DISTRIBUTION OF HYRAX (MAMMALIA/HYRACOIDEA/PROCAVIIDAE) IN THE CENTRAL REGIONS OF BENIN.

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ISSN: 2320-5407 Int. J. Adv. Res. 4(11), 2327-2339
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Journal Homepage: - www.journalijar.com
Article DOI: 10.21474/IJAR01/2343
DOI URL: http://dx.doi.org/10.21474/IJAR01/2343
RESEARCH ARTICLE
DISTRIBUTION OF HYRAX (MAMMALIA/HYRACOIDEA/PROCAVIIDAE) IN THE CENTRAL
REGIONS OF BENIN.
*Daouda A. I. H.1, Adjanohoun M.1, Hennou A.1, Atindogbe G.2, Mensah G.A.3 and Sinsin B.4.
1. Departement d’Amenagement and Gestion des Ressources Naturelles, Faculte d’Agronomie, Universite de
Parakou (Benin).
2. Centre de Bio-statistique et Informatique Generale, Fac. Sc. Agronomiques, U.A.-C. Cotonou (Benin).
3. Institut National de Recherche Agronomique du Benin-INRAB / CRA – Agonkanmey, Cotonou (Benin).
4. Laboratoire d’Ecologie Appliquee/Fac. Sc. Agronomiques, Univ d’Abomey-Calavi (UA-C) Cotonou (Benin).
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Manuscript Info Abstract
……………………. ………………………………………………………………
Manuscript History
Received: 30 September 2016
Final Accepted: 30 October 2016
Published: November 2016
Key words:-
Hyrax, central regions of Benin,
distribution, Occurrence Area , Lama
Forest.
Few research works were carried out on hyraxes (family of
Procaviidae) in Benin until now. This study provides the first data on
the spatial distribution evolution of Procavia capensis (rock hyrax) and
Dendrohyrax dorsalis subsp sylvestris (tree hyrax) in the central
regions of Benin: if both kinds of hyrax are met in this area of Sudano -
Guinean transition, only Dendrohyrax dorsalis ssp. sylvestris are met in
Guinea zone (South - Benin). In this phytogeographical area, the tree
hyrax occupies essentially the Lama forest, and weakly met in few
forest islands and in the valley of Oueme. The past and present
occurrence areas of Dendrohyrax dorsalis subsp sylvestris were
mapped and their surface area, calculated. Due to the significant
fragmentation and degradation of habitat of tree hyrax in southern
Benin, its occurrence areas increased from 8007 km² to 4003 km²
(current occurrence area) within less than half a century. If the
classified forest of Lama gathers the largest subpopulation, the other
sites are severely fragmented, with small and isolated subpopulations in
the forest islands far between them with a distance of dozens
kilometers. Copy Right, IJAR, 2016,. All rights reserved.
……………………………………………………………………………………………………....
Introduction:-
According to IUCN (2009), 21% of mammals are considered endangered from the earth planet. Several cases have
been recorded in African countries including Benin of which the red-bellied monkey Cercopithecus erythrogaster
erythrogaster, endemic primate of Benin (Nobime and Sinsin, 2003; Sinsin et al., 2002). Similarly, some taxa
continue to die out of Benin including Tragelaphus eurycerus (Neuenschwander et al., 2011); whereas several other
taxa remain to be described or their taxonomic position may be indicated (Colyn et al., 2010; Granjon et al., 2005;
Daouda, 2002; Codjia et al., 1996). This is the case of hyraxes which were a long time mistaken for rodents, but are
now systematized in the particular order of Hyracoidea. From this order, its remains only the family of Procaviidae.
From four species described initially, this number has now increased to eleven, divided into three genus: Procavia,
Heterohyrax and Dendrohyrax. If the described species of these three genus of hyraxes were studied extensively in
East Africa and South Africa, data is rare in the west part of the continent and particularly in Benin (Kingdon, 1997;
Grubb, 1998; Bacho, 2004; Barry et al., 2008; Neuenschwander et al., 2011; Djossa et al., 2012; Daouda et al.,
2015). From their works, it appears that two genus of hyrax are met in Benin: one species of the rock hyrax
Corresponding Author:- Daouda A.I.H.
Address:- Departement d’Amenagement and Gestion des Ressources Naturelles, Faculte d’Agronomie,
Universite de Parakou (Benin) .

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(Procavia, capensis, kerstingi) and one subspecies of tree hyrax, probably similar to the Dendrohyrax dorsalis ssp
sylvestris described in Ghana as their vocalizations were concordant (Akpona et al., 2011). The country of Benin
was excluded by omission from the occurrence area of tree hyrax until 2006 (Kingdon, 2006). Meanwhile genetic
characterization of the tree hyrax in Benin, their geographic distribution area continues to be fragmented and
reduced. This work presents the area of past and current occurrence of D. dorsalis ssp, and an estimation of its
current occurrence area in Benin. Meanwhile genetic characterization of hyrax in Benin, their geographic range is
constantly being reduced and fragmented. Also the geographic distribution of badgers in the Central regions of
Benin, this work presents the past and current occurrence area of D. dorsalis subsp sylvestris in the south of Benin
where the few dense forests islands (its preferred habitat) are highly degraded.
Study Area:-
The study was carried out in two out of three phyto-geographical areas of Benin (Figure 1), including the Guinean
zone that extends from the coast to the Municipality of Djidja (7 ° 15 N), and the Guineo-Sudanese transition zone
(ranges from 7 ° - 7 ° 30 N to 9 ° 30 - 10 ° N). The preselected districts are presented in the study area (Figure 2).
The first phytogeographical zone has a bimodal rainfall with 900 mm of rain per year in the western part to 1300
mm/year in the East. This type of sub-equatorial climate is closely related to the phenomenon of "Dahomey Gap".
Thus, if the semi-deciduous rainforest can still observe at a few forest Islands, "the evergreen unit of land is
completely absent" (Neuenschwander et al., 2011; Sinsin et Kampmann, 2011).
Guinean area was divided into four phytogeographical districts namely the Coastal, the Oueme Valley, Pobè and
Plateau (Adomou et al., 2006). Only the Oueme Valley, Pobè and Plateau phytogeographical districts were taken
into account in the current work. The original vegetation is semi-deciduous dense forests and Guinean savannas.
One can note the dense semi-deciduous rainforest made of Triplochiton scleroxylon and Celti zenkeri with some
variations including the climatically dried Nesogordonia kabingaensis and Mansonia altissima, as swamp forest
with Xylopia rubescens and Mitragyna ciliata and periodically flooded forest with Dialium guineense and Berlinia
grandifolia (Oueme Valley). The forest of Lama that is seasonally flooded is rich in vegetable species such as Ceiba
pentandra, Diospyros mespiliformis, Afzelia africana, Mimusop andongensis, Cynometra megalophylla, etc.
The vegetation of the Guinean-Congolese area is certainly very strongly deteriorated because of the high population
density. The mammalian fauna of the Lama forest is composed of rodent species, reptile, cattle, small carnivores and
primates (Sinsin and Assogbadjo, 2002; Kassa et al., 2007). The second study area (Guineo-Sudanian transition) is
characterized by lower rainfall (1100 mm of rain per year) compared to the first, with a gradual merger of the two
rainfall peaks, gradually as one leaves from its south limit to the north. The Guinean-Sudanese transition area is
covered by granite hills and inselbergs and is divided into three phytogeographical districts: Zou, Bassila and South
Borgou (Adomou et al., 2006).
In Zou phytogeographical District which is located in the south of the Guinean-Sudanese zone and adjacent to the
Guinea-Congolese area, the climax vegetation is of dry deciduous forests type with Hildegardia barteri, Diospyros
mespiliformis Pouteria alnifolia, Anogeissus leiocarpa, and some gallery forests.
The other two phytogeographic districts are essentially marked by the Classified Forests of Oueme, the Classified
Forests of Wari-Maro, the Kouffé Mounts, the mounts of Agoua and Toui-Kilibo of Benin. The granite formations
extend beyond the Guinean-Sudanese transition zone up to the extreme north of Benin (Sudanese zone).
Materials And Methods:-
It is to collect herein data to determine the geographic distribution of hyraxes in southern and central Benin. The
method used was based on investigation through surveys and interviews and walking surveys. The populations
targeted in investigations and interviews are hunters, forest rangers, local residents (including farmers, traditional
healers) and any person resource of both sexes who can provide useful information and recognizing the hyraxes,
their footprints, feces or their cries. Physiognomic planks and well-illustrated documents were enlisted to help the
identification of the species. A questionnaire was administered to a sample of the target population, during group
interviews and individual interviews with resource persons, for extra information. Furthermore, it was also collected
skulls and / or body parts of hyraxes kept by hunters or other resource persons. The field study consisted of two
phases:

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The first phase consisted of an initial investigation into nine administrative departments in the south and center of
Benin (Atlantique, Mono, Couffo, Zou, Collines, Borgou, Donga, Plateau and Oueme valley). Thirty-six cities were
preselected based on ecosystems and habitats they contain, even if hyraxes do not exist or have disappeared
according to the documentation and resource persons. Then, 252 localities, both in villages, suburbs and hamlets
were visited by the authors. 1548 people were interviewed respectively from October 2012, from January to
February 2013, from April to May 2014 and from March to April 2015. This phase was followed by a phase of
surveys, including potential sites of tree hyrax (known or inferred habitats, as well as the village lands or places
mentioned in interviews). Sites (past and present) are georeferenced using GPS. Only sites or actual points of
contact with the taxon (direct observation of the hyrax or recording of its cry) were qualified as current occupation
sites.
The ArcGIS software was used for the treatment of geographic data for the design of the map of current and past
occurrence areas of hyraxes. As exploration habitats is almost complete in the Guineo-Congolese area (southern part
of Benin), the area of the previous range of occurrence and the area of current area were estimated using a digital
electronic planimeter.
To describe the relationship between the knowledge of Daman by people, the kind of hyrax known and their
respective habitats, according to their profession and administrative department, a Multiple Correspondence
Analysis (MCA) was performed. The eight selected variables from the survey sheet for this analysis are presented in
Table 1 with their respective variables. The MCA was performed using the statistical software 'R'.
Results:-
The main survey results are presented in Figures 4, 5, 6 and 7. A fraction of 20.5%, or 318 individuals out of a total
of 1548 people interviewed recognize at least one of the two taxa and their manners. However, this proportion varies
greatly according to the Departments. The Department of Zou was the first (with 24% of relative contribution),
followed by the Cpollines (22%) and Ouémé (14%).
Axis 1 of the plane formed by the axes 1 and 2 opposes clearly the taxon rocks 'Proc' daytime habits ('Midi') to that
tree hyrax 'Dend' often lonely 'soli' with nocturnal habits. This discrimination identifies the Departments associated
exclusively with one of the 2 species or, relatively, to the both. The Departments Zou, Oueme, Plateau, Atlantic and
Couffo are closely associated with tree taxon ('Dend') and are opposed, on axis 1, to the Departments Borgou,
Donga and Collines which were dominates by the rock hyraxes. These three departments are in an intermediate
position between the tree hyrax ('Dend') and the rock hyrax ('Proc'). However, they are nearer to the rock hyrax than
tree hyrax. As for axis 2 which represents the sex and occupation, it combines the female 'F' to the profession of
traditional therapist ('Trad'). This traditional therapist occupation is opposed to the other (farmer and hunter)
dominated by men ('M').
Geographical boundaries map of the distribution of tree hyrax and rock hyrax (Fig. 5) highlights two areas of a large
area of overlap. This sympatric area where both kinds of hyraxes have allotopic live, is entirely within the Sudano -
Guinean phyto-transition area. In the north of this central region as well as in the Sudan region (north - Benin), no
rock hyrax (Procavia capensis) is met.
Tables 2 and 3 indicate the current occupation sites tree hyrax (Dendrohyrax dorsalis subsp sylvestris) respectively
in the Guinean zone (southern part of Benin) and in the Sudano - Guinean transition zone (central regions of Benin).
The 'site' of the Central Nucleus of the classified forest of Lama is designated by the plural ('sites') because it alone
represents several stations or sites (Table 2). As in Table 3, it shows the presence of sites of D. dorsalis subsp
sylvestris (five sites) identified in the central part of Benin which is the Sudano - Guinean transition zone; it is also
the great sympatric area of the two genres of hyrax present in Benin.
In Figure 6, we see a site of rock hyrax (Procavia. Capensis), especially in the area of Kaodji, probably marking the
beginning of its occurrence area which is located indeed at higher latitudes (Figure 7). It may be noted on the map
that past occurrence area of Dendrohyrax dorsalis subsp sylvestris extends along the border with Nigeria.
If we extract from the map in Figure 8 the both sites of tree hyraxes located at the intersection of the Guinean zone
and the Sudano - Guinean (Kaodji and Okpa), it would be only five current sites and two past sites of tree hyrax to
fifteen current sites and ten past sites for rock hyraxes.

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Discussion:-
The percentage of interviewees who recognize at least one of the two taxa by the Department, is even lower than the
species is rare. The Zou had recorded the highest percentage (24%) because the forest of the Lama is the only habitat
of the tree hyrax which is landscaped and well protected in the study area. It confirms that the taxon is still quite
common in this department. This also confirms that the taxon is still quite common in this Department. The sites of
the Central Nucleus of the classified Lama forest actually include ten stations (Djossa et al., 2012). In contrast, in
Atlantic and Couffo, the percentage obtained is less than 7% since the tree hyrax is now rare.
All these five Departments are strongly associated with Dendrohyrax dorsalis subsp sylvestris because they are all
located in the Guinean zone, habitat of this taxon, while habitat of Procavia capensis kerstingi is virtually non-
existent. The sole site of the rock hyrax is located at the northern limit of the Guinean phytogeographic area, at its
intersection with the Sudano-Guinean zone. This is the site Kaodji (7 ° 25 'N). Beyond this first site of Procavia
capensis extends the phytogeographical zone of Sudano - Guinean transition, less humid vast area covered by hills
and rock-dwelling formations. This explains the dominance of colonies of rock hyraxes (presence fifteen sites)
compared to the sites that host the tree hyrax (5 sites). The locality of Kaodji is probably the southern boundary of
this sympatric area which extends until to the latitude of the Penessoulou forest (9 ° 30'N), which host an ancient
tree hyrax site; but only 30% of the sample population interviewed in this central region (Department of Collines
and South of the Departments of Borgou and Donga) recognizes the simultaneous presence of both taxa, while the
rock hyrax is mentioned by 100% of the sampled population. This justifies the offset position of the Departments
Collines, Borgou and Donga, closer to the rock hyrax than the tree hyrax on the factorial map (Figure 5). The
distance between some hyrax sites trees up to 80 km found herein, suggests that past sites or present, would remain
to be discovered in this vast Sudano -guinean area. Most of these inferred or projected sites are located around and
inside of the classified forests of Kouffe Mountains, along the Oueme River and its tributaries. The protected areas
are considered in fact to be the "last refuge" of wildlife in Africa (Tehou and Sinsin 2000). These environments are
still host much of the dense dry forests and gallery forests, habitats of tree hyrax, as outcrops, granite hills and
inselbergs which are habitats of rock hyrax . Thanks to ecotones or habitat diversity (sites Kaodji and Koda), two
different kinds of hyraxes can live in the same environment albeit allotopic way as in the Serengeti park in Tanzania
(Hoeck 1982, 1989). The localities of Kaodji and Koda host each occupation sites of Procavia capensis and
Dendrohyrax dorsalis subsp sylvestris located at 1 to 2 km from each other. By contrast, heterospecific associations
were also mentioned in southern Africa in Heterohyrax brucei (hyrax steppes) and Procavia capensis (Barry and
Mundy, 2002; Deniro and Epstein, 1978).
Area of occurrence of tree hyrax:-
The past and present occurrence areas of the tree hyrax were evaluated in the Guinean zone since sites identification
being there almost exhaustive. The occurrence area of Dendrohyrax dorsalis ssp sylvestris was reduced by half in
three decades from 8007 km² (past occurrence area) to 4003 km² (current occurrence area). This significant loss to
the taxon habitat in southern Guinean zone in Benin is comparable to that recorded for the red-bellied monkey
(Sinsin et al., 2002). However, the taxon distribution area extends beyond the Guinean zone and much of the Sudano
- Guinean transition zone or even South - West of Nigeria. Indeed, the authors had listened to two calls of tree hyrax
near to a forest island preserved in the town of Ijebu - Ode (100 km from the border Benin - Nigeria) and whose
vocalization has proved to be very similar to that of Benin taxon. Considering the well concordant vocalizations
specimens of Ghana and Benin (Akpona et al., 2011), we would be tempted to say that this is the same subspecies
represented from Ghana to Nigeria, pending extensive genetic and molecular studies. Those genetic and molecular
studies could show discrimination or not in the various subpopulations of D. dorsalis subsp sylvestris of West
Africa.
The largest subpopulation of hyraxes trees still probably that of the classified forest of Lama (16250 ha), Central
ecosystem whose core is well preserved for several decades. Most other occupancy sites of tree hyrax correspond to
smaller forest patches, a few hectares. Occupation sites of less than a hectare were observed in isolated hyraxes
groups in Tanzania (Hoeck, 1982). It is not only the classified forests, sacred or community forests usually degraded
and used as last refuges by the taxon but also inaccessible gallery forests, swamps and flood forests. Like the red-
bellied monkey Cercopithecus erythrogaster erythrogaster (Sinsin et al., 2002), the taxon as refuge there probably
because these habitats in the Oueme River Valley are generally difficult to access more than half of the year. These
small colonies of D. dorsalis subsp sylvestris are not spared from a possible local extinction due to their sometimes
irreversible isolation from other metapopulation (development of settlements and agricultural systems ....)

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Conclusion:-
The greatest population recorded in southern Benin and human activities had destroyed and fragmented the habitat
of tree hyrax over the last fifty years. This pressure on this taxon has created isolation of its subpopulations of Benin
from one another but also from those of Nigeria. Nevertheless, this study reveals the wide geographic distribution of
hyraxes in Benin, particularly that of Dendrohyrax dorsalis subsp sylvestris which was omitted from Dahomey -
Gap until the late 20th century by zoologists. Despite the reduction in its area of half of occurrence in the Guinean
zone (preferred habitat), extensive studies on the behavioral ecology, population, habitat, its ethnozoology and
genetic characterization, will better reflect its real status. This is indeed a prerequisite for the development of a
conservation program that may be accompanied by a conservation strategy of Dendrohyrax dorsalis subsp sylvestris
species which is the least prolific species of three kinds of hyraxes.
Figure 1:- phytogeographic Districts of Benin (Source : Neuenschwander et al., 2011)

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Figure 2: Study area and surveyed Communes in the South and Central regions of Benin for hyraxes study.

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Figure 3: Dendrohyrax dorsalis (Fraser, 1855)
Fig. 4:- Relative Contribution of each administrative Department to the survey database on the hyraxes.

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Figure 5: Factorial Map on hyraxe species identification according to their habits, interviewees occupation and their
provenance (department).
-2 -1 012
-1 0 1 2 3
MCA factor map
Dim 1 (19.77%)
Dim 2 (9.70%)
Atl
Borg
Col
Couf
Dong
Oué
Plat
Zou
F
MAd
J
V
Aut
Chas
Cult
Trad
Dend
Proc
Arb
Roch
Trou
Mat
Midi
Soir
Coup
Grp
Soli

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Figure 6: Geographical boundaries of the distribution area (known or inferred, past and present) of hyrax in the
study area. (Years 1961-2011): first phase of the study.

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Figure 7: Past and present occurrence areas of the tree hyrax in the South Benin (Guinean Zone).

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Figure 8: past and present sites of hyrax in the sympatric zone entirely within the Sudano - Guinean (Central
regions of Benin).

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Table 1: Variables used for multivariate analysis, with their respective terms
Variables
Terms
Departments
Atlantic (Atl), Borgou (Borg), Collines (Col), Couffo (Couf), Plateau (Plat)*, Zou
and Oueme (Oue)
Age
Young (J; <25 years) ; Adult (Ad; 25≤ age <60) ; Old (V; ≥ 60 ans)
Sex
Male (M) ; Female (F)
Main activity
Hunter (Chas), Farmer (Cult), Traditionnal therapist (Trad), Other (Aut)
Observation moment
Morning** (At dawn), Noon*** (09 o’clock to 17 o’clock), evening (18 o’clock to
21 o’clock)
Observation localisation
Rock (Roch); Tree (Arb);
Type of observed hyrax
Dendrohyrax (Dend) ; Procavia (Proc)
Life style or habits
Loner (Soli) ; Couple (Coup) ; Group (Grp)
***"Noon" means any period of daytime for characterizing a species with diurnal habits. **"Morning" and
"Evening" indicate herein a nocturnal species, probably of the kind Dendrohyrax (Figure 3).
Table 2: Sites and localities where Dendrohyrax dorsalis ssp still exists and whose presence was confirmed by
authors from 2012 to 2013.
Localities or Sites of current
occupation: 2012-2014
Communes
Ecology: phytogeographical
district
Occupied habitats
Classified Forest of Lama
Zogbodome and
Toffo
‘District of PLATEAU’:
depression of Lama
Dense and semi-deciduous Forest
Tovlamè
Zogbodomè
‘District of PLATEAU’
Border of stream
Gbadagba
Djidja
‘District of ZOU’:
degraded Forest
Border of stream
Cove
Cove
‘District of PLATEAU’:
degraded Forest
Border of stream
Kaodji (7°25
Zagnanado
District of PLATEAU’:
degraded Forest
Border of stream (sympatric area
with P. Capensis)
Forest of Itchede
Adja-Ouere
‘District of PLATEAU’ :
Classified Forest
Border of stream
Classified Forest of
Sakete (Gbamè)
Sakete
‘District of POBE’ Depression
Semi deciduous and
dense Forest
(Akpadanou)
Adjohoun
‘OUEME VALLEY :
Periodically flooded gallery forest
Okpa
Savè
‘District of ZOU : gallery forest
Periodically flooded gallery forest
(Oueme stream border)
Niaouli
Allada
District of PLATEAU
Semi deciduous and
dense Forest
Zè-Aokpa
Zè
District of PLATEAU
Degraded Forest
Table 3: Sites and localities where Dendrohyrax dorsalis subsp sylvestris still exists (in Sudano-Guinean zone) and
confirmed by the authors: 2013-2015.
Localities or Sites of current
occupation: 2012-2014
Communes
Ecology: phytogeographical district
Occupied habitats
Okounfo
Save
Soudano-Guinean transition Zone (District
of Zou)
Border of Oueme stream
Koda
Tchaourou
Soudano-Guinean transition Zone District
of Zou (South Borgou)
Border of stream
Baanon
Bantè
Soudano-Guinean transition Zone
District of Zou (South Borgou)
Border of stream Adjiro
Ouanou
Bassila
Soudano-Guinean transition Zone
District of Zou (South Borgou)
Degraded Forest
Sites of Kouffe Mountain
Forest
Bassila
Soudano-Guinean transition Zone
District of Zou (South Borgou)
Degraded Forest

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