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Mammalia 72 (2008): 116–122 2008 by Walter de Gruyter • Berlin • New York. DOI 10.1515/MAMM.2008.015

Article in press - uncorrected proof

2007/15

A survey of the small mammals of Minziro Forest, Tanzania,

with several additions to the known fauna of the country

William T. Stanley 1,* and Charles A.H. Foley 2

1 Division of Mammals, Field Museum of Natural

History, 1400 South Lake Shore Drive, Chicago,

IL 60605, USA, e-mail: [email protected] Wildlife Conservation Society, P.O. Box 2703,

Arusha, Tanzania

*Corresponding author

Abstract

Complete faunal lists are vital for understanding the nat-

ural history of countries and unique habitats within them,

as well as the design of effective conservation strategies.

Although many mammal species of Tanzania are familiar,

the fauna of some areas of the country has not been

properly documented. We surveyed the small mammals

(shrews, bats and rodents) of Minziro Forest in north-

western Tanzania. Six species are documented here for

the first time for Tanzania. Other records include Tanza-

nian species poorly represented as vouchers in collec-

tions. Minziro is a unique forest worthy of conservation

priority and further study.

Keywords: bats; Minziro; rodents; shrews; Tanzania.

Introduction

Although the larger mammals of Tanzania are reasonably

well documented, the smaller fauna is relatively poorly

known, particularly in forested habitats. Recent surveys

in montane areas have augmented our understanding of

the shrews and rodents of these habitats (Stanley et al.

1998, Stanley and Hutterer 2000, 2007, Carleton and

Stanley 2005, Stanley and Olson 2005), but the lowlandforests of Tanzania remain poorly known. One such forest

is the Minziro Forest Reserve, situated in the northwest-

ern corner of Tanzania (Figure 1). Contiguous with Mala-

bigambo Forest Reserve in Uganda, Minziro is a mosaic

of groundwater-forest and grassland habitats, which is

unique within Tanzania, and is also one of very few exam-

ples of Guinea-Congo forest type within the country.

Although recent surveys have focused on the mammals

of forests in southern Uganda, such as Sango Bay and

Kalinzu Forest (Dickinson and Kityo 1996, Lunde and

Sarmiento 2002) and the birds in Minziro Forest (Baker

and Hirslund 1987), the mammalian fauna has not been

documented. A survey of the small mammals (shrews,bats, and small to medium-sized rodents) was conducted

in August 2006 to assess the small mammal fauna.

Materials and methods

Study area

Minziro Forest Reserve is in Bukoba Region, Minziro Dis-

trict, and is bordered by the Tanzania-Uganda border to

the north and the Kagera River along much of its eastern

boundary (Figure 1). The reserve covers approximately

29,000 ha, and ranges in elevation from 1140 to 1180 m.

It rings the village of Minziro, which is above the reserve

at 1330 m. Much of the reserve is Baikiaea-Podocarpus

seasonal swamp forest (Davenport and Howard 1996),

with historically two rainy seasons: March–May and

October–November. Seasonal flooding of the river is an

important aspect of the Minziro ecosystem. Small-scale

logging has occurred in the area (Baker and Hirslund

1987), as well as burning of the grassland (observed

during this study).

This study, carried out between 16 and 25 August

2006, focused on three microhabitats in the southern half

of the reserve, including seasonal swamp forest

(1.087498 S, 31.535538 E, 1150 m; hereafter referred to

as the forest site), grassland (1.084468 S, 31.524158 E,

1149 m; grassland site), and seasonal swamp forest

bordering a permanent pond (1.094188 S, 31.515388 E,

1161 m; pond site).

Trapping methodology

Shrews and rodents were sampled using two different

trapping techniques: pitfall lines and trap lines. Six pitfall

lines were installed, each consisting of 11 15-l buckets,

26 cm deep, buried in the ground so that the top of each

bucket was flush with the ground. The buckets were

spaced 5 m apart and had a 50 cm high plastic fence

installed in a horizontal position that passed over the

center of each bucket. Further details are given in Stanley

et al. (1996). Trap lines utilized three different types of

traps: Museum Specials (14 cm=7 cm), Victor Rat Traps

(17.5 cm=8.5 cm) and medium-sized Sherman Traps(23 cm=9.5 cm=8 cm). The size of these different traps

does not allow for the capture of larger rodents such as

squirrels and Cricetomys. For this reason, we restrict our

analysis and discussion of the trapping effort to smaller

rodents of-200 g. The bait used in the traps was freshly

fried coconut coated in peanut butter, and traps were

re-baited every afternoon. Trap and pitfall lines were

checked twice each day (early morning and afternoon)

and specimens were handled in accordance with the

American Society of Mammalogists guidelines (Animal

Care and Use Committee 1998). The terms ‘‘trap night’’

and ‘‘bucket night’’ refer to one trap or bucket in opera-

tion for a 24-h period (07:00–07:00 h).Bats were sampled with a 12 m mist net, which was

set at one end of the water body at the pond site. The

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W.T. Stanley and C.A.H. Foley: Small mammals of Minziro Forest, Tanzania 117


Figure 1 Map of Minziro Forest, and its location within Tanzania.

net was opened at 18:00 h and monitored for 2 h eachnight for four consecutive nights starting on 20 August.

Other bats were sampled opportunistically from buildings

within Minziro village and nearby caves. A ‘‘net hour’’ is

defined as a 12 m net left open for 1 h.

Because not all traps and buckets were employed for

equal amounts of time (some trap lines were set on the

first day of the survey, whereas others were installed on

the second or third day), we quantify the sampling effort

at each site. We refer to the success rate of each method

as either ‘‘trap success’’ or ‘‘bucket success’’, and cal-

culated these values by dividing the number of individ-

uals captured by the number of trap or bucket nights and

multiplying by 100. When discussing the two trappingmethodologies combined, we use the term ‘‘sampling

night’’ to refer to either one trap night or one bucket

night. We use ‘‘capture success’’ to refer to the successrate for the two methodologies combined, and calculated

this value by dividing the number of individuals captured

by the number of sampling nights and multiplying by 100.

Specimen preparations included skins and skeletons

and formalin-fixed cadavers. The standard body meas-

urements taken include total length (TL), length of head

and body (HB), length of tail vertebrae (TV), length of hind

foot (HF), length of ear (EAR), length of forearm (FA) for

bats, and weight (WT) (DeBlase and Martin 1974). All

measurements were recorded in millimeters except

weight, which was in grams. Tissues including heart, liv-

er, and kidney were preserved in EDTA buffer. All speci-

mens are deposited in the Field Museum of NaturalHistory (FMNH; catalogue numbers are listed in Appen-

dix), and a portion will be returned to the Department of






118 W.T. Stanley and C.A.H. Foley: Small mammals of Minziro Forest, Tanzania


Table 1 Distribution of shrews and rodents in three different microhabitats in Minziro Forest Reserve in August 2006.

Forest Grassland Pond Totals

Crocidura hildegardeae Thomas 1904 0 1 0 1

Crocidura nanilla Thomas 1909 0 1 0 1

Crocidura nigrofusca Matschie 1895 0 2 2 4

Crocidura olivieri (Lesson 1827) 0 0 1 1

Suncus megalura (Jentink 1888) 0 5 0 5

Sylvisorex johnstoni (Dobson 1888) 9 0 0 9

Lophuromys aquilus (True 1892) 0 1 6 7

Lophuromys sikapusi (Temminck 1853) 0 4 0 4 Deomys ferrugineus Thomas 1888 0 0 4 4

Dendromus mystacalis Heuglin 1873 0 1 0 1

Dasymys incomtus (Sundevall 1847) 0 7 0 7

Grammomys macmillani (Wroughton 1907) 0 0 6 6

Hybomys univittatus (Peters 1876) 8 0 12 20

Hylomyscus stella (Thomas 1911) 56 1 31 88

Mus musculoides Temminck 1853 0 1 0 1

Mus triton (Thomas 1909) 0 1 0 1

Oenomys hypoxanthus (Pucheran 1855) 0 0 1 0

Praomys jacksoni (de Winton 1897) 35 0 38 73

Graphiurus murinus (Desmarest 1822) 0 0 1 1

Total number of individuals 100 32 102 234

Total number of species 3 10 10 15

Total number of sampling nights 664 328 1063 2055

Trap success (%) 15.1 9.7 9.6 11.4

Only specimens caught in traps or buckets are included in totals. Species that were recorded for the first time in Tanzania are in

bold font.

Zoology, University of Dar es Salaam. We follow the tax-

onomy of Hutterer (2005), Musser and Carleton (2005),

Schlitter (2005), and Simmons (2005).

Results

During the survey, a total of 2055 sampling nights were

accrued, including 550 pitfall nights and 1505 trap nights.

By sites, 1063 sampling nights (198 pitfall nights, 865

trap nights) were accrued at the pond site, 328 sampling

nights (88 pitfall nights, 240 trap nights) at the grassland

site, and 664 sampling nights (264 pitfall nights, 400 trap

nights) at the forest site.

Nineteen shrews representing six species were cap-

tured in 550 pitfall nights for an overall bucket success

of 3.4%. By site the bucket success was 1.0% for the

pond site, 9.1% for the grassland site, and 3.4% for the

dry forest site. For the 1505 trap nights, 213 mammals

representing two shrew species and 12 rodent species

were captured, representing an overall trap success of

14.1% (Table 1). Three of these captures were shrews,

and thus the overall trap success was 13.9% for rodents.

The trap success by site for rodents exclusively was

11.4% for the pond site, 7.1% for the grassland site, and

24.2% for the dry forest site. Shrews were captured in

traps at both the grassland and pond sites. Twelve spe-

cies of bat were netted at the pond site and two other

species were taken opportunistically in buildings in

Minziro and nearby caves (Table 2).

An interpretation of the species accumulation curves

suggests that most rodent species were documented at

a given site (Figure 2). The one exception was the grass-

land site, where the only specimen of Dendromus mys-

tacalis captured during the entire survey was obtained on

the last day of sampling. The capture of a Dendromus in

a Museum Special is unusual compared to similar sur-

veys conducted by WTS using the same methodology

(Stanley and Hutterer 2007). Our results indicate that the

grassland habitat may not have been adequately sam-

pled to document the entire fauna of this habitat in the

Minziro area.

Six species of shrews were recorded from Minziro

(Table 1), including Crocidura hildegardeae, C. nanilla,

C. cf. nigrofusca, C. olivieri , Suncus megalura, and Syl-

visorex johnstoni . Four specimens of Crocidura cf. nigro-

fusca, a medium-sized shrew with significant pilosity on

the tail, were trapped: two at the grassland site and two

at the pond site. Following the example of Hutterer et al.

(1987), we compared ratios of the width of the two upper

unicuspid teeth (the canines and third upper incisor) and

the least interorbital width of the cranium to those pro-

vided by Dippenaar (1982) for this taxon. The Minziro

specimens cluster with the specimens Dippenaar called

C. zaodon, now referred to as C. nigrofusca (Hutterer

2005), and not C. turba. Dickinson and Kityo (1996) listed

C. turba as one of the species collected in Sango Bay,

Uganda. Hutterer et al. (1987) recorded C. nigrofusca

from Rwanda, and we echo their call for more systematic

work on both C. turba and C. nigrofusca to resolve the

alpha-taxonomy of this difficult group. One specimen of

C. nanilla was taken at the grassland site. This species

has also been recorded in Sango Bay (Dickinson and

Kityo 1996) and Rwanda (Hutterer et al. 1987). One juve-

nile specimen (based on lack of fusion between basioc-

cipital and basisphenoid bones) of Crocidura sp. was

collected at the pond site, but is not included in Table 1.

Hutterer (1986) published the first record of Sylvisorex

johnstoni from Tanzania based on specimens from near

Lusahanga (Figure 1). No other published records exist








Table 2 Bats recorded during the survey of Minziro Forest Reserve and their external measurements (see materials and methods

for definition of abbreviations).

Species TL TV HF EAR FA WT

(mm) (mm) (mm) (mm) (mm) (g)

Epomops franqueti (Tomes 1860) (ns10) 147.1"13.3 21.7"0.5 26.1"1.6 90"3.4 101.3"21

(128–168) (21–22) (25–29) (85–95) (55–130)

Hipposideros caffer (Sundevall, 1846) (ns11) 82.9"4.0 28.9"3.8 8.5"0.7 16.3"1.3 47.8"1.9 8.2"1.0(76–89) (24–35) (8–10) (15–18) (45–51) (6.5–9.8)

Nycteris grandis Peters 1865 (ns1) 143 64 13 28 60 26

Nycteris cf. thebaica E. Geoffroy 1818 (ns1) 48

Chaerephon pumilus (Cretzschmar 1830) (ns2) 87.5"4.9 27.5"4.9 8.5"0.7 14.0"1.4 39.0"1.4 8.2"1.3

(84–91) (24–31) (8–9) (13–15) (38–40) (7.3–9.2)

Glauconycteris gleni Peterson & Smith 1973 (n56) 103.8"4.2 43.7"2.3 7.8"1.2 14.0"1.3 39.5"1.4 8.3"0.4

(98–109) (40–46) (6– 9) (13– 16) (38– 41) (7.7–8.8)

Glauconycteris humeralis J.A. Allen 1917 (n511) 89.7"3.9 42.5"1.9 6.4"0.7 11.0"1.4 35.4"0.5 4.4"0.5

(85–96) (40–45) (6–8) (10–15) (35–36) (4.0–5.7)

Glauconycteris poensis (Gray, 1842) (n59) 91.5"4.4 39.0"3.1 8.0"1.0 14.9"0.3 36.9"1.6 6.0"0.9

(85–100) (34–43) (6–9) (14– 15) (35– 40) (5.2–8.0)

Neoromicia nanus (Peters 1852) (ns8) 75.4"3.9 27.4"2.7 6.1"0.6 11.0"2.2 28.2"2.1 3.7"0.5

(70–81) (24–31) (5–7) (9–16) (25–31) (3.1–4.3)

Neoromicia tenuipinnis (Peters 1872) (ns1) 70 24 6 12 30 3.6

Mimetillus moloneyi (Thomas 1891) (ns1) 93 33 7 13 31 9

Data are presented as mean"SD (range). Species recorded for the first time in Tanzania are in bold font.

for Tanzania. This species was recorded in the Sango

Bay area (Dickinson and Kityo 1996), and Mabira Forest,

both in Uganda (Hutterer 1986). This was the most com-

mon shrew species recorded during our survey, but we

only caught it at the dry forest site. The second most

common species encountered was Suncus megalura,

which was only seen at the grassland site.

A total of eight net hours generated 57 specimens rep-

resenting 12 species (Table 2). All ten specimens of

Megachiroptera collected were Epomops franqueti ,which was confirmed by examination of ridges on the

soft palate (Bergmans 1997). Although known from west-

ern Tanzania, Minziro is on the eastern edge of the known

distribution of this fruit bat. Within the Microchiroptera,

there were notable records, including Glauconycteris gle-

ni and G. poensis, which are the first for Tanzania, and

Mimetillus moloneyi , a little-known species that is poorly

represented in collections (Cotterill 2001). Two speci-

mens of Chaerephon pumilus were collected from a

building in Minziro village, and one specimen of Nycteris

cf. thebaica was obtained in a cave near the village.

Three species of rodents were documented for the first

time in Tanzania during the Minziro survey. These includeDeomys ferrugineus, Lophuromys sikapusi , and Hybomys

univittatus. Another notable species was Oenomys

hypoxanthus, which has only been documented at a few

localities within western Tanzania (Kingdon 1974, Rod-

gers et al. 1984).

Of the three microhabitats sampled, the pond site gen-

erated the highest number of rodent species (eight) and,

notably, genera such as Deomys and Oenomys were only

captured there. Lophuromys sikapusi was only seen at

the grassland site, and although the lowest number of

rodent species was observed at the forest site, this area

had the highest abundance (based on trap success) of

Hylomyscus and Praomys.One Manis tricuspis skeleton was obtained from a res-

ident of Minziro village. The specimen was a sub-adult,

based on open sutures of the skull. Although the distri-

bution map in Kingdon (1971) shows a record of this spe-

cies in Tanzania, the text states that, within eastern

Africa, the species is restricted to Uganda and Kenya.

Swynnerton and Hayman (1951) did not record the spe-

cies as occurring in Tanzania. Although we cannot be

sure that this specimen was actually captured within Tan-

zania given the proximity of Minziro to the Ugandan

border, M. tricuspis was documented by a camera trap

placed within the forest on the Tanzanian side during

August 2006 (Tanzania Mammal Atlas Project, 2006).

Discussion

The results of this survey underscore the importance of

Minziro Forest in both improving information on Tanza-

nia’s mammalian list and understanding the natural

history of various shrew, bat, and rodent species within

eastern Africa. Six species captured represent the first

published records for the country: Glauconycteris gleni ,

G. humeralis, G. poensis, Deomys ferrugineus, Lophu-

romys sikapusi , and Hybomys univittatus. The specimensof G. gleni , G. humeralis, and G. poensis are particularly

noteworthy because of the paucity of records across

their known distributions. Although previously recorded

within Tanzania, Sylvisorex johnstoni and Mimetillus

moloneyi are poorly known. Finally, the known ranges of

Epomops franqueti and Oenomys hypoxanthus have their

eastern boundary barely jutting into western Tanzania.

Musser and Carleton (2005) reported that the eastern lim-

its of Lophuromys sikapusi are still unresolved. Kingdon

(1974) shows a distribution of this species that extends

throughout much of Tanzania and close to the eastern

coast, but no specimen record to confirm the identifica-

tion is mentioned. Hence, the specimens collected duringthis survey represent the most southeasterly locality of

this species in Africa based on documented museum

vouchers.








Figure 2 Species accumulation curves for each of the trap lines set at the three different habitats sampled within Minziro Forest.

Although species accumulation curves suggest that

we caught most of the rodent species occurring at the

pond and forest sites during our survey, capture of the

first (and only) Dendromus in the grassland on the last

day suggests that further sampling is necessary. We also

suggest that additional sampling should be conducted in

these same microhabitats during other times of the year

(i.e., the wet season), and that other habitats in and

around Minziro Forest Reserve be the focus of future

surveys.

The closest low-elevation forests to Minziro that have

been the focus of recent published faunal surveys

include the Ugandan sites of the Kalinzu Forest in the

southeast (Lunde and Sarmiento 2002) and Sango Bay,

just north of the Tanzanian border near the edge of Lake

Victoria (Dickinson and Kityo 1996). Although many of the








species we found in Minziro were also documented dur-

ing these surveys, there were other species that were

found in one or both of the Ugandan sites, but not in

Minziro. The small to medium-sized rodent species doc-

umented in the Kalinzu survey but not in Minziro (for

sample sizes greater than five) include Hylomyscus aeta

and Malacomys longipes. Malacomys and grassland-

dwelling rodents such as Lemniscomys and Mastomys

were documented in Sango Bay, but not in Minziro. Pos-

sible explanations for differences between the faunal lists

generated among the three surveys include differences

in bait (Kalinzu) and in the season sampled (Sango Bay).

The Malacomys observed in Sango Bay were captured

in swamp-forest in March. Subsequent surveys in Minziro

during wetter times of the year may reveal this rodent in

Tanzania. Given the significant number of species previ-

ously unknown from Tanzania before this brief survey,

further study and conservation of the Minziro forest are

warranted. Based on the paucity of such information, fur-

ther botanical study would be especially informative.

Although the fauna and habitats of Minziro are found (and

may be common) in other countries surrounding Tanza-

nia, conservation efforts vary in each of these countries,

both over time and in approach. Hence, conservation

efforts targeting Minziro by Tanzanian scientists and land

managers will broaden the conservation potential of an

important habitat and its biota.

Acknowledgements

We thank Sarah Durant, Chediel Khazael, Philip Kihaule, Zawadi

Mbwambo, Maiko Munissi, Daudi Peterson, Trude Peterson,

Mwemezi Rwiza, Annette Simonson, and John Simonson for

assistance in the field. Steven Goodman offered helpful advice

on the manuscript. The Tanzania Wildlife Research Institute and

Commission of Science and Technology granted permits for this

research project.

Appendix

Field Museum of Natural History (FMNH) catalogue num-

bers for specimens collected during survey of Minziro

Forest.

Crocidura hildegardeae (ns1): FMNH 192911; C. nanilla(ns1): FMNH 193039; C. nigrofusca (ns4): FMNH

192913–192915, 193037; C. olivieri (ns1): FMNH

192912; Crocidura sp. (ns1): FMNH 193038; Suncus

megalura (ns5): FMNH 192916–192917, 193040–

193042; Sylvisorex johnstoni (ns9): FMNH 192918–

192923, 193043–193045; Epomops franqueti (ns10):

FMNH 192924–192929, 193046–193049; Hipposideros

caffer (ns11): FMNH 192930–192935, 193050–193054;

Nycteris grandis (ns1): FMNH 192936; Nycteris cf. the-

baica (ns1): FMNH 192937; Chaerephon pumilus (ns2):

FMNH 192938, 193055; Glauconycteris gleni (ns6):

FMNH 192939–192941, 193056–193058; G. humeralis

(ns11): FMNH 192942–192946, 193059–193064;

G. poensis (ns9): FMNH 192947–192950, 193065–193069; Neoromicia nanus (ns7): FMNH 192952–

192955, 193070–193072; N. tenuipinnis (ns1): FMNH

192956; Mimetillus moloneyi (ns1): FMNH 192951;

Lophuromys aquilus (ns7): FMNH 192960–192962,

193078–193081; L. sikapusi (ns4): FMNH 192963–

192964, 193082–193083; Deomys ferrugineus (ns4):

FMNH 192958–192959, 193075, 193077; Dendromus

mystacalis (ns1): FMNH 193074; Dasymys incomtus

(ns7): FMNH 192965–192966, 193084–193088; Gram-

momys macmillani (ns6): FMNH 192967–192969,

193089–193091; Hybomys univittatus (ns20): FMNH192970–192977, 193092–193103; Hylomyscus stella

(ns86): FMNH 192978–193006, 193104–193162,

193194; Mus musculoides (ns1): FMNH 193007; M. tri-

ton (ns1): FMNH 193163; Oenomys hypoxanthus (ns1):

FMNH 193008; Praomys jacksoni (ns73): FMNH

193009–193036, 193144, 193164–193193, 193195–

193208; Graphiurus murinus (ns1): FMNH 193209.

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A Survey of the Small Mammals of Minziro Forest, Tanzania, With Several Additions to the Known Fauna...

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