Adulis and the transshipment of baboons during classical antiquity
Abstract
Adulis, located on the Red Sea coast in present-day Eritrea, was a bustling trading centre between the first and seventh centuries CE. Several classical geographers—Agatharchides of Cnidus, Pliny the Elder, Strabo—noted the value of Adulis to Greco-Roman Egypt, particularly as an emporium for living animals, including baboons (Papio spp.). Though fragmentary, these accounts predict the Adulite origins of mummified baboons in Ptolemaic catacombs, while inviting questions on the geoprovenance of older (Late Period) baboons recovered from Gabbanat el-Qurud (‘Valley of the Monkeys’), Egypt. Dated to ca. 800–540 BCE, these animals could extend the antiquity of Egyptian–Adulite trade by as much as five centuries. Previously, Dominy et al. (2020) used stable isotope analysis to show that two New Kingdom specimens of Papio hamadryas originate from the Horn of Africa. Here, we report the complete mitochondrial genomes from a mummified baboon from Gabbanat el-Qurud and 14 museum specimens with known provenance together with published georeferenced mitochondrial sequence data. Phylogenetic assignment connects the mummified baboon to modern populations of P. hamadryas in Eritrea, Ethiopia, and eastern Sudan. This result, assuming geographical stability of phylogenetic clades, corroborates Greco-Roman historiographies by pointing toward present-day Eritrea, and by extension Adulis, as a source of baboons for Late Period Egyptians. It also establishes geographic continuity with baboons from the fabled Land of Punt (Dominy et al., 2020), giving weight to speculation that Punt and Adulis were essentially the same trading centres separated by a thousand years of history.
Editor's evaluation
This fundamental Research Advance sheds new light on the ancient baboon trade in the Red Sea. Combining ancient DNA methods from a mummified baboon with historical accounts, this work provides compelling evidence connecting the ancient Egyptian trade of baboons with the ancient port city of Adulis. The results will be of significance to a broad range of scholars interested in applying ancient DNA to improve our knowledge of historical events.
https://doi.org/10.7554/eLife.87513.sa0Introduction
Adulis, on the coast of present-day Eritrea, was an important hub during the rise of cross-ocean maritime trade, connecting ships, cargoes, and ideas from Egypt, Arabia, and India (Burstein, 2002; Munro-Hay, 1982; Seland, 2008). Trade peaked between the fourth and seventh centuries CE, propelling the rise and expansion of the Aksumite kingdom, but its occupation history extends, at minimum, to the first millennium BCE (Zazzaro et al., 2014). Corroborating this archaeological record are written accounts that draw attention to the importance of Adulis as one of the foremost sources of African animals or animal products during the Hellenistic period (323–31 BCE). In Topographia Christiana, a sixth-century text, the Nestorian merchant Cosmas Indicopleustes recounts his own visit to Adulis in 518 CE (Fauvelle-Aymar, 2009; Hatke, 2013). There he copied the text of a stele inscribed in Greek and known today as the Monumentum Adulitanum I. The text celebrates the military conquests of Ptolemy III Euergetes (reign: 246–222 BCE) and notes the local availability of war elephants for himself and his predecessor, Ptolemy II Philadelphus (reign: 284–246 BCE) (Bowersock, 2013).
Echoing this account is the first-century Periplus Maris Erythraei, an anonymous text focused on maritime trade across the Red Sea Basin: ‘practically the whole number of elephants and rhinoceros that are killed live in the places inland, although at rare intervals they are hunted on the seacoast even near Adulis’ (Casson, 1989; Casson, 1993). Pliny the Elder described Adulis as a thriving emporium in his Naturalis Historia, another first-century text, and commented on the availability of ivory, rhinoceros horn, hippopotamus hides, tortoise shell, and sphingia—or ‘sphinx monkeys,’ a term that probably refers to the gelada, Theropithecus gelada (Jolly and Ucko, 1969). Pliny’s account relied heavily on the writings of Agatharchides of Cnidus (ca. 145 BCE), who described ‘Aithiopia’ (meaning the Red Sea coast and African hinterlands) as a source of sphinx monkeys, cepi (probably patas monkeys, Erythrocebus patas; Burstein, 1989), and cynocephali—or ‘dog-heads.’ Strabo’s Geographica references the worship of cynocephali at Hermopolis (Egypt), making it clear that the animal in question is the hamadryas baboon (Papio hamadryas), the traditional sacred animal of the Egyptian god Thoth (Figure 1). The source of baboons in ancient Egypt is an enduring question (Dominy et al., 2020) as the current distribution of baboons excludes Egypt (Figure 2) and there is no prehistoric evidence of baboons occurring in Egypt naturally (Geraads, 1987).

Strabo’s reference (17.1.40) to the worship of cynocephali at Hermopolis Magna makes clear that the animal in question is the hamadryas baboon (Papio hamadryas).
The sanctuary and temple complex featured several 35-tonne statues of P. hamadryas as the embodiment of Thoth. One of the oldest deities in the Egyptian pantheon, Thoth is best known as a god of …

Present-day distributions of the six baboon species, major mitochondrial clades, and provenance of samples analysed in this study.
(a) Overview of species distributions according to the IUCN (2020) and coloured by species (red: P. papio; brown: P. ursinus; yellow: P. cynocephalus; orange: P. kindae; green: P. anubis; purple: P. …
Though fragmentary, this historiography points to Adulis as a commercial source of mummified baboons in Ptolemaic catacombs, such as those at Saqqara and Tuna el-Gebel (Goudsmit and Brandon-Jones, 1999; Peters, 2020) [or those of their progenitors if Ptolemaic Egyptians maintained captive breeding programs; (von den Driesch et al., 2004)]. At the same time, these accounts invite questions focused on the source of pre-Ptolemaic baboons recovered from Gabbanat el-Qurud, Egypt (Lortet and Gaillard, 1907) and dated to ca. 800–540 BCE (Richardin et al., 2017), a span that corresponds to the 25th Dynasty and Late Period of Egyptian antiquity. If these older specimens can be traced to Eritrea, and by extension Adulis, then they have the potential to extend the time depth of Egyptian–Adulite trade by as much as five centuries.
Mummified baboons have been investigated morphologically, revealing species-level taxonomic assignments as well as individual details, such as age, sex, and pathological condition (Boessneck, 1987; Brandon-Jones and Goudsmit, 2022; Goudsmit and Brandon-Jones, 1999; Peters, 2020). Such data are telling, but insufficient for determining fine-scale geographic origins. Recent oxygen and strontium stable isotope evidence suggests that mummified hamadryas baboons were imported from a region encompassing northern Somalia, Eritrea, and Ethiopia (Dominy et al., 2020), a level of geographic precision with limited practical value. Another limitation concerns the captive breeding of some animals. For instance, stable isotopes can reveal a lifetime in Egypt but not the geoprovenance of the source population, as shown for olive baboons from the Ptolemaic catacombs of North Saqqara (Dominy et al., 2020). The analysis of ancient DNA (aDNA) recovered from baboon mummies and compared to the current distribution of baboon genetic diversity has the potential to provide more detailed insights on the geographic origin of baboons in ancient Egypt. To explore this possibility, we sequenced the mitochondrial genome (mitogenome) of a mummified baboon to infer its geographic origin through phylogenetic assignment.
Gabbanat el-Qurud
In Topography of Thebes, Wilkinson, 1853 noted a site called Gabbanat el-Qurud (‘Valley of the Monkeys’) located ca. 2.5 km north–northwest of Medinet Habu, the mortuary temple of Ramses III. Intrigued by this observation, French Egyptologists Louis Lortet and Claude Gaillard sought and found the site in February 1905, along with the remains of mummified baboons. They recovered ‘17 skulls and a large quantity of bones,’ which they attributed to Papio anubis and P. hamadryas (Lortet and Gaillard, 1907). The assemblage includes juvenile and adult males and females buried in jars, sarcophagi, or wooden coffins. Now accessioned in the Musée des Confluences, Lyon, France, the linen wrapping of one mummified individual (MHNL 90001206) was dated radiometrically to 803–544 cal. BC (95.4%) (Richardin et al., 2017).
Ottoni et al., 2019 sampled dental calculus from 16 individuals in this same assemblage and reported the preservation of ancient microbial DNA in a subset of six. Their success motivated us to extract DNA from the remaining tooth material of ten individuals (Table 1, Supplementary file 1). In addition, we obtained samples (skin, bone, or tooth) from 21 modern historic specimens of baboons available in museum collections and representing the northeast African distribution of Papio (Table 1, Figure 2). These specimens were collected between 1855 and 1978, and we denote them ‘historic samples’ in the remainder of the article to distinguish them both from the older mummified specimens (‘mummified samples’) and recently collected material (‘modern samples’). Latitude–longitude information on the origin of the specimens was either derived from the respective museum database or assigned based on the listed provenance (Table 1).
Information on samples analysed in this study.
| Taxon | Origin | Museum ID | Country | Latitude | Longitude | MitoClade | AccNo | Reference |
|---|---|---|---|---|---|---|---|---|
| P. hamadryas | MNHN | MO-1972–357 | ETH | 9.320 | 42.119 | G3-X | OQ538080 | This study |
| P. hamadryas | SMNS | SMNS-Z-MAM-001034* | ETH | 11.500 | 39.300 | G3-X | OQ538076 | This study |
| P. hamadryas | MfN | ZMB_Mam_025647_(2) | ETH | 14.164 | 38.891 | G3-X | OQ538079 | This study |
| P. hamadryas | SMNS | SMNS-Z-MAM-000960 | ERI | 15.783 | 38.453 | G3-X | OQ538078 | This study |
| P. hamadryas | NHMUK | ZD.1910.10.3.1 | SOM | 9.933 | 45.200 | G3-X | MT279063 | Roos et al., 2021 |
| P. hamadryas | MfN | ZMB_Mam_012808 | ETH | 9.314 | 42.118 | G3-X | OQ538089 | this study |
| P. anubis | Wild | ETH | 8.968 | 38.571 | G3-X | JX946196 | Zinner et al., 2013 | |
| P. hamadryas | MfN | ZMB_Mam_042543_(1) | ETH | 9.593 | 41.866 | G3-Z | OQ538084 | this study |
| P. hamadryas | MfN | ZMB_Mam_074849 | DJI | 11.589 | 43.129 | G3-Z | OQ538085 | this study |
| P. hamadryas | MNHN | MO-1972–359 | ETH | 6.998 | 40.478 | G3-Z | OQ538086 | this study |
| P. hamadryas | SMNS | SMNS-Z-MAM-001288 | SDN | 19.110 | 37.327 | G3-Y | OQ538081 | this study |
| P. hamadryas | Wild | ERI | 15.011 | 38.971 | G3-Y | JX946201 | Zinner et al., 2013 | |
| P. hamadryas | SMNS | SMNS-Z-MAM-007509† | - | - | - | G3-Y | OQ538082 | this study |
| P. hamadryas | MHNL | 51000172 | EGY | - | - | G3-Y | OQ538083 | this study |
| P. anubis | SMNS | SMNS-Z-MAM-000584 ‡ | SDN | 13.460 | 33.780 | G3-Y | OQ538075 | this study |
| P. cynocephalus | Wild | TNZ | 7.347 | 37.165 | G1 | JX946199 | Zinner et al., 2013 | |
| P. cynocephalus | MNHN | ZM-MO-1977-5 | SOM | 3.243 | 45.471 | G1 | OQ538088 | this study |
| P. anubis | NHMUK | ZD1929.4.27.2 | COD | 0.800 | 26.633 | J | MT279061 | Roos et al., 2021 |
| P. anubis | NHMUK | ZD1929.4.27.1 | COD | 1.183 | 27.650 | J | MT279062 | Roos et al., 2021 |
| P. anubis | Wild | 19GNM2220916 | TNZ | 4.679 | 29.621 | J | MG787545 | Roos et al., 2018 |
| P. anubis | SMNS | SMNS-Z-MAM-032128 | SSD | 4.281 | 33.555 | J | OQ538087 | this study |
| P. anubis | SMNS | SMNS-Z-MAM-000583 | SDN | 13.333 | 32.729 | J | OQ538077 | this study |
| P. anubis | MfN | ZMB_Mam_074869 | CMR | 5.533 | 12.317 | F | OQ538071 | Kopp et al. in prep |
| P. anubis | Wild | NGA | 7.317 | 11.583 | F | JX946198 | Zinner et al., 2013 | |
| P. anubis | MfN | ZMB_Mam_074887 | CMR | 9.328 | 12.946 | F | OQ538069 | Kopp et al. in prep |
| P. anubis | MfN | ZMB_Mam_074885 | NGA | 7.298 | 10.318 | F | OQ538064 | Kopp et al. in prep |
| P. anubis | MfN | ZMB_Mam_074883 | CMR | 6.334 | 9.961 | F | OQ538072 | Kopp et al. in prep |
| P. papio | Wild | SEN | 12.883 | 12.767 | E | JX946203 | Zinner et al., 2013 | |
| P. anubis | NHMUK | ZD.1947.586 | SLE | 8.917 | 11.817 | E | MT279064 | Roos et al., 2021 |
| P. anubis | MfN | ZMB_Mam_075043 | TGO | 9.260 | 0.781 | D | OQ538066 | Kopp et al. in prep |
| P. anubis | MfN | ZMB_Mam_011198 | TGO | 6.228 | 1.478 | D | OQ538067 | Kopp et al. in prep |
| P. anubis | Wild | CIV | 8.800 | 3.790 | D | JX946197 | Zinner et al., 2013 | |
| P. anubis | MfN | ZMB_Mam_007396_(1) | TGO | 6.950 | 0.585 | D | OQ538065 | Kopp et al. in prep |
| P. anubis | NHMUK | ZD.1939.1022 | NER | 17.000 | 7.933 | D | MT279065 | Roos et al., 2021 |
| P. anubis | NHMUK | ZD.1939.1020 | NER | 17.683 | 8.483 | D | MT279066 | Roos et al., 2021 |
| P. anubis | MNHN | ZM-MO-1960-476 | TCD | 20.344 | 16.786 | K | MT279067 | Roos et al., 2021 |
| P. anubis | MNHN | MO-1996-2511 | CAF | 3.905 | 17.922 | K | OQ538068 | Kopp et al. in prep |
| P. anubis | NHMUK | ZD.1907.7.8.11 | CAF | 8.000 | 20.000 | K | MT279068 | Roos et al., 2021 |
| P. anubis | MNHN | MO-1996-2510 | CAF | 4.966 | 18.701 | K | OQ538070 | Kopp et al. in prep |
| P.ursinus | Wild | ZAF | 24.680 | 30.790 | B | JX946205 | Zinner et al., 2013 | |
| P. cynocephalus | Wild | TNZ | 11.261 | 37.514 | B | JX946200 | Zinner et al., 2013 | |
| P. kindae | ZMB | 12.591 | 30.252 | C | JX946202 | Zinner et al., 2013 | ||
| P. cynocephalus | Wild | 04MNM1300916 | TNZ | 6.119 | 29.730 | H | MT279069 | Roos et al., 2021 |
| P. ursinus | Wild | ZAF | 34.456 | 20.407 | A | JX946204 | Zinner et al., 2013 | |
| P. cynocephalus | Wild | 24UNF1150317 | TNZ | 7.815 | 36.895 | MT279060 | Roos et al., 2021 | |
| Theropithecus gelada | ||||||||
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