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

aMuséum national d’Histoire naturelle, Département Systématique et Evolution (entomologie), UMR 7205 du CNRS – Origine, Structureet 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 etenvironnements. CP 56, 55 rue Buffon F-75005 Paris, France; cUMR 5199 du CNRS, PACEA – Anthropologie des populations passéeset 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 CollegeHeights Blvd. Bowling Green, KY 42101-3576, USA; gEntomología, Centro de Estudios en Zoología, Departamento de Botánica yZoología, Centro Universitario de Ciencias Biológicas y Agropecuarias, Universidad de Guadalajara, Km. 15.5 Carr. Nogales, LasAgujas, 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

(Accepté le 23 mai 2013)

Summary. The archaeoentomological study of insect remains recovered from a pre-Columbian funerary bundle (10th–11thcenturies AD) is presented and illustrated. Among this material, 12 species belonging to 10 families and four distinct ordershave been identified. From the biological data of the different taxa, some hypotheses about the funerary practices of thehunter–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 ordresdistincts d’insectes ont pu être identifiées. A partir des données biologiques des différents taxons en présence, deshypothèses 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 karsticenvironment, allows an exceptional preservation oforganic matter and other perishable materials. For twomillennia prior to the Spanish conquest (1519–1521),numerous natural caves in northern Mexico were used asburial sites by hunter–gatherer populations who buriedtheir dead in these locations over several generations. Inthe dry environment of these natural sanctuaries, thebodies placed in mortuary bundles made of plaited vegetalmaterials have been preserved over the centuries, as wellas the fragile remains of necrophagous insects linked toprocesses of thanatomorphosis.

The Candelaria funerary cave (Coahuila State,Mexico) excavated in the early 1950s has provided manyhuman remains, often remarkably preserved. These popu-lations have been the subject of numerous archaeologicaland bioarchaeological studies (Martínez del Río 1953a,1953b; Romano 1956; Taylor 1968; Mansilla & Pijoan

2000). While the presence of insects had been previouslynoted, the first archeoentomological analysis based onsome of these remains was performed very recently byVergara-Pineda et al. (2009). These last authors providedthe first inventory of entomofauna associated with humanremains at Candelaria, basing their analysis both on asample recovered from an adult cranium and from thefunerary bundle of a young child. A second sample setrecovered from the same bundle is presented and illu-strated here.

Beyond a strictly taxonomic interest, the study ofinvertebrate necrofauna in an archaeological funerarycontext also sheds light on the immediate environmentof the site at the moment of its use. From a “paleo-forensic” perspective, it may provide a new insight intothe reconstruction and understanding of the funerarypractices of semi nomadic groups from northernMexico.

*Corresponding author. Email: huchet@mnhn.fr

Annales de la Société entomologique de France (N.S.), 2013Vol. 49, No. 3, 277–290, http://dx.doi.org/10.1080/00379271.2013.845474

© 2013 Société entomologique de France

Geographical setting

The Candelaria funerary cave is located near San Pedro delas Colonias, in the semidesert zone of the State ofCoahuila, northern Mexico [approximately 25°01′ N,102°46′ W] (Figure 1). It is a natural cavity that formedin Cretaceous limestone of the Sierra de la Candelaria, asmall mountain formation that borders the Las Deliciasendorheic basin (Maldonado-Koerdell 1956). The regionis characterized by an arid climate (less than 200 mm ofannual precipitation) and Chihuahuan desert vegetation(primarily cacti, yucca, and thorn shrubs).

The cavity occurs at 1000 m asl. It is accessible througha shaft approximately 9 m high and 1 m wide. The funerarydeposits were located in two adjoining rooms where thefloors show a regular inclination (Figure 2).

Archaeological context

The Candelaria funerary cave is an exceptional site inthe archaeology of northern Mexico for the spectacularstate of conservation of its organic remains. The siteoffers a particularly rich image of funerary practicesand the material culture of the hunter–gatherers of this

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

Figure 1. Geographic location of the funerary cave of Candelaria in Coahuila State, Mexico.

Figure 2. Candelaria Cave, seen in cross section, with thelocation of the funerary bundles (adapted from Maldonado-Koerdell 1956).

278 J.-B. Huchet et al.

Antropología e Historia (Aveleyra Arroyo de Anda,Maldonado-Koerdell and Martínez del Río), after evi-dence of looting activities had been reported. Thedeposits have been partially disturbed prior to the begin-ning of the excavations. While the possibility of post-depositional contamination cannot be totally excluded,the composition and the biology of the faunal composi-tion presented here leads us to believe that all of thesestudied insect remains are linked to processes of post-mortem decay, and are therefore contemporaneous withthe deposit.

Despite the perturbations, we can discern precise infor-mation regarding the funerary practices. The cavity func-tioned as a collective burial place during the prehispanicperiod, with at least 116 individuals (83 adults and 33subadults) buried there. The skeletal collection comprisesmainly a deposit of mixed bones and two intact infantmortuary bundles. Commingling of bones was causedapparently by natural rock falls inside the cave and thedestruction of originally individual mortuary bundles bylooters. These recurring burials took place over severalgenerations. The bodies were brought into the cave asmortuary bundles (Romano 1956) and placed on thefloor in the cave, without making a pit or covering themwith earth (González Arratia 2007). The corpses werewrapped in a fetal position with pieces of cloth madefrom yucca fibers maintained by a network of fine cords(Weitlaner-Johnson 1977) (Figures 3, 4), and the bundleswere tied with a cord to keep in place the bent legs andarms. Each bundle was placed on a scaffold comprised of

varied perishable materials: mats, trays, frames of wood,and various objects (bows, digging sticks, etc.) and indi-vidually separated by vegetative matter (cactus pads,yucca leaves, etc.).

Many other objects deposited around the deceasedwere also collected (Martínez del Río 1953a, 1953b;Maldonado-Koerdell 1956; Aveleyra Arroyo de Anda1964; González Arratia 1999): wooden tools and huntingweapons, arrows, hafted chert knives, basketry, sandals,cordage, stone ornaments, shell, bone, etc. The character-istics of some of these objects and the first radiocarbondates obtained (Aveleyra Arroyo de Anda 1964; GonzálezArratia 1999, p. 48–49) suggested that the cave’s occupa-tion occurred around the twelfth and thirteenth centuriesAD. New C14 dating obtained by one of the authors (J.Mansilla, in: Pineda et al. 20091) indicates that the funer-ary deposits are considerably more ancient (AD 940 ± 24and 1020 ± 28).

Given the excellent conservation of the human skeletalremains, numerous studies have been conducted (e.g. cra-niometry: Romano 1956; pathology: Mansilla & Pijoan2000, 2005; Pineda et al. 2009). However, despite theexceptional preservation of the organic material, the softtissues were only very rarely preserved.

Only three bundles were recovered intact: those of oneadult and two children. A radiological study carried out onone of the two juvenile bundles by Mansilla & Pijoan(2000) (Figure 5) determined that the deceased individualwas three years old. The second child died before he orshe 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 secondchild. 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 ancientMexico

Although the precise meaning behind the funerary bundlescreated by the hunter–gatherers at Candelaria Caveremains unknown, it is important to point out that thispractice was not only restricted to this site and culture.The custom of wrapping a corpse within a cloth or matbound with cords occurred widely throughout theAmericas, from the Southwestern USA to the Andes. InMexico, this practice has been observed archaeologicallyin the northern semi-arid regions where the climate favorsthe preservation of the organic remains. While the first aimof this system of wrapping was probably to facilitate thetransport of the dead to the burial place, the bundle is notrestricted to this single function and, in different cultures,it also carried a symbolic meaning. Among the Aztec andother Late Postclassic societies (AD 1200–1521), funerarybundles were considered to represent the chrysalis of abutterfly (Taube 2002, p. 308–309). Representations frompictographic manuscripts (codices) show that these bun-dles were often decorated with butterfly imagery or withspecific anatomic attributes of the chrysalis. Those des-tined for flames were themselves symbolized by a butter-fly, the symbol of both fire and the souls of the deaddoomed to the celestial beyond (Beutelspacher 1988,p. 16–20). Regardless, there is no evidence to date thatthe inhabitants of Candelaria Cave shared this particularsymbolism, which corresponds to very distinct groupswho did not practice cremation burials. Instead, it appearsthat this form of funerary preparation served a more prag-matic purpose, such as, in the present case, facilitating thetransport of the dead to a cave that was difficult to access.

The insect remains

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

younger child (Figure 3). The collected residues have beenprocessed to dry sieving (mesh size: 300 µm), examinedunder a stereomicroscope and mounted according to stan-dard entomological procedures (glued on card-points andlabeled).

The faunal composition recovered from this context ishardly surprising since, to a large extent, the different taxaare those found to most frequently colonize carrion. Thesampling includes 12 species belonging to 10 families andfour distinct orders (Diptera, Coleoptera, Hymenopteraand Lepidoptera).

Diptera

The remains or “sclerites” attributable to this order (nearlycomplete or fragmented puparia) correspond to threesarco-saprophagous families of cyclorraphous flies:Muscidae, Fanniidae and Sarcophagidae. These threefamilies traditionally figure among the most relevant taxafor estimating the postmortem interval (PMI) in forensicentomology.

Muscidae

Synthesiomyia nudiseta Van der Wulp 1883(Figures 6, 7)

This species usually colonizes cadavers a few days afterthe first settlement of necrophagous flies (Skidmore 1984).According to Buxton & Hopkins (1927), S. nudisetainvades carrion at the active decay stage (the “secondwave” according to the terminology of Easton & Smith[1970]). When pupation occurs, the mature larvae exude afrothy and sticky substance which solidifies to form aprotective cocoon upon which sand particles and otherexogenous 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 Family Taxa Number of insect remains MNI*

Diptera Muscidae Synthesiomyia nudiseta van der Wulp > 10 4Diptera Fanniidae Fannia sp. 10 5Diptera Sarcophagidae Genus ? sp. ? > 3 3Coleoptera Dermestidae Dermestes (Dermestinus) carnivorus Fabricius > 9 5Coleoptera Histeridae Saprinus (s. str.) alienus J.L. LeConte 1 1Coleoptera Histeridae Xerosaprinus (s. str.) coerulescens (J.L. LeConte) 1 1Coleoptera Histeridae Xerosaprinus (s. str.) vitiosus (J.L. LeConte) 1 1Coleoptera Staphylinidae Genus ? sp. ? 1 1Coleoptera Anobiidae Niptus n. sp. > 10 8Coleoptera Trogidae Omorgus sp. 2 1Hymenoptera Formicidae Acromyrmex versicolor (Pergande) > 10 3Lepidoptera Tineidae ? Genus ? sp. ? 1 1

Total > 59 34

* MNI = minimal number of individuals.

280 J.-B. Huchet et al.

pupate, S. nudiseta usually completes the entire biologicalcycle in situ (in close contact with the cadaver). In theNeotropical region, this species among the flies is mostfrequently associated with human remains in archaeologi-cal contexts (Huchet & Greenberg 2010). In this respect,the unidentified muscid species associated with sacrificedhuman bodies from Pacatnamu (Peru) mentioned byFaulkner (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 carrionearly on and then replace the pioneer blowflies.

Due to its great dispersal ability, S. nudiseta is nowwidely distributed throughout the warmer parts of theglobe, and sometimes very distant from the supposedarea of origin (South America). In the NationalMuseum of Natural History (Paris), the species is repre-sented by specimens from various origins, including NewCaledonia, Thailand and Japan. The species has beenrecently collected in Europe from both Spain (Ebejer &Gatt 1999) and Malta (Carles-Tolrá 2002). In the field offorensic entomology, S. nudiseta is frequently used in theestimation of the PMI (Jirón et al. 1983; Lord et al.1992; Oliveira-Costa et al. 2001; Calderón-Arguedaset al. 2005; Kumara et al. 2009). A recent discovery ina pre-Columbian grave in Peru (Mochica civilization)(Huchet & Greenberg 2010) provided evidence of com-plex funeral practices, notably a long exposure of thecorpse prior to burial, as evidenced by the presence ofthis species.

The characteristic shape of the anal spiracular slits(Eldridge & James 1957; Siddons & Roy 1942; Skidmore1984) combined with the presence of distinct remains ofsandy cocoons recovered from the sample leaves no doubtabout the specific identification of this species.

Fanniidae

Fannia sp.(Figure 8)

The Fanniidae species usually develop in relativelyadvanced cadavers (Nuorteva et al. 1974; Smith 1986;Lee & Marzuki 1993). The immature stages of the speciesbelonging to the genus Fannia Robineau-Desvoidy 1830are morphologically characteristic. The larva and pupariaare flattened dorsoventrally and bear many well-developedplumose lateral and dorsal processes. The Neotropicalfauna includes 66 species (Carvalho et al. 2003).

Remains of Fannia canicularis (L. 1761) wererecently recovered from an Egyptian mummy of thePtolemaic Period (Gerisch 2001). As with the speciespreviously quoted, the presence of a representative of thegenus Fannia is undoubtedly linked with the postmortemdecay of the young child.

Sarcophagidae

Genus ? sp. ?(Figure 9)

The Sarcophagidae or “flesh flies” includes nearly 2600species worldwide (Pape et al. 2009). Although probablymore diversified than in any other region of the world,little is known regarding the Neotropical fauna (Pape1989). Unlike the Calliphoridae, the Sarcophagidaefemales are larviparous and directly deposit first instarlarvae on carrion. The females usually arrive at corpsesslightly later than the pioneer blowflies, attracted by thefirst odors of decay.

The identification at the family level is based on thepresence of a puparium fragment (Figure 9). Contrary toother sarcosaprophagous diptera families, the posteriorspiracles are hidden in a crateriform-shaped cavity locatedin the upper part of the last abdominal segment. Noexhaustive work on puparia is available and the onlyexisting reference concerning the Sarcophagidae (Greene1925) is incomplete and now obsolete. Ten additionalpuparia attributed to this family that were recoveredfrom the funerary bundle are mentioned by Vergara-Pineda et al. (2009).

Hymenoptera

Formicidae

The presence of Formicidae on carrion can be linked totwo distinct ecological habits: predation upon eggs,larvae or pupae of thanatophagous insects, or necro-phagy (ants feeding on exudates or decomposing tis-sues) (Moretti de Carvalho & Ribeiro 2006; Clark &Blom 1991). When ants are found in association withhuman remains, they are sometimes used as part offorensic investigations (Goff & Win 1997; Martínezet al. 1997). The presence of a fungus-growing antspecies within the bundle raises many questions.Different hypotheses on how and why such colonizationoccurs are proposed below.

Myrmicinae Lepeletier 1835

Attini Smith 1858

The Attini is a New World endemic tribe primarilyNeotropical in distribution which includes 95% of theknown species, with the other 5% restricted to theNearctic region (Fernández 2003, 2006; Mayhé-Nunes& Jaffe 1998). Attini comprises a monophyletic groupof approximately 230 described species (Schultz &Brady 2008) arranged in 14 distinct genera (Boltonet al. 2006). These fungus-growing ants use the organicdetritus they collect as a substrate for their fungal culti-vars, the main source of their diet (basidiomycete fun-gus of the genus Leucocoprinus (Pat.), notably

Annales de la Société entomologique de France (N.S.) 281

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

Acromyrmex Mayr 1865

Acromyrmex versicolor (Pergande 1894)(Figures 10, 11)

The genus Acromyrmex shares many morphological affi-nities with Atta. The Neotropical region includes 26 spe-cies. The workers are polymorphous, bearing numerousspines (more than three pairs) on the mesosoma (Figure10). Four species occur in Mexico: Acromyrmex echinatiorForel 1899 (Chihuahua), A. lundii (Guérin-Méneville 1838)(without precise locality data), A. octospinosus (Reich1793) (including two subspecies) (Guerrero and Yucatan)and A. versicolor (Lower California, Durango and Sonora)(Rojas 2001; Alatorre-Bracamontes & Vásquez-Bolaños2010). To a large extent, Acromyrmex species live mainlyin tropical forests but also occur in open environments(pasture). An exception to the rule, A. versicolor lives inarid and desert environments, building gigantic nestsincluding many mounds and entrances. In most cases, thefounding of ant colonies results from the alliance of severalant queens; this singular behavior is called “pleometrosis”(Fisher & Cover 2007).

In Mexico and in several other countries of CentralAmerica, the leaf-cutting ants Acromyrmex and Atta,from their activity of defoliation, represent true pestsof cultivated and sylvan plants. From the significantplant biomass consumed, the ecological role ofAcromyrmex and Atta take the place of large herbivor-ous mammals.

Presence of A. versicolor in the funerary bundle

A. versicolor is recorded for the state of Coahuila for thefirst time. The presence of this species in a funerarycontext raises several questions. Firstly, the Attini arestrictly fungus-eaters and do not include any necropha-gous or necrophilous taxa.3 Secondly, the tribe does notinclude any troglobite or troglophilous forms. Thereforetheir presence in such context is linked with other phe-nomena we elucidate below.

Acromyrmex ants build large and complex nests, andhence the hypothesis that an entrance (or an exit) mayhave once opened into the cave cannot to be ruled out.Another important fact to consider is that the funerarybundle, mainly composed of vegetable fiber cords, mayhave attracted these insects. In this hypothesis, the

colonization by ants would have to have taken placewithin a short period after the placement of the funerarybundle in the cave given that freshly-cut leaves and otherplant parts are required for fungi cultivation. Additionally,unintentional transportation of ants within the traditionalofferings following burial remains a possibility. Lastly, thepresence of ants may be linked with some prehistoricintent carrying unknown symbolism.

Lepidoptera

Tineidae?

The presence of this order is attested by the remains ofmoth cocoons including various exogenous debris. Thesemainly dermatophagous insects in the larval stage areadapted to consume fabrics, feathers, integuments andother organic debris. As in the trogid and dermestid bee-tles, these moths very likely colonized the funerary bundleattracted by the epidermal remains and also by the fabricsthat they consumed and shredded. Distinct signs of activ-ity attributed to this order have been reported by Vergara-Pineda et al. (2009).

Coleoptera

Dermestidae

The genus Dermestes L. 1758, which means etymologi-cally “skin-feeder”, includes 92 species and sub-speciesworldwide (Háva 2010). Among the representatives of thisgenus, many species have a cosmopolitan distribution(passive transportation via the international trade ofleather, hides, and various stored products). In regards totheir economic importance, their biology has been thesubject of a significant amount of literature (Lepesme1938; Hinton 1945; Osuji 1975; Woodroffe & Coombs1979; Delobel & Tran 1993).

Adults are medium-sized beetles (6–10 mm), oblong-shaped, the dorsal surface reddish brown to black, and theelytra sometimes with a pale transverse band and with orwithout setose spots. In many species, the abdominalsternites are covered with short, dense, chalk-white setaeand dark brown patches in the middle and on the sides.

In their natural habitat, both adults and larvae feed oncarrion at different stages of postmortem decay with amarked predilection for desiccated cadavers (Byrd &Castner 2009). From their relative frequency on humancorpses, Dermestes beetles have forensic significance inhelping 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 archaeologicalcontext, a few species figure among the classic hosts ofthe Egyptian mummies (Hope 1834, 1836; Neolitzky1911; Lesne 1930; Strong 1981; Adams 1990; Taylor1995).

282 J.-B. Huchet et al.

Dermestes L. 1758

Subgenus Dermestinus Zhantiev 1967

Dermestes (Dermestinus) carnivorus Fabricius 1775(Figures 12, 13)

This species has been identified from several nearly com-plete adult specimens perfectly preserved (Figures 12, 13)and from larval remains notably with the terminal abdom-inal segment bearing typical urogomphi (horn-like protru-sions) on the upper surface. Oddly enough, Vergara-Pineda et al. (2009) identified the dermestid remains asDermestes (Dermestinus) maculatus Degeer 1774, fromboth the adult head (one single adult) and funerary bundle(three adults). Given that both species are close morpho-logically, we are inclined to have some reservations aboutthe simultaneous occurrence of both taxa in the samecontext. D. maculatus can be reliably distinguished fromD. carnivorus by the presence of a small acuminate toothlocated at the inner apical angle of the elytron. Moreover,only the fourth abdominal sternite bears a small medianstout bristle in males (this characteristic occurs on both thethird and fourth segments of the abdominal sternites inmales of D. carnivorus).

Although Dermestes carnivorus is probably native tothe Neotropical region, the species can currently be foundthroughout most zoogeographical regions of the world(Fauvel 1889; Lepesme 1946). In Mexico in its naturalhabitat, this species is frequently reported to be found incaves where it seems to be particularly attracted by batguano (Ryckman 1956; Constantine 1958). Experimentsconducted by Ryckman (1956) in the Ney cave providedevidence that a D. carnivorus colony could skeletonize adead bat within six hours. Jirón & Cartín (1981) men-tioned this species as frequently present on mammalcarcasses in Costa Rica. In an archaeological context,D. carnivorous was recovered from funerary depositsand offerings in the mortuary cave “La Chagüera”(Morelos state, Mexico) (Muñiz Vélez 2001). The recur-rence of this species in a karstic environment wouldsuggest that the colonization of the bundle by theseinsects took place in situ, and was conducted by a resi-dent population.

Staphylinidae

Aleocharinae Fleming 1821

Staphylinidae or “rove beetles” is an immensely ecologi-cally diverse group including almost 47,000 speciesarranged in nearly 3300 genera (Newton et al. 2000).Navarrete-Heredia & Zaragoza-Caballero (2006) men-tioned 1522 described species for Mexico including 725endemics. The distribution by state shows marked differ-ences as Coahuila appears to be relatively lower with only19 known species for an area of 151,571 km2

(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 necro-phagous (Fichter 1949). Most of the taxa frequentingcarrion are predacious in both the larval and adult stagesand prey on other arthropods (mainly eggs, maggots andbeetle larvae). According to Gennard (2007), rove beetlesusually colonize a body in the bloat stage of decomposi-tion. Within the Aleocharinae sub-family (notably thegenus Aleochara Gravenhorst 1802), many species intheir larval form are solitary parasitoids of cyclorrhaphousdiptera (Peschke & Fuldner 1977). The first instar larvaeactively seek a puparium to parasitize, then perforate thecuticle and finally devour its host. Depending on thespecies, pupation occurs in situ or in the ground after thelarva leaves the puparium.

In all likelihood, the presence of one species ofAleocharinae in this context is linked with the presenceof sarcosaprophagous flies within the funerary bundle.Only an exhaustive and careful examination of the funer-ary bundle would allow us to reasonably conclude thepresence of perforated puparia due to the parasitoid beha-vior of these staphylinids.

Histeridae

This relatively small family of beetles includes nearly 4000species worldwide (Mazur 1997). Most of the species areconsidered rare and their specific identification can be verydifficult. Apart from aquatic environments, the Histeridae or“clown beetles” have colonized most ecosystems and habi-tats and some groups exhibit noteworthy ecological adapta-tions (e.g. sub-cortical saproxylic, myrmecophilous,sabulicolous, and cavernicolous species; Degallier & Gomy1983; Gomy 2010). Their ethology is poorly known and fewlarvae have been observed and described. It is generallyaccepted that all species are predacious and that both adultsand larvae of the most widely distributed genera (e.g. HisterL. 1758, and Saprinus Erichson 1834) are useful auxiliaryagents to fight against fly proliferation (notably Muscidae,Calliphoridae, and Sarcophagidae). In light of their ecology,histerid beetles are commonly associated with carrion andsometimes used in forensic cases for establishing the PMI(Introna et al. 1998; Byrd & Castner 2001). In a funeraryarchaeological context, several histerid species have beenfound in association with human remains from distinct geo-graphical, cultural or chronological contexts (Faulkner 1986;Huchet & Gallis 1996; Muñiz Vélez 2001).

The Histerid remains recovered from the funerarybundle consist of three species belonging to theSaprininae: Saprinus (s. str.) alienus J.L. LeConte 1851(Figure 14), Xerosaprinus (s. str.) vitiosus (J.L. LeConte

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Figures 6–19. Insect remains recovered from the funerary bundle of the Candelaria cave. 6, Synthesiomyia nudiseta Van derWulp (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).

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 inNew Mexico, Texas and Nevada and is also recorded inMexico. The only ecological record available regardingthis species is attributed to David S. Verity who collectedmany specimens using rotten-meat traps (pitfall traps).

Xerosaprinus Wenzel in Arnett 1962

Subgenus Xerosaprinus s. str.

Xerosaprinus (Xerosaprinus) vitiosus (J. L. Le Conte1851)

(Figure 15)

Described initially from Colorado, this species occurs inCalifornia, Arizona and Mexico (Mazur 1997). The ecolo-gical data available concerning this species are: goat’s cada-ver; on the beach, under rotten algae (G. Arriagada, personalcommunication); 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. LeConte 1851)(Figure 16)

Described initially fromCalifornia, this species also occurs inNevada and Mexico. The ecology of this species remainspoorly known. Like Saprinus alienus, the species was col-lected by David S. Verity in rotten-meat traps (pitfall traps).

Unlike Niptus n. sp. or Omorgus sp. (see below) whichprobably visited the bundle during the later stages of decom-position, the presence of histerid beetles is closely linkedwith sarcosaprophagous fly activity which usually colonizescarrion 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 Niptusare scavengers, and frequently feed on dry dung. Taxaoften are associated with desert rodents and occasionally

are found in or around packrat (Neotoma spp.) nests,kangaroo rat burrows (Dipodomys deserti Stephens 1887,and Dipodomys spectabilis Merriam 1890), and also areasfrequented by mice (Peromyscus spp.) (Aalbu & Andrews1992; Spilman 1976; T. K. Philips, personal observations).

In xeric habitats, rodents bring in a variety of plantmaterial into their nesting sites, such as seeds, twigs,grass, leaves and pieces of cacti. While small fragmentsof accumulated plant material might be used for feedingby both adults and larvae, more important are the fecalpellets from the rodents. Dung can build up to a largedegree in various latrine locations and are used as food forboth larvae and adults. Larvae are typically not free-livingand feed inside pellets or accumulated organic debris.

Distribution of North American species of Niptus

The eight native species in this genus (excluding the twoPseudeurostus Heyden 1906 species sometimes placedwithin Niptus) are restricted to the drier portions of thesouthwest USA and Mexico. Niptus is a North Americanclade that does not extend into Central or South America,and the one European species, N. hololeucus (Faldermann1836), is probably not closely related (Philips 2000).

A possible new species of Niptus from CandelariaMortuary Cave?

The discovery of a potentially undescribed species inCandelaria Mortuary Cave leads one to hypothesize thatit might be a troglophile or even a troglobite and thereforerestricted to this cave environment. In addition to this newtaxon, other species of Niptus have been reported fromcaves. For example, Ashworth (1973) reports fragments ofindividuals of N. abstrusus Spilman 1968, in a 12,000-year-old fossil Neotoma Say & Ord 1825 nest from a cavein western Texas. This extant species is known fromseveral caves in southwestern Texas and north-centralMexico (see Aalbu & Andrews 1992) and hence is likelyonly a troglophile. In contrast, a true troglobite is Niptusarcanus Aalbu & Andrews 1992, known only from ElPakiva Cave, in Mitchell Caverns State Park, California,in the Mojave Desert.

It is likely that in caves, populations of at least somespecies of spider beetles can build up in numbers andhence are more easily sampled in these habitats. Thismight lead one to assume that the species is restricted tothe cave habitat; hence one needs to be cautious of assum-ing cave endemicity and the characterization of being atroglobite. Cave associations for species of Niptus mightappear to be relatively recent as no species has typicaltroglobite features such as lack of eyes. Reduced eye sizein both N. arcanus and N. abscondidus Spilman suggestthat these species are restricted to caves.

Annales de la Société entomologique de France (N.S.) 285

Based on some similarities to N. ventriculus LeConte1859, this potentially new species from CandelariaMortuary Cave may have evolved from an isolated popu-lation of the former. This is also true for several additionalspecies of Niptus that, based on morphological similarity,likely speciated from isolated populations of N. ventricu-lus. One wonders how many species of Niptus restricted toisolated caves in southwest USA and Mexico still remainto be discovered and described.

Spider beetles have been reported to colonize corpsesduring the ultimate stages of postmortem decay, notablywhen the cadaver reaches an advanced stage of desiccation(Smith 1986; Byrd & Castner 2009). The beetles usuallyscavenge upon the remnant tissue and other organic remainsor debris left by the previous waves of necrophilous insects.Different ptinid species have been recovered from funeraryarchaeological contexts, notably Gibbium psylloides(Czempinski 1778), found in large numbers in the wrappingsof the “Two Brothers” mummies (Nekht-Ankh and Khnum-Nakht) (David 1978; Curry 1979). This species was alsopresent among the Tutankhamum tomb material (Alfieri1931) and also in the sarcophagus attributed to the Frenchcount of Toulouse, Guillaume Taillefer, in association withseveral other species of Ptinus L. (Huchet & Gallis 1996).Recently, one of the authors (JBH) recovered the remains ofPtinus sp. (cf. fur (L. 1758)) from the grave of the FrenchKingLouis XI who died in 1483 (Huchet, forthcoming).

In the same way as D. carnivorus, previously men-tioned, we are inclined to think that the colonization of themortuary bundle by this ptinid took place within the caveby 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 primar-ily necrophilous and colonizes carcasses during theadvanced stages of postmortem decay, feeding notablyupon scleroproteins (skin, hair, fur, horns, hooves, etc.)and other desiccated organic remains (e.g. tendons, andligaments) (Reed 1958; Palestrini et al. 1992).

Deloya (2000) reported 27 species of Trogidae forMexico, including 18 Omorgus species. Amongthese taxa, only five species occur in Coahuila state:O. fuliginosus Robinson 1941, O. punctatus (Germar1824), O. scutellaris (Say 1823), O. suberosus(Fabricius 1775) and O. umbonatus LeConte 1854.Several species have been reported from natural caveswhere they exploit bat guano (Scholtz 1986). MuñizVélez (2001) mentioned three Omorgus species

inhabiting the funerary caves of Morelos state inMexico (O. suberosus, O. rubricans (Robinson 1946),and O. fuliginosus). O. suberosus is undoubtedly themost common Mexican species, occurring in 20 of the31 states (Deloya 2000). Trogid beetles, like speciesbelonging to the genus Dermestes (Coleoptera:Dermestidae), only feed on desiccated carcasses thatare subaerially exposed. Omorgus females oviposit inthe sediment subjacent to the primary source of food ata depth ranging between 15 and 25 cm depending onthe species (Baker 1968). In an archaeological context,their association with buried human remains indicatesthat the corpse was temporarily exposed over a rela-tively long period (Ubelaker & Willey 1978; Huchet &Greenberg 2010). The presence of Trogid remains in thefunerary bundle indicates that the corpse was directlyaccessible to these insects and consequently remainedunburied. In all likelihood, the colonization took placeduring the dry decay stage, the insects attracted bydesiccated skin and ligaments.

The identification of the remains at the generic levelwas made from an abdominal sternite (Figure 18) and afragment of pronotum (Figure 19).

Discussion

From the composition of the thanatophilous assemblagerecovered within the funerary bundle, some hypothesesabout the funerary treatment of the young child can beproposed.

Surprisingly, no calliphorid remains were recoveredfrom the sampling, though this might be attributable tothe fact that the sample studied here represents a tiny partof the entomofauna from the bundle. Additionally, nocalliphorid remains were reported in the Vergara-Pinedaet al. (2009) study. Due to their ability to locate andcolonize a cadaver soon after death, calliphorids (blow-flies) are very often involved in forensic investigations inthe estimation of the PMI (Smith 1986; Greenberg 1991;Greenberg & Kunich 2002; Szpila 2010 inter alia). In thepresent case, their absence may be explained in two ways:either a short exposure of the body prior to burial and/or arapid wrapping of the child in the funerary bundle, or dueto the seasonality of death.4

With regard to the biological data concerningSarcophagidae and Fanniidae, we are inclined to thinkthat these flies probably arrived at a later stage, after thecorpse exhaled the first odors of decay. Although it isgenerally agreed that flies rarely oviposit in the dark(Nuorteva 1977; Greenberg 1990), we are inclined tothink that, in the present case, the location of the bodiesin the cave, at a few meters from the entrance, did notprevent the colonization of the funerary bundle. In thisrespect, the garment, heavily impregnated with bodily

286 J.-B. Huchet et al.

fluids and other liquefying tissues, probably played a veryattractive role.

Concerning the colonization by the different beetletaxa, some hypotheses regarding the chronology of theirarrival can be suggested. In all likelihood, histerid andstaphylinid beetles invaded the bundle more or less simul-taneously with the necrophagous flies (the presence ofthese later being inherent with the persistence of damporganic matter). Both families of beetles actively preyupon maggots and their presence probably dates fromthe first few weeks following the young individual’sdeath. On the other hand, the colonization of the bundleby the Dermestidae, Ptininae and Trogidae correspond to alater stage, after the human remains reached a significantdegree of desiccation. However, among these latter taxa,dermestid beetles are known to sometimes invade within ashort period after death (Early & Goff 1987;VanLaerhoven & Anderson 1999; Oliva 2001).

In the present case, the most relevant data arebased upon the biology of the muscid fly Synthesiomyianudiseta. As previously mentioned, at the postfeedingstage the larvae secrete a sticky substance which, underthe effect of the last movement prior to molting, covers thepuparium with soil particles or any other exogenous mate-rial nearby. Fragmented remains of S. nudiseta cocoonsrecovered within the funerary bundle proved to be of greatvalue and raised interesting questions. A careful examina-tion reveals that they include many remains of dermestidlarval skins (exuviae) and presumed dermestid fecal pel-lets, indicating that S. nudiseta probably invaded thecadaver at a later stage than usual, probably after dermes-tid infestation. This would suggest that the bundle hadbeen, intentionally or not, temporarily rehydrated, prob-ably in the few weeks following death. From this hypoth-esis, different scenarios might be envisaged. A firsthypothesis would be that the bundle had been placed inthe cave after a certain period of time, perhaps the timetaken to transport the bundle to the burial site from theplace of death. The corpse could have been exposed in theopen air before its final placement in the mortuary cave.However, in this case, we are inclined to think that thepioneer calliphorid flies would have probably colonizedthe corpse and some of their remains would be present inthe samplings. A second hypothesis would be that thehunter–gatherer semi nomads had temporarily or evenperiodically removed the deceased ancestors from thecave in order to satisfy some funerary practices whichwould have left no trace in the archaeological record (thecave entrance having been able to play the role of a“connecting zone” between the World of the Living andthe World of the Dead). However, it is reasonable toexpect that the cave entrance, located on a steep slope,would have probably restricted such practices. We couldnot totally preclude that the bundle had been rehydratedby isolated or repeated episodes of water seepage. Finally,

the existence of different funerary rituals in the core of thecave (e.g. ritual aspersion of the deceased?) could be at theorigin of the late colonization by the muscid fly S.nudiseta.

Conclusion

This archaeoentomological study of a sample recoveredfrom a Mexican funerary bundle shows a standard combi-nation of the main invertebrate thanatofauna associatedwith decaying cadavers. The high specific diversity defi-nitely indicates that several colonization episodes occurredand thus a relatively easy access to the cadaver was likelythroughout the process of bodily decay. In the presentcase, the lack of the pioneer flies (Calliphoridae) indicatethat the corpse was apparently shielded early on frominsect activity (rapid wrapping in the bundle withoutbody exposure prior to burial, or the death may haveoccurred during a period when fly activity is low).Finally, the late or postponed colonization by the muscidfly S. nudiseta, evidenced by the nature of the exogenouselements used to build the cocoons, may indicate that thebundle may have been occasionally rehydrated, suggestingsecondary handling of the deceased (e.g. occasionalremoval of the bundle(s) from the funerary cave?).Further studies of the invertebrate thanatofauna recoveredfrom Mexican funerary bundles may result in furtherinsights into the funerary practices of these prehistoricpopulations.

Acknowledgments

We would like to thank our colleagues G. Arriagada (Chili) andD. S. Verity (California, USA) who verified the identification ofthe 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 photo-graphs of the insect remains illustrating this paper. Finally, weare especially indebted to K. Irby who kindly assisted in improv-ing the English of this paper.

Notes1. Dating carried out on teeth collected from several crania

from Candelaria, using accelerator mass spectrometry atthe Oxford University.

2. Taxonomic identifications of the insect remains have beenrespectively performed by Y. Gomy (Coleoptera:Histeridae), T.K. Phillips (Coleoptera: Anobiidae:Ptininae), C.E. Alatorre-Bracamontes and M. Vásquez-Bolaños (Hymenoptera: Formicidae), J.L. NavaretteHeredia (Mexico) (Coleoptera: Staphylinidae) and J.-B.Huchet (Diptera: Muscidae, Fannidae, Sarcophagidae;Coleoptera: Dermestidae and Trogidae).

3. Ants of some basal Attini genera (e.g. Cyphomyrmex)occasionally use small arthropod cadavers as substratumfor fungi cultivation.

4. Recent (unpublished) experiments conducted by one of theauthors (JBH) in Peru evidenced that, depending on the

Annales de la Société entomologique de France (N.S.) 287

season of the year, different muscid or sarcophagid fliescan replace the pioneer flies (calliphorid) during the firststage of decay.

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