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Mammalia 72 (2008): 116122  2008 by Walter de Gruyter Berlin New York. DOI 10.1515/MAMM.2008.015

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


with several additions to the known fauna of the country

William T. Stanley1,* and Charles A.H. Foley2


Division of Mammals, Field Museum of Natural
History, 1400 South Lake Shore Drive, Chicago,
IL 60605, USA, e-mail: bstanley@fieldmuseum.org
2
Wildlife Conservation Society, P.O. Box 2703,
Arusha, Tanzania

Materials and methods

Study area

Complete faunal lists are vital for understanding the natural 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 northwestern Tanzania. Six species are documented here for
the first time for Tanzania. Other records include Tanzanian species poorly represented as vouchers in collections. Minziro is a unique forest worthy of conservation
priority and further study.

Minziro Forest Reserve is in Bukoba Region, Minziro District, 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: MarchMay and
OctoberNovember. 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).

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

Trapping methodology

*Corresponding author

Abstract

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 lowland
forests of Tanzania remain poorly known. One such forest
is the Minziro Forest Reserve, situated in the northwestern corner of Tanzania (Figure 1). Contiguous with Malabigambo 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 examples 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.

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 operation for a 24-h period (07:0007:00 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 each


night 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 calculated these values by dividing the number of individuals captured by the number of trap or bucket nights and
multiplying by 100. When discussing the two trapping
methodologies 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 success


rate 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 measurements 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, liver, and kidney were preserved in EDTA buffer. All specimens are deposited in the Field Museum of Natural
History (FMNH; catalogue numbers are listed in Appendix), and a portion will be returned to the Department of

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

Zoology, University of Dar es Salaam. We follow the taxonomy 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 captured 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 species 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 grassland site, where the only specimen of Dendromus mystacalis 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 surveys conducted by WTS using the same methodology
(Stanley and Hutterer 2007). Our results indicate that the
grassland habitat may not have been adequately sampled 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 Sylvisorex johnstoni. Four specimens of Crocidura cf. nigrofusca, 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 provided 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 juvenile specimen (based on lack of fusion between basioccipital 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 1 Distribution of shrews and rodents in three different microhabitats in Minziro Forest Reserve in August 2006.

Crocidura hildegardeae Thomas 1904


Crocidura nanilla Thomas 1909
Crocidura nigrofusca Matschie 1895
Crocidura olivieri (Lesson 1827)
Suncus megalura (Jentink 1888)
Sylvisorex johnstoni (Dobson 1888)
Lophuromys aquilus (True 1892)
Lophuromys sikapusi (Temminck 1853)
Deomys ferrugineus Thomas 1888
Dendromus mystacalis Heuglin 1873
Dasymys incomtus (Sundevall 1847)
Grammomys macmillani (Wroughton 1907)
Hybomys univittatus (Peters 1876)
Hylomyscus stella (Thomas 1911)
Mus musculoides Temminck 1853
Mus triton (Thomas 1909)
Oenomys hypoxanthus (Pucheran 1855)
Praomys jacksoni (de Winton 1897)
Graphiurus murinus (Desmarest 1822)
Total number of individuals
Total number of species
Total number of sampling nights
Trap success (%)

Forest

Grassland

Pond

Totals

0
0
0
0
0
9
0
0
0
0
0
0
8
56
0
0
0
35
0
100
3
664
15.1

1
1
2
0
5
0
1
4
0
1
7
0
0
1
1
1
0
0
0
32
10
328
9.7

0
0
2
1
0
0
6
0
4
0
0
6
12
31
0
0
1
38
1
102
10
1063
9.6

1
1
4
1
5
9
7
4
4
1
7
6
20
88
1
1
0
73
1
234
15
2055
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.

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W.T. Stanley and C.A.H. Foley: Small mammals of Minziro Forest, Tanzania 119

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
(mm)

Epomops franqueti (Tomes 1860) (ns10)


Hipposideros caffer (Sundevall, 1846) (ns11)
Nycteris grandis Peters 1865 (ns1)
Nycteris cf. thebaica E. Geoffroy 1818 (ns1)
Chaerephon pumilus (Cretzschmar 1830) (ns2)
Glauconycteris gleni Peterson & Smith 1973 (n56)
Glauconycteris humeralis J.A. Allen 1917 (n511)
Glauconycteris poensis (Gray, 1842) (n59)
Neoromicia nanus (Peters 1852) (ns8)
Neoromicia tenuipinnis (Peters 1872) (ns1)
Mimetillus moloneyi (Thomas 1891) (ns1)

TV
(mm)

HF
(mm)

EAR
(mm)

FA
(mm)

WT
(g)

147.1"13.3
(128168)
82.9"4.0
(7689)
143

28.9"3.8
(2435)
64

21.7"0.5
(2122)
8.5"0.7
(810)
13

26.1"1.6
(2529)
16.3"1.3
(1518)
28

101.3"21
(55130)
8.2"1.0
(6.59.8)
26

87.5"4.9
(8491)
103.8"4.2
(98109)
89.7"3.9
(8596)
91.5"4.4
(85100)
75.4"3.9
(7081)
70
93

27.5"4.9
(2431)
43.7"2.3
(4046)
42.5"1.9
(4045)
39.0"3.1
(3443)
27.4"2.7
(2431)
24
33

8.5"0.7
(89)
7.8"1.2
(69)
6.4"0.7
(68)
8.0"1.0
(69)
6.1"0.6
(57)
6
7

14.0"1.4
(1315)
14.0"1.3
(1316)
11.0"1.4
(1015)
14.9"0.3
(1415)
11.0"2.2
(916)
12
13

90"3.4
(8595)
47.8"1.9
(4551)
60
48
39.0"1.4
(3840)
39.5"1.4
(3841)
35.4"0.5
(3536)
36.9"1.6
(3540)
28.2"2.1
(2531)
30
31

8.2"1.3
(7.39.2)
8.3"0.4
(7.78.8)
4.4"0.5
(4.05.7)
6.0"0.9
(5.28.0)
3.7"0.5
(3.14.3)
3.6
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 common 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 representing 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 western Tanzania, Minziro is on the eastern edge of the known
distribution of this fruit bat. Within the Microchiroptera,
there were notable records, including Glauconycteris gleni 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 specimens 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 include
Deomys 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, Rodgers et al. 1984).
Of the three microhabitats sampled, the pond site generated 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 resident 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 species 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 species as occurring in Tanzania. Although we cannot be
sure that this specimen was actually captured within Tanzania 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 Tanzanias 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, Lophuromys sikapusi, and Hybomys univittatus. The specimens
of 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 limits 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 identification is mentioned. Hence, the specimens collected during
this survey represent the most southeasterly locality of
this species in Africa based on documented museum
vouchers.

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

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

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W.T. Stanley and C.A.H. Foley: Small mammals of Minziro Forest, Tanzania 121

species we found in Minziro were also documented during 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 documented in the Kalinzu survey but not in Minziro (for
sample sizes greater than five) include Hylomyscus aeta
and Malacomys longipes. Malacomys and grasslanddwelling rodents such as Lemniscomys and Mastomys
were documented in Sango Bay, but not in Minziro. Possible 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 previously unknown from Tanzania before this brief survey,
further study and conservation of the Minziro forest are
warranted. Based on the paucity of such information, further botanical study would be especially informative.
Although the fauna and habitats of Minziro are found (and
may be common) in other countries surrounding Tanzania, 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 numbers for specimens collected during survey of Minziro
Forest.
Crocidura hildegardeae (ns1): FMNH 192911; C. nanilla
(ns1): FMNH 193039; C. nigrofusca (ns4): FMNH
192913192915, 193037; C. olivieri (ns1): FMNH
192912; Crocidura sp. (ns1): FMNH 193038; Suncus
megalura (ns5): FMNH 192916192917, 193040
193042; Sylvisorex johnstoni (ns9): FMNH 192918
192923, 193043193045; Epomops franqueti (ns10):
FMNH 192924192929, 193046193049; Hipposideros
caffer (ns11): FMNH 192930192935, 193050193054;
Nycteris grandis (ns1): FMNH 192936; Nycteris cf. thebaica (ns1): FMNH 192937; Chaerephon pumilus (ns2):
FMNH 192938, 193055; Glauconycteris gleni (ns6):
FMNH 192939192941, 193056193058; G. humeralis
(ns11): FMNH 192942192946, 193059193064;
G. poensis (ns9): FMNH 192947192950, 193065
193069; Neoromicia nanus (ns7): FMNH 192952
192955, 193070193072; N. tenuipinnis (ns1): FMNH
192956; Mimetillus moloneyi (ns1): FMNH 192951;

Lophuromys aquilus (ns7): FMNH 192960192962,


193078193081; L. sikapusi (ns4): FMNH 192963
192964, 193082193083; Deomys ferrugineus (ns4):
FMNH 192958192959, 193075, 193077; Dendromus
mystacalis (ns1): FMNH 193074; Dasymys incomtus
(ns7): FMNH 192965192966, 193084193088; Grammomys macmillani (ns6): FMNH 192967192969,
193089193091; Hybomys univittatus (ns20): FMNH
192970192977, 193092193103; Hylomyscus stella
(ns86): FMNH 192978193006, 193104193162,
193194; Mus musculoides (ns1): FMNH 193007; M. triton (ns1): FMNH 193163; Oenomys hypoxanthus (ns1):
FMNH 193008; Praomys jacksoni (ns73): FMNH
193009193036, 193144, 193164193193, 193195
193208; Graphiurus murinus (ns1): FMNH 193209.

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