The Holocene vertebrate fauna from Guenfouda site, Eastern Morocco

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The Holocene vertebrate fauna from Guenfouda site, Eastern MoroccoHassan Aouraghea; Jordi Agustib; Brahim Ouchaouc; Salvador Bailond; Juan Manuel Lopez-Garciae;Hamid Haddoumia; Kamal El Hammoutia; Aicha Oujaaf; Bouchra Bougarianec

a Faculté des Sciences, Centre Universitaire de Recherches en Archéologie et Laboratoire desGéosciences Appliquée, Université Mohamed 1er, Oujda, Morocco b ICREA, Institut de PaleoecologiaHumana i Evolució Social, Universitat Rovira i Virgili, Tarragona, Spain c Faculté des Sciences,Université Moulay Ismaïl, Meknès, Morocco d Muséum National d'Histoire Naturelle, Paris, France e

Institut de Paleoecologia Humana i Evolució Social, Universitat Rovira i Virgili, Tarragona, Spain f

Institut National des Sciences de l'Archéologie et du Patrimoine, Rabat, Morocco

First published on: 27 April 2010

To cite this Article Aouraghe, Hassan , Agusti, Jordi , Ouchaou, Brahim , Bailon, Salvador , Lopez-Garcia, Juan Manuel ,Haddoumi, Hamid , El Hammouti, Kamal , Oujaa, Aicha and Bougariane, Bouchra(2010) 'The Holocene vertebrate faunafrom Guenfouda site, Eastern Morocco', Historical Biology, 22: 1, 320 — 326, First published on: 27 April 2010 (iFirst)To link to this Article: DOI: 10.1080/08912961003701193URL: http://dx.doi.org/10.1080/08912961003701193

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The Holocene vertebrate fauna from Guenfouda site, Eastern Morocco

Hassan Aouraghea*, Jordi Agustib, Brahim Ouchaouc, Salvador Bailond, Juan Manuel Lopez-Garciae, Hamid Haddoumia,

Kamal El Hammoutia, Aicha Oujaaf and Bouchra Bougarianec

aFaculte des Sciences, Centre Universitaire de Recherches en Archeologie et Laboratoire des Geosciences Appliquee, UniversiteMohamed 1er, Oujda, Morocco; bICREA, Institut de Paleoecologia Humana i Evolucio Social, Universitat Rovira i Virgili, Tarragona,Spain; cFaculte des Sciences, Universite Moulay Ismaıl, Meknes, Morocco; dMuseum National d’Histoire Naturelle, Paris, FranceeInstitut de Paleoecologia Humana i Evolucio Social, Universitat Rovira i Virgili, Tarragona, Spain; fInstitut National des Sciencesde l’Archeologie et du Patrimoine, Rabat, Morocco

(Received 20 January 2010; final version received 11 February 2010)

The site of Guenfouda, located 30 km South of Oujda in the Jebel Metsila, which forms part of the Monts d’Oujda, wasdiscovered in 2003. The site is of significant archaeological importance and systematic excavations have been undertakenannually since 2004. The upper levels of the cave deposits, dated to the Holocene, have yielded a very rich and well-preserved faunal assemblage comprising several taxa from the Mediterranean area, particularly within the sub-humid, semi-arid and arid bioclimatic stages. The most important feature of the faunal assemblage is that it includes species recognisablefor the first time as fossils in Morocco. The associated lithic industry is of good quality and includes cores, flakes and varioustools. The ceramic remains are very abundant, and some of these probably attest to occupation in the ancient Neolithic.

Keywords: vertebrate fauna; Holocene; Guenfouda; Eastern Morocco

Les vertebres holocenes de la grotte de Guenfouda (Maroc oriental)La grotte de Guenfouda se trouve dans un massif des Monts d’Oujda; elle est situee a 30 km au Sud de la ville d’Oujda. Lespremieres fouilles ont eu lieu en 2004 et se poursuivent chaque annee par une equipe de chercheurs et d’etudiants del’Universite Mohamed Premier d’Oujda en collaboration avec des institutions nationales et internationales. Le decapage desniveaux superieurs du remplissage d’age Holocene, montre une grande richesse en vestiges fauniques bien conservesappartenant a plusieurs taxons evoluant dans le domaine mediterraneen, principalement a l’interieur des etagesbioclimatiques subhumide, semi-aride et aride.

Un des points caracterisant cette faune, c’est l’identification de certaines especes pour la premiere fois a l’etat fossile auMaroc.

La faune a ete associee dans tous les niveaux fouilles du remplissage a une industrie lithique riche et variee et a des tessonsde ceramique tres abondants, dont certains portent des motifs de type Cardial, evoluant dans le Neolithique ancien.

Mots cles: faune de Vertebres; Holocene; Guenfouda; Maroc oriental

1. Introduction

Guenfouda cave (also called Ghar Zebouj) is located 6 km

from the village of Guenfouda in the province of Jerada in

Eastern Morocco (Figure 1).

The cave was formed in the Aalenian–Bajocian

(Jurassic) dolomites and oolitic limestones of the cliff of

Jebel Metsila, in the Monts d’Oujda (Figure 2). The cave

is a large gallery consisting of two main chambers

separated by a central stalagmite pillar and lies at an

altitude of 930 m.

The archaeological and paleontological importance of

Guenfouda was recognised in 2003 by A. Billy and

H. Aouraghe. It was confirmed during the first

fieldwork conducted in 2004, during which important

archaeological material has been discovered (Aouraghe

et al. 2005, 2008).

The infill deposits of Guenfouda cave appear to be

deep. At the surface there is an ashy level with stones bearing

signs of having been heated, which is rich in archaeological

material paralleling the upper Neolithic levels of Rhafas

cave, located at 15 km northeast of Guenfouda, and the Ibero-

Maurusians levels of Pigeons Cave in Tafoughalt, in the

Beni-znassen Mountains (Figure 1).

The excavated Neolithic upper levels of the cave

reveal a rich lithic assemblage, shards of pottery variously

decorated and abundant wild or domestic faunal bones, in

association with human remains. Preliminary results of the

archaeological and paleontological explorations of

ISSN 0891-2963 print/ISSN 1029-2381 online

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DOI: 10.1080/08912961003701193

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*Corresponding author. Email: [email protected]

Historical Biology

Vol. 22, Nos. 1–3, March–June–September 2010, 320–326

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the cave have been the subject of a brief publication

(Aouraghe et al. 2008). The present work treats the study

of all excavated vertebrate fauna including the remains

unearthed at the excavation campaign of 2009.

2. The vertebrate faunas

The vertebrate fossil material excavated from the

Guenfouda cave deposits attests the existence of a rich

and varied fauna, composed of more than 40 species or

genera belonging to many groups:

Hominids (Homo sapiens); Carnivores (Canis famil-

iaris, Vulpes vulpes, Caracal caracal, Mustelide);

Herbivores (Equus asinus, Equus sp. (? mauritanicus),

Sus scrofa ou Sus scrofa f. domestica, Bos primigenius,

Alcelaphus buselaphus, Oryx sp., Ammotragus lervia,

Gazella (two species), Ovis aries, Capra hircus); Rodents

(Hystrix cristata, Mus spretus, Meriones shawi, Dipodillus

campestris, Eliomys sp., Atlantoxerus getulus, Ctenodac-

tylus sp.); Lagomorphs (Lepus capensis, Oryctolagus

cuniculus); Insectivores (Atelerix algirus, Crocidura

russula, Crocidura whitakeri, Elephantulus rozeti);

Amphibians (Discoglossus pictus, Bufo mauritanicus, Bufo

viridis, Rana sp.); Reptiles (Testudo, Emys ou Mauremys,

Trogonophis wiegmanni, Agama cf. bibronii, Chamaeleo

chamaeleon cf. Eumeces, Chalcides, Malpolon, Coronella

cf. girondica, Natrix maura, Gekkonidae (species ?) and

Lacertidae (species ?)); Birds (Struthio camelus and other

birds, including raptors, recovered in the sieve).

The site has yielded a significant number of bones and

teeth in varied states of preservation. There are two types

of fossils present, depending on the nature of the sediment

and the presence of concretion carbonates from karstifica-

tion: the first type are bones coated with a layer of

carbonate concretion located in the interior of the cave,

and the second type are those without concretions from the

Figure 1. Location of the Guenfouda cave (Eastern Morocco).

Figure 2. Position of the Guenfouda cave within Jebel Metsila(Monts d’Oujda).

Historical Biology 321


square at the entrance of the cave, the sediments of which

are more earthy and powdery.

2.1 Rodents, insectivores, chiropters andmacroscelideans

(Jordi Agustı & Juan Manuel Lopez-Garcıa)

Rodents

The site of Guenfouda includes the following rodent species:

Mus spretus,Mus shawi,Dipodillus campestris,Eliomys sp.,

A. getulus, Ctenodactylus sp. and H. cristata (Figure 3).

Meriones spretus Lataste, 1883 (Figure 3(C))

Number of identified specimens (NISP): 161.

Minimal number of individuals (MNI): 70.

Remarks. This is by far the most common species of the

site. In Guenfouda, the first lower m 1 of this species

presents a clearly tetralobated anterocone, with well-

developed labial lobe of the anterocone (tE), differing in

this respect from the domestic species Mus musculus

domesticus that today lives in the area. Another difference

between M. spretus and M. musculus is that the former

colonises rocky and sandy areas, in addition to the

mountains, being much more resistant to dryness and

being able to persist in zones with poor vegetation.

Meriones shawi (Rozet, 1833) (Figure 3(D))



Remarks. This rather large gerbil represents the second

most common species from the sample. Today, it inhabits

quite diversified environments including clay and sandy

soils, more rarely rocky environments.

Dipodillus campestris Le Vaillant, 1867 (Figure 3(B))



Remarks. The species D. campestris has been allocated in a

number of genera or subgenera, including Hendecapleura

and Dipodillus (we follow this last group here). This species

inhabits a variety of environments (for example daıas,

sebkha, rocky biotopes, etc.), and is frequently commensal

with humans where there is grain accumulation.

Eliomys sp.



Remarks. Glirids (dormice) are very rarely represented at

Guenfouda. We identified only an upper maxilla retaining the

P 4 and a very worn upper M 3 in which the dental design

cannot be recognised, both showing the typical morphology

of the genus Eliomys. Until recently, all the specimens from

this genus in North Africa were assigned to the European

species Eliomys quercinus, although a number of subspecies

were recognised. Now it seems clear that the North African

populations belong to two different species, Eliomys

munbyanus and Eliomys melanurus. Both tend to colonise

zones with quite abundant vegetation as well as rocky or

mountain environments. Given the scarce representation of

this genus at Guenfouda, it is impossible for the moment to

assign the examples to one or another species.

Atlantoxerus getulus (Linnaeus, 1758) (Figure 3(A))



Remarks. A. getulus is a ground squirrel which today

occupies most of the western Maghreb. It inhabits rocky

Figure 3. Small mammals from the Guenfouda cave. (A)A. getulus, left P4-M1 (occlusal view); (B) Dipodillus campestris,left M1-M3 (occlusal view); (C) Mus spretus, right M1-M2(occlusal view); (D) Mus shawi, right M1-M2 (occlusal view); (E)Crocidura russula, right mandible (lingual view); (F) Crocidurawhitakeri, left mandible (lingual view); (G)Elephantulus rozeti, leftM2 (occlusal view); (H) Myotis blythii, left M3 (occlusal view).

H. Aouraghe et al.322


environments without trees, where it can be found at

altitudes of up to 3,000 m.

Ctenodactylus sp.



Remarks.Gundis are represented at Guenfouda by an upper

P 4. Although singular, its presence is significant. The

most common species,Ctenodactylus gundi, today inhabits

rocky environments in arid areas in the south of Morocco.

A second smaller species, Ctenodactylus vali, lives in

similar places. The size of the P 4 seems to relate to the

former, most common, species. However, given the

scarcity of this species in Guenfouda, we await further

material to provide specific determination to continue.

Hystrix cristata



Remarks. H. cristata (porcupine) is represented in

Guenfouda by 10 dental remains from at least four

individuals including two young. The presence of the

porcupine is also attested by numerous typical incisor bite

marks left on the bones of large mammals.

Rare in the early Quaternary, H. cristata became very

common in the North African middle and upper

Pleistocene as its presence is attested in several sites

including Carriere Thomas (Bernoussi 1997), El Harhoura

1 (Aouraghe and Abassi 2002) and Bouknadel, Doukkala

II (Michel 1990). It currently exists in Morocco all

the way from the North to the borders of the Sahara

(Aulagnier 1992).

2.2 The Lagomorphs

(B. Bougariane)



The lagomorphs are represented by two species:

O. cuniculus (MNI ¼ 2) and L. capensis (MNI ¼ 2).

These species are attested by two pelvic fragments of

different sizes. Among the remains of lagomorphs, we

note the presence of two unfused distal epiphyses

of femurs. They are noticeably large, despite coming

from young individuals. Based on dimensions, these

two fossils can be attributed to L. capensis (hare).

However, two other remains recall the rabbit

O. cuniculus. These are a distal end of a humerus

with a very prominent medial tubercle and a distal tibia

with a depression on the shaft above the epiphysis.

These two anatomical characters are typical of bones

from O. cuniculus (Callou 1997).

2.3 The Amphibians and Reptiles

(S. Bailon)

Amphibians and reptiles are represented by at least 17

taxa belonging to 4 anuran amphibians (D. pictus,

B. mauritanicus, B. viridis et Rana sp.) and 13 reptiles

(Testudo, Emys or Mauremys, T. wiegmanni, Agama cf.

bibronii, C. chamaeleon cf. Eumeces, Chalcides, Malpolon,

Coronella cf. girondica, N. maura. The taxonomic affinities

ofGekkonidae and Lacertidae still remain to be established).

The number of taxa represents approximately 53% of the

total herpetofauna (about 32 species) which are currently

present within a 50 km radius from Guenfouda [data on

extant species obtained from Bons and Genier (1996).

Agames and chameleons are represented, respectively, by

Agama cf. bibronii (Figure 4) and C. Chamaeleon (Figure 5).

These are the first fossils of these animals in Morocco.

2.4 Birds

The study of fossilised bird remains from the excavations

is underway. The species S. camelus is represented by a

large amount of eggshell fragments. These were some-

times sliced and perforated for necklaces. Figure shows the

various stages of manufacture of these discs of ostrich eggs

discovered at Guenfouda.

It is worthwhile to note that only fragments of

eggshells are present, but no bone remains of this species

have been discovered so far. It is not clear whether the

ostrich was among the species hunted by prehistoric man or

if only its eggs were used for decorative purposes (Figure 6).

The presence of ostrich in the Neolithic levels of Guenfouda

is synonymous with an arid climate.

2.5 Carnivores and herbivores

(Ouchaou, Aouraghe and Bougariane)

Large mammals are abundant at Guenfouda.

Figure 4. Agama cf. bibronii from Guenfouda, 1: right dental,(A) medial view and (B) side view.



https://www.researchgate.net/publication/240816387_Les_rongeurs_du_site_aterien_d'El_Harhoura_1_Temara_Maroc?el=1_x_8&enrichId=rgreq-131d7012cd0675cbc0d9b7bc0310b59e-XXX&enrichSource=Y292ZXJQYWdlOzIzMjk2Njg2NDtBUzo5OTI2MDcxNjU1MjIwMEAxNDAwNjc3MDA3NTIx

https://www.researchgate.net/publication/34811424_Contribution_a_l'etude_paleontologique_des_Vertebres_fossiles_du_Quaternaire_marocain_a_partir_de_sites_du_Maroc_atlantique_central_et_oriental?el=1_x_8&enrichId=rgreq-131d7012cd0675cbc0d9b7bc0310b59e-XXX&enrichSource=Y292ZXJQYWdlOzIzMjk2Njg2NDtBUzo5OTI2MDcxNjU1MjIwMEAxNDAwNjc3MDA3NTIx

Carnivores (NISP: 41).

Forty-one fossilised remains are from carnivores, rep-

resented by four species, including two Canidae: Canis

lupus f. familiaris, V. vulpes (Figure 7(A)), a medium-

sized Felida with morphology similar to C. caracal, and a

small carnivore that could be a mustelid.

Herbivores (NISP: 336)

The herbivores are abundant and comprise diverse families,

including equids, suids and bovids. The accumulation of

bones in the infilling is mainly anthropogenic.

Equidae (NISP: 24) (Figure 7(C))

The Equids are represented by teeth and bone fragments

(NISP: 24). Based on the degree of tooth wear, they can be

classified into three classes: young, adult and older.

The remains attributed to horses belong to two species:

one small, E. asinus, and one medium-sized Equus sp.,

probably a Zebrine. The latter is recognised in several sites

in North Africa under the name of Equus mauritanicus.

Most paleontologists doubt on this attribution, and it will

be necessary to review the equid materials from old

excavations in North Africa.

Suidae (NISP: 11) (Figure 7(D))

The suids include a few remnants belonging to young and

adult individuals. However, in the absence of canine and

third molars, the distinction between Sus scrofa

(Wild Boar) and Sus scrofa f. domestica (pig) is not

always obvious.

Bovidae (NISP: 301)

The bone remains assigned to Bovidae are the most

frequent and most diverse in the Guenfouda cave deposits.

They include five subfamilies: Bovini, Alcelaphini,

Hippotraginids, Antilopini and Caprini.

The Bovini include B. primigenius f. taurus (Figure

7(G)). The Antilopini include two different-sized species

of Gazella (Figure 7(F)). The Caprini are the most

abundant, with A. lervia (Figure 7(E)), C. hircus and O.

aries (Figure 7(I)).

Among the Alcelaphini remains, there are three

isolated lower molars (M1, M2 and M3) of A. buselaphus,

representing the same individual. The dimensions of A.

buselaphus found at Guenfouda (Table 1) show some

resemblance with those known from the sites of

Bouknadel, Doukkala II (Michel 1990), Kehf-el-Baroud

(Ouchaou 2000) and El Harhoura 1 (Aouraghe 2001).

Similarly, the dimensions of astragals of A. buselaphus

from Guenfouda fall within the variation limits of

specimens from several prehistoric sites and extant ones,

but they are smaller than those found at the sites Doukkala

II and El Harhoura 1 (Figure 8).

Hippotraginids include one lower molar (M1-2)

distinguished by a characteristic caprin fold (anterosty-

lide), detached from the mesial lobe and carinated external

pillars. The dental remains probably belong to the genus

Oryx. Its dimensions are (DMD £ DVL ¼ 22.5 £ 10.2

mm). These dimensions are comparable to those of Oryx

M1 of El Harhoura 1, whose values are DMD £ DVL:

23.2 £ 12 mm (Aouraghe 2001).

2.6 Hominids

(A. Oujaa)

Human remains discovered between 2004 and 2007

include post-cranial bones, some bones of hands and feet,

long bone fragments (tibia, fibula and femur) and a piece

of collarbone. These bones are characterised by a general

slenderness, all belonging to mature individuals in good

health, with no evidence of bone disease. Identification of

sex and age is not possible at the moment.

During the 2007–2009 campaigns, other human remains

were brought to light, including a mostly intact burial,

damaged by the weight of the pile of boulders (Figure 9).

Figure 5. C. chamaeleon from Guenfouda, (2) right maxillary,lateral view and (3) left dental fragment, lateral view.

Figure 6. Fragments of eggshell of S. camelus transformed intoadornment objects (Scale: £ 1).




3. Conclusion

The archaeological and paleontological importance of the

Guenfouda cave site is its abundant fossil and lithic

assemblages. The fauna is very rich, varied, and includes

more than 40 species or genera.

This faunal assemblage includes fossilised examples

of the species C. chamaeleon (Chamaeleonidae) and

Agama cf. bibronii (Agamidae) (Figures 5 and 6), and this

is the earliest known example of fossils of these two

species in Morocco.

Table 1. Comparative dimensions of A. buselaphus lower molars from different Quaternary sites.

M1 M2 M3

Sites DMD DVL DMD DVL DMD DVL

Guenfouda 21.0 11.0 23.0 13.0 31.0 11.0Bouknadel (1) 16.3–22.9 9.8–11.3 20.4–22.3 10.9–12.4 29.5–31.2 10.0–10.8Doukkala II (1) 17.0–20.5 10.9–11.7 23.9–25.0 11.0–12.7 29.5–31.2 10.0–12.3Kehf-el-Baroud (2) 17.0 10.3 22.0 12.0 30.5–31.2 11.9El Harhoura I (3) 19.5–24.7 8.4–11.4 22.1–24.0 9–13.4 30.3–32.1 10.8–12.5

Notes: (1) Michel 1990, (2) Ouchaou 2000 and (3) Aouraghe 2001.

Figure 7. Large mammal remains from Guenfouda. (A) Canis familiaris and Vulpes vulpes, (B) Alcelaphus buselaphus, (C) Equussp. and Equus asinus, (D) Sus scrofa or Sus scrofa f. domestica, (E) Ammotragus lervia, (F) Gazella, (G) Bos primigenius, (H) lagomorphsand (I) Ovis aries, Capra hircus.



The quaternary infilling seems very deep. The date the

upper levels to the Holocene, corresponding to Neolithic

culture, is established by the presence of pottery sherds with a

Cardial type pattern, dated to the ancient Neolithic, by

typical Neolithic tools (polished axes and grinding wheels),

and by the presence of domesticated fossil animal species.

Most faunal taxa recovered at Guenfouda, especially

small vertebrates, evolved in the Mediterranean region,

mainly in the sub-humid, semi-arid and arid bioclimatic

zones.

That there were water points in the vicinity of the site is

attested by the presence of aquatic species such as D. pictus,

Rana, N. maura and freshwater turtles. The landscape seems

to have alternated between areas with vegetation cover type

or bushy scrubland and more or less open areas.

Further excavations and the ongoing multidisciplinary

studies will certainly provide additional data on the

paleontological significance of the site, and without doubt

will bring more information details of the vertebrate

Quaternary vertebrate fauna from Guenfouda.

Acknowledgements

These investigations conducted by the University Center forArchaeological Research (CURA), fall under the GlobalResearch Project (PGR), funded by the University Mohamed1er, Oujda, for the recovery of the archaeological heritage ofEastern Morocco. J.M.L.-G. has been supported by a postdoctoralcontract of the Subprogram Juan de la Cierva (Spanish Ministryof Science and Innovation).

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Mouhsine T. 2003. Contribution a la connaissance de la faune duPleistocene superieur du Rif oriental: la grotte Ifri N’Ammar.Paleontologie, Archeozoologie, Paleoecologie, Biochronologie etBiogeographie. These de 3eme Cycle a l’INSAP, Rabat, 305 p., 30 pl.

Ouchaou B. 2000. Les faunes mammalogiques holocenes des gisements dunord du Maroc: etude paleontologique et observations archeozoologi-ques. These d’Etat, Universite Moulay Ismaıl, Meknes, 369 p.

Peters J, Van Neer W, Plug I. 1997. Comparative postcranial osteology ofHartebeest (Alcelaphus buselaphus), Scimitar oryx (Oryx dammah)and addax (Addax nasomaculatus), with notes on the osteometry ofgemsbok (Oryx gazella) and arabian oryx (Oryx leucoryx). Ann SciZool. Musee Royale de l’Afrique Centrale, Tervuren, Belgique280:1–83.

Figure 8. Comparative dimensions (height of external side) ofastragals of A. buselaphus mm. (1) Michel (1990), (2) Aouraghe(2001), (3) Ouchaou (2000), (4) Mouhsine (1997), (5) Mouhsine(2003), (6) Merzoug (2005) and (7) Peters et al. (1997).

Figure 9. Partial sketch of the Neolithic burial from Guenfouda(Haddoumi).





https://www.researchgate.net/publication/292812344_Comparative_postcranial_osteology_of_hartebeest_Alcelaphus_buselaphus_scimitar_oryx_Oryx_dammah_and_addax_Addax_nasomaculatus_with_notes_on_the_osteometry_of_gemsbok_Oryx_gazella_and_Arabian_oryx_Oryx?el=1_x_8&enrichId=rgreq-131d7012cd0675cbc0d9b7bc0310b59e-XXX&enrichSource=Y292ZXJQYWdlOzIzMjk2Njg2NDtBUzo5OTI2MDcxNjU1MjIwMEAxNDAwNjc3MDA3NTIx











The Holocene vertebrate fauna from Guenfouda site, Eastern Morocco

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