, Arizona and Mexico (Mazur 1997). The ecological data available concerning this species are: goat’s cadaver; on the beach, under rotten algae (G. Arriagada, personal
communication); on the beach, under a dead fish (Coll.
Y.G.); at light trap (Coll. Y.G.); in cow dung (Coll. Y.G.).
Xerosaprinus (Xerosaprinus) coerulescens (J. L. Le
Conte 1851)
(Figure 16)
Described initially from California, this species also occurs in
Nevada and Mexico. The ecology of this species remains
poorly known. Like Saprinus alienus, the species was collected by David S. Verity in rotten-meat traps (pitfall traps).
Unlike Niptus n. sp. or Omorgus sp. (see below) which
probably visited the bundle during the later stages of decomposition, the presence of histerid beetles is closely linked
with sarcosaprophagous fly activity which usually colonizes
carrion during both the active and advanced stages of decay.
Anobiidae

Ptininae Latreille 1802
Genre Niptus Boieldieu 1856
Niptus n. sp.
(Figure 17)
Niptus biology
Like most members of the spider beetles, species of Niptus
are scavengers, and frequently feed on dry dung. Taxa
often are associated with desert rodents and occasionally

285

are found in or around packrat (Neotoma spp.) nests,
kangaroo rat burrows (Dipodomys deserti Stephens 1887,
and Dipodomys spectabilis Merriam 1890), and also areas
frequented by mice (Peromyscus spp.) (Aalbu & Andrews
1992; Spilman 1976; T. K. Philips, personal observations).
In xeric habitats, rodents bring in a variety of plant
material into their nesting sites, such as seeds, twigs,
grass, leaves and pieces of cacti. While small fragments
of accumulated plant material might be used for feeding
by both adults and larvae, more important are the fecal
pellets from the rodents. Dung can build up to a large
degree in various latrine locations and are used as food for
both larvae and adults. Larvae are typically not free-living
and feed inside pellets or accumulated organic debris.

Distribution of North American species of Niptus
The eight native species in this genus (excluding the two
Pseudeurostus Heyden 1906 species sometimes placed
within Niptus) are restricted to the drier portions of the

southwest USA and Mexico. Niptus is a North American
clade that does not extend into Central or South America,
and the one European species, N. hololeucus (Faldermann
1836), is probably not closely related (Philips 2000).

A possible new species of Niptus from Candelaria
Mortuary Cave?
The discovery of a potentially undescribed species in
Candelaria Mortuary Cave leads one to hypothesize that
it might be a troglophile or even a troglobite and therefore
restricted to this cave environment. In addition to this new
taxon, other species of Niptus have been reported from
caves. For example, Ashworth (1973) reports fragments of
individuals of N. abstrusus Spilman 1968, in a 12,000year-old fossil Neotoma Say & Ord 1825 nest from a cave
in western Texas. This extant species is known from
several caves in southwestern Texas and north-central
Mexico (see Aalbu & Andrews 1992) and hence is likely
only a troglophile. In contrast, a true troglobite is Niptus
arcanus Aalbu & Andrews 1992, known only from El
Pakiva Cave, in Mitchell Caverns State Park, California,
in the Mojave Desert.
It is likely that in caves, populations of at least some
species of spider beetles can build up in numbers and
hence are more easily sampled in these habitats. This
might lead one to assume that the species is restricted to
the cave habitat; hence one needs to be cautious of assuming cave endemicity and the characterization of being a
troglobite. Cave associations for species of Niptus might
appear to be relatively recent as no species has typical
troglobite features such as lack of eyes. Reduced eye size
in both N. arcanus and N. abscondidus Spilman suggest

that these species are restricted to caves.


286

J.-B. Huchet et al.

Based on some similarities to N. ventriculus LeConte
1859, this potentially new species from Candelaria
Mortuary Cave may have evolved from an isolated population of the former. This is also true for several additional
species of Niptus that, based on morphological similarity,
likely speciated from isolated populations of N. ventriculus. One wonders how many species of Niptus restricted to
isolated caves in southwest USA and Mexico still remain
to be discovered and described.
Spider beetles have been reported to colonize corpses
during the ultimate stages of postmortem decay, notably
when the cadaver reaches an advanced stage of desiccation
(Smith 1986; Byrd & Castner 2009). The beetles usually
scavenge upon the remnant tissue and other organic remains
or debris left by the previous waves of necrophilous insects.
Different ptinid species have been recovered from funerary
archaeological contexts, notably Gibbium psylloides
(Czempinski 1778), found in large numbers in the wrappings
of the “Two Brothers” mummies (Nekht-Ankh and KhnumNakht) (David 1978; Curry 1979). This species was also
present among the Tutankhamum tomb material (Alfieri
1931) and also in the sarcophagus attributed to the French
count of Toulouse, Guillaume Taillefer, in association with
several other species of Ptinus L. (Huchet & Gallis 1996).
Recently, one of the authors (JBH) recovered the remains of
Ptinus sp. (cf. fur (L. 1758)) from the grave of the French King

Louis XI who died in 1483 (Huchet, forthcoming).
In the same way as D. carnivorus, previously mentioned, we are inclined to think that the colonization of the
mortuary bundle by this ptinid took place within the cave
by an autochthonous population.

Trogidae
Omorginae Nikolajev 2005
Omorgus Erichson 1847
The Trogidae (“hide beetles” or “carcass beetles”)
includes nearly 300 species worldwide (Scholtz 1986).
The taxon is notably diversified in the Neotropical,
Afrotropical and Australian regions. This family is primarily necrophilous and colonizes carcasses during the
advanced stages of postmortem decay, feeding notably
upon scleroproteins (skin, hair, fur, horns, hooves, etc.)
and other desiccated organic remains (e.g. tendons, and
ligaments) (Reed 1958; Palestrini et al. 1992).
Deloya (2000) reported 27 species of Trogidae for
Mexico, including 18 Omorgus species. Among
these taxa, only five species occur in Coahuila state:
O. fuliginosus Robinson 1941, O. punctatus (Germar
1824), O. scutellaris (Say 1823), O. suberosus
(Fabricius 1775) and O. umbonatus LeConte 1854.
Several species have been reported from natural caves
where they exploit bat guano (Scholtz 1986). Muñiz
Vélez (2001) mentioned three Omorgus species

inhabiting the funerary caves of Morelos state in
Mexico (O. suberosus, O. rubricans (Robinson 1946),
and O. fuliginosus). O. suberosus is undoubtedly the
most common Mexican species, occurring in 20 of the

31 states (Deloya 2000). Trogid beetles, like species
belonging to the genus Dermestes (Coleoptera:
Dermestidae), only feed on desiccated carcasses that
are subaerially exposed. Omorgus females oviposit in
the sediment subjacent to the primary source of food at
a depth ranging between 15 and 25 cm depending on
the species (Baker 1968). In an archaeological context,
their association with buried human remains indicates
that the corpse was temporarily exposed over a relatively long period (Ubelaker & Willey 1978; Huchet &
Greenberg 2010). The presence of Trogid remains in the
funerary bundle indicates that the corpse was directly
accessible to these insects and consequently remained
unburied. In all likelihood, the colonization took place
during the dry decay stage, the insects attracted by
desiccated skin and ligaments.
The identification of the remains at the generic level
was made from an abdominal sternite (Figure 18) and a
fragment of pronotum (Figure 19).

Discussion
From the composition of the thanatophilous assemblage
recovered within the funerary bundle, some hypotheses
about the funerary treatment of the young child can be
proposed.
Surprisingly, no calliphorid remains were recovered
from the sampling, though this might be attributable to
the fact that the sample studied here represents a tiny part
of the entomofauna from the bundle. Additionally, no
calliphorid remains were reported in the Vergara-Pineda
et al. (2009) study. Due to their ability to locate and

colonize a cadaver soon after death, calliphorids (blowflies) are very often involved in forensic investigations in
the estimation of the PMI (Smith 1986; Greenberg 1991;
Greenberg & Kunich 2002; Szpila 2010 inter alia). In the
present case, their absence may be explained in two ways:
either a short exposure of the body prior to burial and/or a
rapid wrapping of the child in the funerary bundle, or due
to the seasonality of death.4
With regard to the biological data concerning
Sarcophagidae and Fanniidae, we are inclined to think
that these flies probably arrived at a later stage, after the
corpse exhaled the first odors of decay. Although it is
generally agreed that flies rarely oviposit in the dark
(Nuorteva 1977; Greenberg 1990), we are inclined to
think that, in the present case, the location of the bodies
in the cave, at a few meters from the entrance, did not
prevent the colonization of the funerary bundle. In this
respect, the garment, heavily impregnated with bodily


Annales de la Société entomologique de France (N.S.)
fluids and other liquefying tissues, probably played a very
attractive role.
Concerning the colonization by the different beetle
taxa, some hypotheses regarding the chronology of their
arrival can be suggested. In all likelihood, histerid and
staphylinid beetles invaded the bundle more or less simultaneously with the necrophagous flies (the presence of
these later being inherent with the persistence of damp
organic matter). Both families of beetles actively prey
upon maggots and their presence probably dates from
the first few weeks following the young individual’s

death. On the other hand, the colonization of the bundle
by the Dermestidae, Ptininae and Trogidae correspond to a
later stage, after the human remains reached a significant
degree of desiccation. However, among these latter taxa,
dermestid beetles are known to sometimes invade within a
short period after death (Early & Goff 1987;
VanLaerhoven & Anderson 1999; Oliva 2001).
In the present case, the most relevant data are
based upon the biology of the muscid fly Synthesiomyia
nudiseta. As previously mentioned, at the postfeeding
stage the larvae secrete a sticky substance which, under
the effect of the last movement prior to molting, covers the
puparium with soil particles or any other exogenous material nearby. Fragmented remains of S. nudiseta cocoons
recovered within the funerary bundle proved to be of great
value and raised interesting questions. A careful examination reveals that they include many remains of dermestid
larval skins (exuviae) and presumed dermestid fecal pellets, indicating that S. nudiseta probably invaded the
cadaver at a later stage than usual, probably after dermestid infestation. This would suggest that the bundle had
been, intentionally or not, temporarily rehydrated, probably in the few weeks following death. From this hypothesis, different scenarios might be envisaged. A first
hypothesis would be that the bundle had been placed in
the cave after a certain period of time, perhaps the time
taken to transport the bundle to the burial site from the
place of death. The corpse could have been exposed in the
open air before its final placement in the mortuary cave.
However, in this case, we are inclined to think that the
pioneer calliphorid flies would have probably colonized
the corpse and some of their remains would be present in
the samplings. A second hypothesis would be that the
hunter–gatherer semi nomads had temporarily or even
periodically removed the deceased ancestors from the
cave in order to satisfy some funerary practices which

would have left no trace in the archaeological record (the
cave entrance having been able to play the role of a
“connecting zone” between the World of the Living and
the World of the Dead). However, it is reasonable to
expect that the cave entrance, located on a steep slope,
would have probably restricted such practices. We could
not totally preclude that the bundle had been rehydrated
by isolated or repeated episodes of water seepage. Finally,

287

the existence of different funerary rituals in the core of the
cave (e.g. ritual aspersion of the deceased?) could be at the
origin of the late colonization by the muscid fly S.
nudiseta.
Conclusion
This archaeoentomological study of a sample recovered
from a Mexican funerary bundle shows a standard combination of the main invertebrate thanatofauna associated
with decaying cadavers. The high specific diversity definitely indicates that several colonization episodes occurred
and thus a relatively easy access to the cadaver was likely
throughout the process of bodily decay. In the present
case, the lack of the pioneer flies (Calliphoridae) indicate
that the corpse was apparently shielded early on from
insect activity (rapid wrapping in the bundle without
body exposure prior to burial, or the death may have
occurred during a period when fly activity is low).
Finally, the late or postponed colonization by the muscid
fly S. nudiseta, evidenced by the nature of the exogenous
elements used to build the cocoons, may indicate that the
bundle may have been occasionally rehydrated, suggesting

secondary handling of the deceased (e.g. occasional
removal of the bundle(s) from the funerary cave?).
Further studies of the invertebrate thanatofauna recovered
from Mexican funerary bundles may result in further
insights into the funerary practices of these prehistoric
populations.
Acknowledgments
We would like to thank our colleagues G. Arriagada (Chili) and
D. S. Verity (California, USA) who verified the identification of
the three Histeridae species recovered from the funerary bundle.
We are also indebted to our friend and colleague H.-P. Aberlenc
(CIRAD-CBGP, Montpellier, France) for the noteworthy photographs of the insect remains illustrating this paper. Finally, we
are especially indebted to K. Irby who kindly assisted in improving the English of this paper.

Notes
1.
2.

3.
4.

Dating carried out on teeth collected from several crania
from Candelaria, using accelerator mass spectrometry at
the Oxford University.
Taxonomic identifications of the insect remains have been
respectively performed by Y. Gomy (Coleoptera:
Histeridae), T.K. Phillips (Coleoptera: Anobiidae:
Ptininae), C.E. Alatorre-Bracamontes and M. VásquezBolaños (Hymenoptera: Formicidae), J.L. Navarette
Heredia (Mexico) (Coleoptera: Staphylinidae) and J.-B.
Huchet (Diptera: Muscidae, Fannidae, Sarcophagidae;

Coleoptera: Dermestidae and Trogidae).
Ants of some basal Attini genera (e.g. Cyphomyrmex)
occasionally use small arthropod cadavers as substratum
for fungi cultivation.
Recent (unpublished) experiments conducted by one of the
authors (JBH) in Peru evidenced that, depending on the


288

J.-B. Huchet et al.
season of the year, different muscid or sarcophagid flies
can replace the pioneer flies (calliphorid) during the first
stage of decay.

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