Annales de la Société entomologique de France (N.S.), 2013 Vol 49, No 3, 277–290, http://dx.doi.org/10.1080/00379271.2013.845474 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 10 familles et 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: huchet@mnhn.fr © 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 278 J.-B Huchet et al 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 m high and 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 Figure 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 and DAF/ INAH; Figure R Enríquez DAF/INAH) 280 J.-B Huchet et al 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 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 > 10 > 10 > 59 5 1 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 282 J.-B Huchet et al 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 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 284 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 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; 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