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Rodents of the MenageshaState Forest, Ethiopia, withan emphasis on the endemicPraomys albipes Rüppell 1842Afework Bekele aa Department of Biology , Addis Ababa University , P.O. Box 1176, Addis Ababa , EthiopiaPublished online: 01 Aug 2012.

To cite this article: Afework Bekele (1996) Rodents of the Menagesha State Forest,Ethiopia, with an emphasis on the endemic Praomys albipes Rüppell 1842, TropicalZoology, 9:1, 201-212, DOI: 10.1080/03946975.1996.10539308

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Tropical Zoolog;y 9: 201-212, 1996

Rodents of the Menagesha State Forest, Ethiopia, with an emphasis on the endemic Praomys albipes Riippell 1842

AFEWORK BEKELE

Department of Biology, Addis Ababa University, P.O. Box 1176, Addis Ababa, Ethiopia

Received 26 August 1994, accepted 28 January 1995

A shrew and 12 species of rodents were recorded from the Menagesha State Forest, Ethiopia, from November 1991 to July 1993. A total of 293 specimens were trapped in 3209 trap nights. Trap success varied from 4.7 to 26.9%. Praomys albipes Riippell 1842 was the most ubiquitous, comprising 54.3% of the total rodents captured in all habitats and 70.8% of the rodents captured in the forest. Species number and richness was low in disturbed habitats. Composition and distribution of the rodent fauna were compared with studies in other localities of Ethiopia. The effect of habitat degradation on the distribution of species and its concern was stressed.

KEY WORDS: rodents, Menagesha State Forest, endemic, snap-trap, Ethiopia, Praomys albipes.

Introduction The study area Materials and methods Results Discussion Acknowledgements References

INTRODUCTION

201 202 204 205 208 211 211

The diverse fauna of Ethiopia is associated with the varied topographical features of the country. The rodent fauna of Ethiopia consists of 70 species, of which 15 are endemic (YALDEN & LARGEN 1992, HILLMAN 1993). Rodents comprise 25% of the Ethiopian mammal fauna and contribute about 50% of the total endemic species. Out of the total 70 species of rodents grouped under nine families, the family Muridae alone comprises 81% of the rodent species and 93% of the endemics. One of these murids is the endemic Praomys albipes, misappropriately also known as the white-footed rat, since the species possesses a dark patch on the upper metatarsal region. There are still uncertainities concerning the systematics of the Praomys-

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202 Afework Bekele

Myomys complex. This study follows the nomenclature adopted by Y ALDEN et al. (1976), RuPP (1980), YALDEN & LARGEN (1992).

Of all the species in the genus Praomys Thamas 1915 in Ethiopia, P. albipes is the most widespread and abundant. The species occurs widely in the forest and bush between altitudes of 1500-3300 m a.s.l. It has also been recorded at 1285 m (DoRST 1972) and 820 m a.s.l. in the humid densely forested region of the southwest (YALDEN et al. 1976). INGERSOL (1968) has collected the species from the southeast part of the country at altitudes of 2000 and 2700 m a.s.l.

Although extensive investigations have been conducted on the ecology of closely related species in different regions of Africa, practically no work has been conducted on the ecology of this widespread and distinct species within Ethiopia. The few ecological studies on Ethiopian rodents include MuLLER's (1977) account on popula­tion ecology of Arvicanthis abyssinicus Riippell 1842 in the highlands and RuPP's (1980) work on the rodents of Ethiopia. More recently, YALDEN (1988a, 1988b) surveyed the high alpine moorland and dense forest in the southern part of the country by examining the species composition, distribution and altitudinal zonation of small mammals. Despite the extensive plateaux comprising about 50% of the African highground above 2000 m (YALDEN 1983), very little is known of the ecology of the mountain fauna of Ethiopia.

The present study site is a remnant forest block in the central part of the country known as the Menagesha State Forest. The area is part of the central plateau covering an altitudinal range from 2200-3385 m a.s.l. This natural forest covers about 2500 ha and is described by FRIIS (1992) as one of the undifferentiated Afro-montane forests comprising ]uniperus-Podocarpus forest with an element of broad-leaved species. The floral composition of this forest was surveyed by BREITENBACH (1962), BREITENBACH & KOUKAL (1962), GILBERT (1970), HENRY (1973) and SEBSIBE DEMISSEW (1980, 1988). The conservation practice and history of the Menagesha State Forest date at least as far back as 5 centuries. This forest has been protected by Imperial edict since 1600. In 1981, it was accepted as one of the 13 national forest priority areas to ensure optimum protection of the natural forest and the planting of both indigenous and exotic conifers on degraded areas.

Using data obtained by snap-trapping surveys from November 1991 to July 1993, the present study focuses on the distribution, abundance and species composition of the rodent fauna of the Menagesha State Forest, with an emphasis on P. albipes.

THE STUDY AREA

The study area is an isolated forest block located in central Ethiopia, about 43 km southwest of the capital, Addis Ababa, between longitudes 38°30' to 38°39' East and latitudes 08°55' to 09°04' North (Fig. 1). The area is part of the volcanic dome with an elevation extending from 2200-3385 m a.s.l. It experiences 7 months of rain from March to September with the highest concentration in July and August (DANIEL GAMACHU 1977). The mean annual rainfall is about 1150 mm. The mean annual temperature varies from 11-22 °C with lower temperatures at higher altitudes.

Juniperus procera Endlicher 1847, Podocarpus fakatus Mirb 1825 and Olea euro­paea subsp. cuspidata Ciffieri 1942 comprise the main vegetation zone while Erica

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Rodents of the Menagesha State Forest 203

~-..... :.·.··:: .. ·. ·:::::::·

... ·.

W· •• v

Fig. 1. - Location of snap-trapping sites at the Menagesha State Forest.

arborea Linnaeus 1753 and Helichrysum odoratissimum Lessing 1832 with mountain grassland dominate the upper slopes. The Menagesha State Forest covers an area of 9248 ha, but the natural forest proper is estimated to be only 2500 ha (ANONYMOUS 1991). Attempts to reafforest the degraded area have resulted in a 1000 ha plantation of the introduced Cupressus lusitanica Miller 1768. The area is still being encroached by the surrounding farmers for timber, fire-wood, grazing and farming. The area is also highly valued as water catchments and a centre of attraction for tourists and schools. In addition, the area harbours about 108 species of birds including seven endemics.

The study area covers different vegetation zones at different altitudes from 2200-3385 m a.s.l. A total of 17 sites in the forest proper and six sites at the periphery of the forest were surveyed (Fig. 1). Some of these sites were trapped frequently, at each trapping occasion, while others were abandoned after an initial trial. Brief comments on the vegetation together with altitudes of the localities (Fig. 1) are described below.

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A: Suba forest station, at 2440 m a.s.l. with grassy patches, Echinops steudneri Hoffm 1898, Solanecio gigas Vatke 1875, Maytenus gracilipes Exell 1952, C. lusitanica, 0. europaea, P. falcatus, Pinus radiata Don 1836, and]. procera.

B: At 2500 m a.s.l., deep gorge with more open grassy patches as well as S. gigas, Kalanchoe densiflora Rolfe 1919 and]. procera.

C: At 2510 m a.s.l. with Rubus apetalus Poir 1804, S. gigas, Senecio ochrocarpus Oliver & Hiern 1877, Myrsine a/ricana Linnaeus 1753, Acacia mearnsii De Wild 1925, P. /akatus and]. procera.

D: At 2540 m a.s.l. with more open grassy patches and]. procera. E: At 2580 m a.s.l. with well covered herb and grassy patches, Dovyalis abyssinica Warburg

1893, Dovyalis vericosa Warburg 1893, R. apetalus, M. gracilipes, M. africana, P. fakatus, 0. europaea and J. procera.

F: At 2740 m a.s.l. with E. arborea forest, M. a/ricana and stunted]. procera. G: At 2860 m a.s.l. near the hotel and tukuls with new plantations of C. lusitanica and Pinus

patula Schlechter 1838, Rosa abyssinica Lindley 1820 and E. steudneri. This locality is very bare because of intensive grazing and farming.

H: Adere mountain at 3010 m a.s.l. with grassy patches, E. arborea and H. odoratissimum with stunted J. procera.

I: Mogle mountain, the highest peak at 3385 m a.s.l. E. arborea and H. odoratissimum comprise the dominant vegetation. Some very stunted]. procera also occur.

J: At 2470 m a.s.l. with dense grass and herb patches, 0. europaea, M. gracilipes and R. apetalus.

K: At 2350 m a.s.l. with Carissa edulis Vahl1790,]. procera, P. fakatus and 0. europaea. L: At 2330 m a.s.l. with]. procera. It also includes an abandoned quarry with elephant grass. M: At 2300 m a.s.l. with C. edulis, Croton macrostachys Richard 1851, Acacia decurrens Willd

1806 and a]. procera plantation. N: At 2330 m a.s.l. with M. gracilipes, 0. europaea and]. procera. 0: At 2630 m a.s.l. with stunted]. procera, grassy patches and dominant E. arborea. P: At 2500 m a.s.l. with C. edulis,]. procera, Eucalyptus plantation and crevices. Q: At 2780 m a.s.l. on the Menagesha mountain with 0. europaea,]. procera, P. fakatus and

stony crevices. R: At 2200 m a.s.l. with young C. lusitanica plantations, C. macrostachys, C. edulis, Solanum

incanum Linnaeus 1753, Lippia adoensis Walp 1844, Measa lanceolata Forsskai 1775 and Pennisetum trachyphyllum Pilger 1901.

S: At 2240 m a.s.l. with Eucalyptus globulus Labill1799, Acacia ahyssinica Benth 1846, Milletia ferruginea Baker 1871, few C. edulis and a new plantation of C. lusitanica.

T: At 2280 m a.s.l. in Suba village and the sawmill compound near the hay meadow. U: At 2280 m a.s.l. in dense bush with C. edulis, C. macrostachys and young C. lusitanica

plantation. V: At 2250 m a.s.l. around and inside wheat fields. W: At 2200 m a.s.l. inside the village tukuls.

MATERIALS AND METHODS

At each trap site, Victor mouse traps and the smaller Museum special traps were used in lines of 10-30 traps alternately at 10 m intervals. A minimum of 30 trap nights were used at each trap site. Some mini-Victor mouse traps were also used occasionally for possible trapping of shrews in addition to locally made pitfall traps. Peanut butter was used as bait. Prebaiting and removal were carried out in the early morning and late afternoon. A total of 3209 trap nights were used during the survey (excluding the sprung or stolen snap-traps).

The captured specimens were weighed using a Pesola spring balance, measured and sexed. Females were examined for pregnancy and lactation and males for testes size and location. Prepared skins and skulls were deposited in the Zoological Natural History Museum, Biology Department, Addis Ababa University.

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Rodents of the Menagesha State Forest 205

RESULTS

A total of 293 specimens were captured in 3209 trap nights (Table 1). Trap success at different sites and occasions varied from 4. 7 to 26.9% with the mean 9.1 %. From the snap-trap surveys, the following nine species of rodents were recorded: Praomys albipes, Desmomys harringtoni Thomas 1903, Lophuromys flavo­punctatus Thomas 1888, Mus mahomet Rhoads 1896, Arvicanthis dembeensis Riippell 1842, Rattus rattus Linnaeus 1758, Otomys typus Heuglin 1877, Stenocephalemys griseicauda Petter 1972 and Dendromus mystacalis Heuglin 1863. A single female Graphiurus murinus Desmarest 1822 was also live-trapped at 2500 m a.s.l. in the open habitat of the forest, under a Juniperus tree. In addition to the above species, the mole-hills and tunnels of the underground dwelling Tachyoryctes splendens Riippell 1836 were very common in more open habitats from 2200-3000 m a.s.l. The quills of porcupine, Hystrix cristata Linnaeus 1758 were also observed dispersed in open habitats within the forest. A young shrew, Crocidura cf. fuscomurina Heuglin 1865 was caught by hand near the edge of the house in the forest station at 2440 m a.s.l., but no shrews were caught in any of the traps.

P. albipes was the only species available at each trapping occasion and comprised 159 (54.3%) of the total 293 rodents collected. Males comprised 83 (52.2%) and females 76 (47.8%). The difference in the rate of capture of males and females was not significant (X.~= 0.30, NS). D. harringtoni comprised 45 (15.4%), L. flavopunctatus

Table 1. Species composition, trap nights and percent trap success of rodents caught by snap-trapping surveys from

different habitats within the Menagesha State Forest.

Species Trap %trap Month

Pa Dh Lf Mm Ad Rr Ot Sg Dm Total nights success

1991 Nov 18 4 3 5 31 405 7.7 Dec 7 5 12 124 9.7

1992 Mar 14 3 4 21 199 10.6 May 5 1 3 3 12 60 20.0 Jul 30 2 4 2 39 145 26.9 Sep 21 21 260 8.1 Nov 17 2 4 2 25 290 8.6

1993 Jan 7 4 3 3 17 248 6.9 Mar 9 18 3 3 4 38 320 11.9 May 21 11 5 6 44 458 9.6 Jul 10 6 2 14 33 700 4.7

Total 159 45 33 32 10 8 3 2 1 293 3209 9.1

Pa = P. albipes, Dh = D. harringtoni, Lf = L. /lavopunctatus, Mm = M. mahomet, Ad = A. dembeensis, Rr = R. rattus, Ot = 0. typus, Sg = S. griseicauda, Dm = D. mystacalis.

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206 Afework Bekele

33 (11.2%) and M. mahomet 32 (10.9%) of the total catch. The remaining five species of rodents comprised 8.2% of the sample. The distribution of P. albipes in Ethiopia from different literature surveys shown in Fig. 2 confirms the high association between the distribution of the species and an elevation above 1500 m a.s.l.

The vegetation zones together with elevational distribution of rodents along the slopes of the forest are given in Fig. 3. Although P. albipes occurred widely in both bushy vegetation and in the forest, it was less common in Erica scrub and open grassy patches. In the Menagesha State Forest, it covered an altitudinal range from 2200-3300 m a.s.l., with frequent occurrences up to 2600 m a.s.l. in the Juniperus zone (Table 2); 61.0% of the total 159 P. albipes were captured in the forest habitat, 25.8% in the bush and only 13.2% in the Erica scrub (Table 3). P. albipes comprised 97 (70.8%) of the total137 rodents captured in the forest habitat while in the bush and in the Erica scrub, it comprised 41 (50.0%) and 21 (31.8%) respectively. R. rattus, A. dembeensis and D. harringtoni could have been trapped more often had their habitats not been frequently avoided. R. rattus was plentiful in areas of human habitations. A. dembeensis was very common in grassland, farmland and nearby scrub at 2200 m a.s.l. below the forest. D. harringtoni occurred in restricted numbers in nearby bush and in open habitats in the forest where Kalanchoe vegetation dominates, but more commonly in Erica scrub at higher altitudes where stony crevices occur. L.

16p-_. __________ _. __________ -M __________ _. ____ ~

12

Fig. 2. - Map of Ethiopia showing the recorded distribution of P. a/bipes (solid circles) and elevation above 1500 m a.s.l. (stippled).

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Rodents of the Menagesha State Forest 207

"' ·- "' "' Q) c :::l :::l a. 0 a. - "' ;., .c Cll

3400 - c (.)

"' "' ·- c:: 0 c a_ ~

:::l Q)

"' a.

"0

"' 0

c:: > 3200 CD a_ "'

a. "'

....... Eric a "' _J :::l

1-

E 3000 "' "' - "' w 0 2800 :::> ---------

Q) 0: "' E

"' 0 :::l (.)

"' c I "' "'

t- Juniperus-Erica :::l

E "'

t- 2600 ....1 <(

2400 Juniperus-Podocarpus-Oiea

E :2

(.!)

• c CD CD -.c "' E (.)

"' CD -"0 "' ;., <{ E

2200 Scrub 0

Fig. 3. - Elevational distribution of rodents along the vegetation zones in the Menagesha State Forest.

flavopunctatus was more widespread in bushy vegetation and open forests with grassy patches. M. mahomet was widespread in bushy vegetation and frequented open forests with grassy patches up to 2580 m a.s.l. 0. typus occurred in open forests with grassy patches and in Erica scrub at higher altitudes. S. griseicauda was captured only in Erica scrub above 3000 m a.s.l. The single pregnant female D. mystacalis and G. murinus

Table 2. The distribution of rodents from the snap-trap survey in the Menagesha State Forest across the vegetation

zones.

Species Catch Altitudinal

Habitat-type range (m)

P. albipes 159 2200-3300 Bush, forest, Erica D. harringtoni 45 2200-3385 Bush, open forest, Erica L. flavopunctatus 33 2200-3300 Bush, open forest, Erica M. mahomet 32 2200-2580 Bush, open forest A. dembeensis 10 2200-2300 Bush, grass, farm R. rattus 8 2200-2860 Human habitations 0. typus 3 2500-3300 Open forest, Erica S. griseicauda 2 3010-3385 Erica D. mystacalis 1 2200 Bush T. splendens * 2200-3010 Open forest, meadow

The asterisk shows the presence of the species from soil pile sightings, but no trap was set to trap the animal.

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208 Afework Bekele

Table 3. The distribution of rodents from snap-trap surveys in each habitat-type, altitude (m a.s.l.) and percent

trap success.

Species

P. albipes D. harringtoni L. jlavopunctatus M. mahomet A. dembeensis 0. typus S. griseicauda D. mystacalis

Total Trap-nights % trap success

Bush <2300

41 3

14 13 10

1

82 868 9.4

Habitat/altitude forest

2300-2800

97 8

11 19

2

137 1712

8.0

Erica Total >2800 catch

21 159 34 45 8 33

32 10

1 3 2 2

1

66 285 604 3184 10.9 9.0

(each with four embryos) were captured in the bush outside the forest proper at 2200 m a.s.l. and in the open forest at 2500 m a.s.l. respectively.

Young J. procera and C. lusitanica plantations supported fewer species and individuals because their dense shade resulted in bare ground, no cover and no berries. At higher altitudes with Erica and stunted ]. procera habitat disturbed by human interference, only a single P. albipes was trapped out of 61 trap nights (1.6%). However, at the highest altitudes with Erica scrub, where human interference is limited, more individuals and species were trapped.

All of the captured P. albipes were from the early morning trap-checks. Thus, P. albipes is exclusively nocturnal. Some individuals of L. flavopunctatus, M. mahomet and D. harringtoni were collected during the late afternoon check. A. dembeensis was exclusively diurnal. The rest of the rodents were collected from the early morning checks.

DISCUSSION

In the present study, the recorded composition of the rodent fauna of the Menagesha State Forest was 12 species and one shrew. Similar studies in Ethiopia by YALDEN (1988b) and HuTTERER & YALDEN (1990) recorded seven species of rodents and four shrews while RuPP (1980) obtained 10 species of rodents and eight shrews (see also HuTTERER 1981). The absence of shrews in the present snap-trap survey was surprising. Variable species richness in different vegetation zones has been revealed by DELANY (1964, 1966) for East Africa, BoND et al. (1980) for South Africa, RAPPOLD & RAPPOLD (1989) for Malawi, RAPPOLD (1974) for Nigeria and DIETERLEN (1967, 1986, 1990) for East and Central Africa. The low species richness in the Menagesha State Forest is comparable to similar studies conducted in other regions of Ethiopia. This view complements the suggestion put forward by DIAMOND & HAMIL-

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Rodents of the Menagesha State Forest 209

TON (1980) concerning the poor forest fauna of Ethiopia, as a result of reduction in the size of forests during the cold and dry periods of the Pleistocene. In addition, some of the species might have been affected due to habitat manipulation by humans prior to the introduction of conservation practices 5 centuries ago and the continua­tion of this encroachment at the present time as well.

On comparing the distribution of rodents along the gradients of vegetation in the Menagesha State Forest the pattern was similar to that observed in related studies in Ethiopia. As part of the central plateau, the lowest altitude of the Menagesha State Forest is 2200 m a.s.l. The range of altitudinal distribution of P. albipes (2200-3300 m a.s.l.) falls within those described by YALDEN (1988b) and RuPP (1980). But, in addition to forest sites, P. albipes also frequents scrub or bushy vegetation outside the forest proper. The scrub area dominated by C. edulis is the preferred habitat outside the forest. D. harringtoni was not collected by YALDEN (1988b) in southern forests of Ethiopia, but RuPP (1980) collected the species from various localities in western Ethiopia from 1800-3200 m a.s.l. In the present study, D. harrittgtoni comprised the largest collection in the Erica scrub above 3000 m a.s.l. Stony crevices at these altitudes were the preferred habitats. L. flavopunctatus was widely distributed in scrub and open habitats in the forest, but in the present study, it was less abundant than in other studies in Ethiopia. The distribution of M. mahomet in grassy patches, bush and open habitats was similar to that found in the previous study by YALDEN (1988a, 1988b), but in the Menagesha State Forest, this species was restricted to areas below 2580 m a.s.l. A. dembeensis was unusually common at 2200 m a.s.l. near grassland and farmland. R. rattus was ubiquitous in areas of human habitations. RuPP (1980) recorded trapping this species in the forest as well. In the present study, only one animal was captured in the garden of the forest station. The record of 0. typus was within the altitudinal range of Rupp' s collection. S. griseicauda was collected in the Erica scrub above 3000 m a.s.l. coinciding with RuPP's trapping. The altitudinal range reported by YALDEN (1988a) was 2400-2920 m a.s.l. It is difficult to state the distribution of D. mystacalis and G. murinus because only a single pregnant female was collected in C. edulis-dominant shrub at 2200 m a.s.l. and open habitat in the forest proper (2500 m) under the Juniperus tree.

P. albipes was found to be ubiquitous in the Menagesha State Forest and comprised 54.3% of the total record. YALDEN (1988a), in southern Ethiopia, also found this species to be the largest contributor (33%), and P. albipes comprised 25~ of RuPP's (1980) collection. In both the collections of YALDEN (1988a) and RUPP (1980), L. flavopunctatus was the second most abundant species. By contrast, in the present study, the second highest component was D. harringtoni, followed by L. flavopunctatus and M. mahomet. Y ALDEN & LARGEN ( 1992) stress that L. flavopunctatus is the most abundant and widespread of the Ethiopian rodents. Although the species was widely distributed in both scrub and forest cover, in the Menagesha State Forest, the number of L. flavopunctatus was limited.

The mean trap success of 9.1% obtained in this study was very low compared to similar studies in Ethiopia. YALDEN (1988a, 1988b) in the Harenna Forest, southern Ethiopia, recorded a mean success rate of 18.7% and a trap success of 38.9% in Erica scrub. RuPP (1980) obtained a 35% trap success. In the present study, the highest trap success recorded was 26.9%. The 10.9% trap success obtained in the Erica scrub includes sites with low and high human interference. The lowest trap success (1.6%) was observed in the Erica forest between altitudinal ranges of 2800-2960 m a.s.l., near

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human habitation. These sites were intensively grazed by livestock, resulting in less ground cover. However, the Erica scrub at 3000-3385 m a.s.l., where human interfer­ence was limited, yielded better trap success (20.3%). YALDEN (1988b) recorded similar low trapping success in the Erica forest diturbed by human habitation and highest trap success in the undisturbed Erica scrub above the disturbed zone. HAPPOLD (1978) emphasized the unusually low number of Praomys tullbergi Thomas 1894 in Nigerian forest, disturbed by human interference.

The distribution of P. albipes in Ethiopia shown in Fig. 2 was modified from YALDEN et al. (1976). However, since most of the regions in Ethiopia have not yet been properly explored, further investigation is likely to reveal more information and extend the range of the species. The localities and altitudinal ranges covered in the present study are limited compared to similar studies in other regions by RuPP (1980) and YALDEN (1988b), but the composition of the rodent fauna shows some similarities and differences. Even though the Erica scrub vegetation zones and Juniperus forests in southern parts of the country show similarities to the present study site, most localities are characterised by different vegetation types. The expected forest rodents such as Dendromus melanotis Smith 1834, Oenomys hypoxanthus Pucheran 1855 and Dasymys incomtus Sundevall1847, recorded by RuPP (1980) from western part of the Rift Valley, are lacking in the Menagesha State Forest. The non-trapping of the common high plateaux grass rat is also surprising. The absence of the endemic Dendromus lovati De Winton 1899 is also odd because old records show its occurrence in the Menagesha area, whereas the absence of the rare endemic mouse, Muriculus imberbis Riippell 1842, is probably the result of habitat degradation. YALDEN's (1988b) record shows the absence of D. harringtoni. At the same time, his collection included the high moorland endemic rodents Arvicanthis blicki Frick 1914, Stenoce­phalemys albocaudata Frick 1914, Lophuromys melanonyx Petter 1972 and Tachyo­ryctes macrocephalus Riippell 1842. The absence of these endemic rodents from the present study site can be explained by their extreme moorland habitat and restricted localised distribution.

P. albipes also occurs in scrub vegetation zones 40 km east and 80 km west of the present study site. These localities are isolated from the Menagesha State Forest by extensive human settlement and degraded areas as a result of farming and grazing. No doubt that these localities were once connected to each other by either woodland or forest vegetation. Although the species is still widely distributed in the forest and bush, its distribution is shrinking due to habitat degradation. According to the estimate of ETHIOPIAN MAPPING AUTHORITY (1988), the continuous forest on Ethio­pian highlands used to cover 37% of the total area. At present, less than 3% of the country is covered with forest. Deforestation in Ethiopia was recognised as a serious issue a century ago (PANKHURST 1989). LARGEN & RAsMUSSEN (1993) stress the deleterious effect on the abundance and distribution of the flora and fauna of Ethiopia of the rapid increase in human population and its associated destruction of the natural habitat in order to provide fuel, timber, and cleared land for agriculture or grazing. Unlike large mammals, poaching is not the cause of population shrinkage and extermination for small mammals, but habitat degradation. Although the relatively small home range sizes of small mammals seem advantageous compared to those of large mammals, the long-term continuous degradation of the habitat due to human pressure can have adverse effects. New management strategies in maintaining and improving this man-induced environment by integrating local people as advocated by

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Rodents of the Menagesha State Forest 211

MIEHE & M.:rEHE (1994) should be implemented by the authorities. To minimize these problems, conservation of forest habitats on a sustainable basis should be given priority and maintaining the biodiversity of this historical site should be the order of the day.

P. albipes is not only confined in the Menagesha State Forest. Along with the other rodents, it is also widely distributed in different localities and habitats of Ethiopia. The pattern of rainfall varies from region to region, so does the vegetation. The composition of the rodent fauna is also variable. In order to arrive at a comprehensive understanding of the population dynamics and distribution patterns of many of these species, further studies should be conducted in different habitats of Ethiopia in addition to a long-term and continuous investigation in the same locality.

ACKNOWLEDGEMENTS

I am extremely grateful to Dr D.W. Yalden for his valuable advice and comment on an earlier draft of the manuscript. I wish to thank Dr J.C. Hillman, the staff members of the Forestry Department and all members of the Ethiopian Flora Project for the identification of plants. I would also like to convey my appreciation to Goitom Redda and Eshetu Assefa for their support in the field work. Financial support for this study was provided by Wildlife Conservation Society -International, ORS Award, The British Council, Universities of Manchester and Addis Ababa.

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