European Journal of Taxonomy 42: 1-18

http://dx.doi.org/10.5852/ejt.2013.42

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ISSN 2118-9773

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2013 • Guy-Crispin Gembu Tungaluna et al.

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Research article

urn:lsid:zoobank.org:pub:4D07035D-79AF-4BFA-8BEE-lAB35EB2C9ED

Rediscovery of Glauconycteris superba Hayman, 1939

(Chiroptera: Vespertilionidae) after 40 years at Mbiye Island,

Democratic Republic of the Congo

Guy-Crispin GEMBU TUNGALUNA 1 , Victor VAN CAKENBERGHE 2 ,

Prescott MUSABA AKAWA 3 , Benjamin DUDU AKAIBE 4 , Erik VERHEYEN 5 ,

Frits DE VREE 6 & Jakob FAHR 7

134 LEGERA, Faculte des Sciences, Universite de Kisangani, B.P. 2012 Kisangani, DRC.

2 6 Functional Morphology Group, Department of Biology, Universiteit Antwerpen,

Campus Drie Eileen, Universiteitsplein 1, B-2610 Antwerpen (Wilrijk), Belgium.

5 Evolutionary Ecology Group, Department of Biology, Universiteit Antwerpen,

Groenenborgerlaan 171, B-2020 Antwerpen, Belgium

5 Vertebrate Department, Royal Belgian Institute of Natural Sciences,

Vautierstraat 29, B-1000 Brussel, Belgium

7 Division of Evolutionary Biology, Zoological Institute, TU Braunschweig, Mendelssohnstr. 4,

D-38106 Braunschweig, Germany

Emails: 1 [email protected]. 2 [email protected] (corresponding author)

3 prescottmusaba@y ahoo.fr . 4 duduakaibe@v ahoo.If . 5 [email protected] .

6 [email protected] . 7 [email protected]

1 um:lsid:zoobank. org: author: 2AE2F96F-A 13 8-419B-B650-6DB44D53 5D14

2 urn:lsid:zoobank.org:author:9A0E 1AF5-C248-4648-9D64-443112890346

3 urn:lsid:zoobank.org:author:4C40421C-lD84-4BDl-A444-360300ECBEEC

4 um:lsid:zoobank.org:author:F983BCBD-121D-4475-AA59-210981D397C2

5 urn: 1 si d: zoobank, org: author: 86B40463-E3D9-4147-9ED3-D7302E0D64B6

6 urn:lsid:zoobank.org:author:8F7DFFBB-099B-48Fl-A427-3FA5CFF9A8E5

7 um:lsid:zoobank, org: author: 866DF6B9-FBDB-4D8B-A438-C2F1564B780D

Abstract. We report the rediscovery of the Pied Butterfly Bat, Glauconycteris superba Hayman, 1939,

40 years after this species was last recorded. The new specimen from Mbiye Island, Democratic Republic

of the Congo, is compared with the type specimens of G. s. superba and G. superba sheila Hayman, 1947

and a specimen from Matonguine, Ivory Coast. The variation in the striking colouration of the pelage

as well as in morphometric data is considered to be individual rather than geographic variation and we

tentatively regard G. s. sheila as a synonym of the nominate form. Despite the wide distribution of this

species in the tropical forest zone of West and Central Africa, only four specimens from four localities

are known to date, which might indicate very specific habitat preferences. Contemporary land cover

information around historic collection sites shows degraded landscapes. Given the highly uncertain area

of occupancy of this species, we suggest changing the status of G. superba in the IUCN Red Fist of

Threatened Species from “Feast Concern” to “Data Deficient”.

Key words. Bats, morphometries, taxonomy, distribution, conservation.

1

European Journal of Taxonomy 42: 1-18 (2013)

Gembu Tungaluna G.-C., Van Cakenberghe V., Musaba Akawa P., Dudu Akaibe B., Verheyen E., De Vree F.

& Fahr J. 2013. Rediscovery of Glauconycteris superba Hayman, 1939 (Chiroptera: Vespertilionidae) after 40

years at Mbiye Island, Democratic Republic of the Congo. European Journal of Taxonomy 42: 1-18. http://dx.doi.

org/10.5852/eit.2013.42

Introduction

Glauconycteris superba Hayman, 1939, the Pied or Superb Butterfly Bat, is one of the most spectacularly

coloured bats found in Africa. It is the largest species of the genus and characterised by black wings and

a predominantly black pelage, which is strikingly marked with white spots and stripes on the head, back

and belly.

Despite this unique and conspicuous colouration, only two additional specimens had been collected since

its discovery in 1938 in Pawa, Democratic Republic of the Congo (DRC): one on 6 Dec. 1946 in Oda,

southern Ghana (and designated as type of Glauconycteris superba sheila Hayman, 1947), and one on 24

Jan. 1972 in Matonguine [sometimes spelled as Matongouine], southwestern Ivory Coast (Fahr 2013).

In this paper we report on a fourth specimen, constituting the rediscovery of this species after 40 years

since the last record, present data on its morphology and distribution, characterize climatic and land cover

conditions at the collection sites, and discuss the taxonomy and conservation status of G. superba.

Material and Methods

Abbreviations used in text and figures

3phl

3ph2

3ph2+3

4phl

4ph2

5phl

5ph2

ad

ale

BBUL

BcH

BDA

Bm

BMNH

Brain

CBF

CBF+

CBFC

CBFC+

C-C

C-M 3

c-m 3

Conang

Concor

Cor

DRC

Ear

length of first phalanx of third finger

length of second phalanx of third finger

length of combined second and third phalanges of third finger

length of first phalanx of fourth finger

length of second phalanx of fourth finger

length of first phalanx of fifth finger

length of second phalanx of fifth finger

adult

alcoholic or fluid-preserved specimen

width between the auditory bullae

height of braincase

Benjamin Dudu Akaibe

body mass

Natural History Museum, Fondon

width of braincase

condylobasal length to base of incisor

condylobasal length to tip of incisor

condylobasal length to base of canine

condylobasal length to tip of canine

width across the upper canines

length of maxillary toothrow

length of mandibular toothrow

distance between condylar and angular processes

distance between condylar and coronoid processes

height at the coronoid process

Democratic Republic of the Congo

ear length

2

GEMBU TUNGALUNA G.-C. et al ., Rediscovery of Glauconycteris superba Hayman, 1939

FA

FA-

FA+

FDV

GCGT

GFS

GFS+

GFSC

GFSC+ =

H&B

HFcu

1 1

1 2

3

lor =

IUCN

FBUFF =

FBUFW =

leg

M 3

M 3 -M 3

Mand =

Mand2 =

Mast =

Meta3 =

Meta4 =

Meta5 =

OCC

PalFen =

PMA

POSTPAF =

PTB

RBLJFF =

RBUFW =

RMCA =

ROSTR =

SHB

SHBB

SHRM

subad

Tail

Thumb =

Tibia

Tot =

Tra =

VVC

WS

z yg

forearm length (used when it is not clear whether or not the wrist is included)

forearm length excluding wrist

forearm length including wrist

Frits De Vree

Guy-Crispin Gembu Tungaluna

greatest length of skull to base of incisor

greatest length of skull to tip of incisor

greatest length of skull to base of canine

greatest length of skull to tip of canine

length of head and body

length of hindfoot with claws

inn er upper incisor

outer upper incisor

first lower incisor

second lower incisor

third lower incisor

least interorbital width

International Union for Conservation of Nature

length of the left auditory bulla

width of the left auditory bulla

legate (collected/donated by)

posterior upper premolar

width across the posterior upper molars

mandibular length from condylar process to base of y

mandibular length from angular process to symphysis between the mandibular rami

mastoid width

length of metacarpal of third finger

length of metacarpal of fourth finger

length of metacarpal of fifth finger

occipitonasal length

length of palatum

Prescott Musaba Akawa

length of postpalatum

width between the pterygoid processes

length of the right auditory bulla

width of the right auditory bulla

Royal Museum for Central Africa, Tervuren

width across rostrum

height of skull over the auditory bullae

height of skull in front of the auditory bullae

height of rostrum over the molars

subadult

tail length

length of the thumb

tibia length

total length

tragus length

Victor Van Cakenberghe

wingspan

zygomatic width

3

European Journal of Taxonomy 42 : 1-18 ( 2013 )

Study site

Field work was conducted at Mbiye, an island situated in the Congo River opposite the city of Kisangani

(Fig. 1). In 2012, the LEGERA team (GCGT, PMA, BDA) deployed ten mist nets (10 m x 2.5 m, mesh

20 mm ) for five nights each month from January to June.

Mbiye Island is located in the Guineo-Congolian lowland forests of the Congo River basin. Dominant

tree species on the island are Coelocaryon botryoides Verm., Gilbertiodendron dewevrei (De Wild) J.

Leonard and Scorodophloeus zenker i Harms (Fabaceae). In the immediate vicinity of the mistnet where

the Glauconycteris specimen was collected, we recorded the following tree species: Agelaeapentagyna

(Lam.) Baill. (Connaraceae), Berlinia grandiflora (Vahl) Hutch, and Dalziel (Fabaceae), Coelocaryon

preussii Warb. (Myristicaceae), Cola digitata Mast. (Malvaceae), Drypetes bipindensis (Pax) Hutch.

(Putranjivaceae), Pterygota bequaertii De Wild. (Malvaceae), and Trilepisium madagascariense DC.

(Moraceae) (for plant identification, see Lejoly et al. 2010; for details on the Mbiye Island, see Nshimba

2008). The natural vegetation of lowland wet forest has been variably degraded on Mbiye Island to

secondary forest and fallow land (“jacheres”) by human land use.

Specimens examined

Besides the new specimen, two additional specimens in the collection of the Royal Museum for Central

Africa (RMCA, Tervuren, Belgium) were examined, and photos of the specimen in the collection of the

Natural History Museum (BMNH), London, were made available for comparison. Age was determined

based on the ossification of the epiphyses of the wing bones. Details of the specimens are given in the

Appendix.

blue point shows the collection locality of G. superba Hayman, 1939.

4

GEMBU TUNGALUNA G.-C. et al ., Rediscovery of Glauconycteris superba Hayman, 1939

Measurements

In the field, eight external measurements were taken by GCGT. Bm was measured with a spring balance

(Pesola; accuracy 1 g). WS was measured with a ruler (accuracy 1 mm). The other measurements were

taken using digital callipers (Mitutoyo; accuracy 0.05 mm, rounded to 0.1 mm): FA, Ear, Tail, Tra, Tot

and Tibia.

Measurements by FDV were taken using a dial callipers (Helios; accuracy 0.05 mm, rounded to 0.1 mm).

Independent measurements were taken by VVC on the specimens available at the RMCA using dial

callipers (Helios; accuracy 0.05 mm, rounded to 0.1 mm). Craniodental measurements include GLS,

GLS+, GLSC, GLSC+, CBL, CBL+, CBLC , CBLC+, OCC, lor, Zyg, Mast, Brain, SHB, SHBB,

ROSTR, C-M 3 , C-C, M 3 -M 3 , PalLen, POSTPAL, PTB, BBUL, LBULL, LBULW, RBULL, RBULW,

Mand, C-M 3 , Mand2, Conang, Concor, and Cor. The external measurements comprise FA+, FA-, Meta3,

3phl, 3ph2+3, Meta4, 4phl, 4ph2, Meta5, 5phl, 5ph2, Tibia, Ear, Tra, Tail, H&B, Thumb, HFcu.

Fig. 2. Pelage pattern of the freshly collected Glauconycteris superba Hayman, 1939 specimen from

Mbiye Island (RMCA al.097-M-3153). Upper left: dorsal view, upper right: ventral view, lower left:

anterior-dorsal view, lower right: detail of head and breast.

5

European Journal of Taxonomy 42 : 1-18 ( 2013 )

Results

Order Chiroptera Blumenbach, 1779

Family Vespertilionidae Gray, 1821

Subfamily Vespertilioninae Gray, 1821

Genus Glauconycteris Dobson, 1875

Glauconycteris superba Hayman, 1939

On 19 Feb. 2012, one of ten mistnets was set up in secondary forest on the island, near a clearing where

crops were cultivated. During this session (from 18:00 to 06:00) one adult male Glauconycteris superba

(Fig. 2) entered the net between 19:00 and 20:00, and was captured alongside a Megaloglossus woermanni

Pagenstecher, 1885, a Rhinolophus landeri Martin, 1838, and a Casinycteris argynnis Thomas, 1910.

The Glauconycteris specimen was preserved in 90% ethanol and deposited in the collections of the

RMCA, where the skull was extracted and cleaned.

Morphological comparison of the four specimens

Figure 3 shows the variation in the white spots among the different specimens. Unfortunately, the

specimen from Matonguine has suffered major damage as it had been dried out for an extended period

of time, which resulted in loss of fur and decolouration of the remaining pelage. Still, it is possible to

Fig. 3. Pelage pattern of the four known specimens of Glauconycteris superba Hayman, 1939 (upper row:

dorsal, lower row: ventral). From left to right: holotype of G. s. sheila Hayman, 1947 (BMNH 47.10),

holotype of G. superba (RMCA 14765), new specimen from Mbiye Island (RMCA al.097-M-3153),

and the specimen from Matonguine (RMCA 97.077-M-7719). Scale bar = 2 cm.

6

GEMBU TUNGALUNA G.-C. et al. , Rediscovery of Glauconycteris superba Hayman, 1939

distinguish the white spots on the belly and the head. Drawings of the head made shortly after the time

of collection (Fig. 4), illustrate that spots on the head of this specimen were almost identical to those

of the other specimens. Am ong the three remaining specimens, the Mbiye specimen is the darkest, but

Hayman (1939, 1947) described his two type specimens as being black, so probably their pelages have

faded over the years. Hayman (1947) also indicated that the type specimen of G. s. sheila differed from

the type of G. s. superba in having “conspicuous unpigmented areas on the upper surface of the elbow,

knee and ankle joints”, suggesting a different colour pattern of G. s. superba. However, as illustrated in

Fig. 3, these unpigmented areas are definitely present on the knees, but not as obvious on the elbows,

although the paler colour of the arms might blur this. In the new specimen, these pale areas are present

on elbows, knees, and ankles.

The various white spots exhibit some individual variation. The two lateral white bands on the belly are

widest in the type of G. s. sheila , followed by the Mbiye specimen, and narrowest in the type of G. s.

superba. However, since a major part of the belly was removed from the latter specimen, the width of

the bands might be underestimated.

The white spot on the throat varies not as much in its width as in the extension to the sides of the head,

where it may or may not become visible on the dorsal side. In Fig. 3, it appears as if the specimen from

Mbiye has a divided throat spot, but this is due to the fact that the throat was cut open to remove the

skull.

The three white spots on the head - one median on the nose and two on the crown of the head near the

bases of the ears - also show some variation in size, especially the ear spots.

The largest spots on the back are fairly similar in all three specimens. The two spots between the

shoulders are widest near the head and tapering to the middle of the back. In the type of G. s. sheila ,

the distance between the two spots seems to be much wider than in the two other specimens, but this

is an artefact due to the position of the shoulder blades, which are fixed close to each other in the latter

specimens, thereby concealing the connecting skin in a cavity. From the middle of the back, about the

height where the two anterior spots end, two bands run to the lower back where they nearly meet. These

two mid-back bands diminish in width from the anterior to the posterior side. In the specimen from

Mbiye, the two bands seem to be subdivided in multiple spots, but this is due to the orientation of the

individual hairs on the photo, where they are stretched to the sides of the body and not along the length

of the body as for the other specimens.

The major difference, however, is in the position, size and number of spots between the mid-back

shoulder spots and the neck region. In the two Congolese specimens, only two spots are present on each

side, whereas the type of G. s. sheila and the specimen from Matonguine have three. These extra spots

were already reported by Hayman (1947). Hayman links the anterior-most spots with the white belly

Fig. 4. Head of Glauconycteris superba Hayman, 1939 from Matonguine (RMCA 97.077-M-7719).

7

European Journal of Taxonomy 42 : 1-18 ( 2013 )

bands, which end just in front of the spots (see also Cansdale’s photo in Nowak 1999). In the type of

G. s. superba, the anterior spots seems to run into belly bands, but in the Mbiye specimen, these ventral

bands do not reach as far anteriorly. Here the throat spot reaches slightly further towards the anterior

spot of the back. So the anterior-most spots on the back might possibly form a connection with either the

belly or the throat spots on the ventral side.

For the specimen from Mbiye, Fig. 3 also shows on the anterior side of the mid-back bands a slight

separation between the main part of these bands and its anterior-most part (again depending on how the

individual hairs are combed). If more dark hairs were present, this could lead to a split off of the anterior-

most part of the mid-back bands. This split-off spot would then be almost similar to the posterior-most

small spot found in the type of G. s. sheila , although in the latter specimen this spot is slightly more

laterally positioned.

Fig. 5. Crania of the four known specimens of Glauconycteris superba Hayman, 1939 (left row: dorsal,

middle row: ventral; right row: lateral). From top to bottom: specimen from Matonguine (RMCA 97.077-

M-7719, (J), Mbiye Island (RMCA al.097-M-3153, $), holotype of G. superba (RMCA 14765, ?), and

holotype of G. s. sheila Hayman, 1947 (BMNH 47.10, $). Scale bar = 1 cm

8

GEMBU TUNGALUNA G.-C. et al. , Rediscovery of Glauconycteris superba Hayman, 1939

The skulls are very similar in size and shape (Tabs 1-2, Figs 5-6) although the type of G. s. superba is

clearly subadult as indicated by the incompletely fused epiphyses of the wing bones. All skulls show

a marked angle at the junction between rostrum and braincase as mentioned by Hayman (1939, 1947).

Hayman (1947) also noted that the posterior upper molar (M 3 ) is considerably reduced. This is indeed

the case for all four of the skulls, and most pronounced in the specimen from Matonguine. All skulls

have weak sagittal and lambdoidal crests, which are least developed in the type of G. 5. superba and

most developed in the type of G. s. sheila. The lack of crest development in the type of G. 5. superba

is probably age-related. Another character mentioned by Hayman (1939) is the presence of the minute

accessory cusplet near the cingulum of the inner upper incisor (I 1 ). The development of this cusplet

is variable, being most pronounced in the type of G. s. sheila and least in the specimen from Mbiye.

The position of the outer upper incisor (I 2 ), described by Hayman (1939) as being “closely crowded

between I 1 and the canine”, is indeed very crowded in both type specimens, but this is less the case in

the two other specimens. The lower incisors are closely crowded as mentioned by Hayman (1939), and

have three cusps. The inner incisors are more or less rectangular in shape, whereas i 0 and i 3 are more

triangular.

Morphometric comparison of the four specimens

When based on the collector’s measurements, the specimen from Mbiye Island has a considerably

shorter forearm (FA) and Tibia compared to published values (Tab. 1). The comparatively small

forearm initially challenged our identification of the Mbiye specimen as G. superba since the FA is

given variably in the literature as 47 mm (Hayman & Hill 1971), greater than 44.0 mm (Peterson &

Smith 1973), and 45 mm (Rambaldini 2010). However, these differences might be partially explained

by individual measurement techniques. Therefore the three specimens currently available at the RMCA

were measured again (Tabs 2-3).

The skull measurements of the three specimens in the RMCA are very similar (Tab. 2). Externally,

the dimensions of the new specimen fit very well with those of the specimen from Matonguine, with

the exception of Tail length, but this is primarily due to the fact that the tail could not be completely

stretched. The tail length appears to be very similar to that of the type from Pawa, but in the latter

specimen the point of origin of the tail could not be determined as the abdominal region (including the

genital area) had been completely removed during preparation. Considering that the type is subadult, it

is remarkable that its long bones (FA and metacarpals) are much larger than the corresponding bones in

Fig. 6. Mandibles of the four known specimens of Glauconycteris superba Hayman, 1939 (upper row:

dorsal view, lower row: lateral view). From left to right: holotype of G. s. sheila Hayman, 1947 (BMNH

47.10, 5) [lateral picture flipped to fit the orientation of the other pictures], holotype of G. superba

(RMCA 14765, ?), new specimen from Mbiye Island (RMCA al.097-M-3153, <$), and the specimen

from Matonguine (RMCA97.077-M-7719, <$). Scale bar = 1 cm.

9

European Journal of Taxonomy 42 : 1-18 ( 2013 )

Table 1. Craniodental and external measurements of Glauconycteris superba Hayman, 1939. Reference:

1. Randolph L. Peterson (courtesy of Judith Eger, pers. comm. 1 Nov. 2001; column 1). 2, 5, 7. FDV.

3. Rosevear (1965). 4. Hayman (1947). 6. Hayman (1939). 8. GCGT (field measurements). For

measurement descriptions see text.

RMCA

97.077-M-7719

BMNH

47.10

RMCA

14765

RMCA

al.097-M-3153

Locality

Matonguine

Oda

Pawa

Mbiye

Sex

6

$

6 s

Reference

l

2

3

4

5

6

7

8

Bm

14.0

Tot

112

88

91

H&B

63

49

66

Tail

49

39

25

Ear

16.0

13.0

13.4

Tra

7.0

7.5

5.5

FA

44.7

44.5

46.0

47.0

46.2

47.5

47.6

42.6

WS

316.0

2nd digit

44.5

Meta3

42.0

40.0

41.0

45.5

44.0

3phl

18.7

18.5

20.7

21.0

3ph2

25.0

24.5

3ph2+3

22.4

24.3

Meta4

40.6

38.2

44.0

43.1

4phl

12.2

12.0

13.3

4ph2

9.7

9.8

10.8

10.2

Meta5

35.5

33.2

40.0

38.3

5phl

7.6

8.2

8.5

5ph2

7.0

8.2

7.5

Tib

20.0

19.0

21.0

21.3

20.3

18.6

Calcar

c. 11.0

HFcu

10.0

8.3

GLS

15.1

16.0

15.3

GLS+

16.4

15.6

16.5

16.1

16.5

16.2

15.9

CBL

15.4

14.9

15.7

15.6

15.5

15.2

PalLen Med

6.9

7.7

5.6

6.5

6.9

Mast

9.5

8.8

9.9

9.5

Brain

8.5

7.9

9.1

9.0

8.5

8.3

BcH

6.7

9.6

8.8

Zyg

11.0

11.9

12.0

11.0

C-C

5.8

5.7

6.2

6.0

5.7

M 3 -M 3

7.5

7.4

8.2

8.1

7.9

7.7

C-M 3

5.8

5.6

6.2

6.0

5.8

lor

4.7

4.8

4.9

5.0

Mand

11.6

11.4

12.9

12.5

12.2

11.7

c-m 3

6.4

6.3

6.7

6.6

6.4

the two other specimens, and - to a lesser extent - this also is the case for its tibia length (Tab. 3). Based

on the skull measurements, however, this specimen had probably nearly reached its adult size.

In 1972, FDV examined the three specimens available at that time. He noticed that the genital area of the

subadult type specimen from Pawa was damaged and that it could not be identified as “probably a male”

as mentioned by Hayman (1939). However, his examination also revealed that the nipples were much

10

GEMBU TUNGALUNA G.-C. et al ., Rediscovery of Glauconycteris superba Hayman, 1939

Table 2. Craniodental measurements (rounded to nearest 0.1 mm). For measurement descriptions see text.

RMCA

97.077-M-7719

RMCA

14765

RMCA

al.097-M-3153

Locality

Matonguine

Pawa

Mbiye

Sex

8

?

8

Age

ad

subad

ad

GLS

14.8

14.9

14.8

GLS+

15.6

15.7

15.9

GLSC

15.1

14.8

15.1

GLSC+

16.2

15.2

16.2

CBL

14.6

15.3

15.1

CBL+

15.3

15.6

15.3

CBLC

14.7

15.0

14.8

CBLC+

15.2

15.6

15.2

OCC

12.4

12.9

12.3

IOR

4.7

4.9

ZYG

11.0

MAST

8.7

9.4

9.1

BRAIN

8.5

8.6

8.3

ROSTR

6.3

6.6

6.7

C-M 3

5.5

5.7

5.5

C-C

5.7

5.5

6.2

M 3 -M 3

7.3

7.5

7.3

PalLen

6.1

5.7

POSTPAL

5.2

5.0

5.6

PTB

1.1

1.4

1.3

BBUL

0.9

1.3

LBULW

2.9

3.0

2.7

LBULL

3.7

RBULW

3.3

2.8

3.1

RBULL

3.8

3.7

SHB

8.0

8.8

8.3

SHBB

6.8

6.7

6.5

SHRM

5.5

5.7

5.5

MAND

11.5

11.6

11.4

C-M,

6.3

6.4

6.0

MAND2

8.9

9.0

9.4

CONCOR

3.0

3.6

3.4

COR

3.8

4.5

4.3

CONANG

2.4

2.7

2.5

better developed than those of the male specimen from Matonguine and that, therefore, the holotype

might represent a female rather than a male.

The forearm length of the type of G. superba agrees very well with that of the type of G. s. sheila (which

is a female) and is larger than the specimens from Mbiye and Matonguine (which are males; Tab. 1).

This lends support to the suggestion that both types are indeed females. However, the limited number of

available specimens does not allow a quantification of the intraspecific variation or sexual dimorphism,

hence we are reluctant in assigning a sex to the type specimen of G. superba. If the type specimen is

indeed a female, then this suggests that sexual dimorphism might also be present in G. superba , the

11

European Journal of Taxonomy 42: 1-18 (2013)

Table 3. External measurements (rounded to nearest 0.1 mm). For measurement descriptions see text.

RMCA

97.077-M-7719

14765

al.097-M-3153

Locality

Matonguine

Pawa

Mbiye

Sex

?

Age

ad

subad

ad

FA-

43.3

46.8

42.8

FA+

43.9

47.3

43.1

Meta3

37.5

42.0

37.7

3phl

17.7

20.2

18.2

3ph2+3

24.8

25.7

26.6

Meta4

36.0

41.1

37.0

4phl

11.8

13.3

11.7

4ph2

8.1

9.1

8.9

Meta5

32.6

36.4

32.9

5phl

7.5

9.3

6.4

5ph2

7.5

6.7

7.2

Tib

16.7*

19.0

16.8

Ear

13.8

13.3

14.3

Tra

6.7

5.3

6.2

Tail

39.0

29.8*

30.7*

H&B

55.8

53.8

Thumb

6.2

6.7

6.5

HFcu

9.7

8.4

9.7

* The tail length of the specimen from Pawa is an approximation as the genital area was removed during preparation of

the specimen. The tail of the specimen from Mbiye could not be stretched completely. Both tibiae of the specimen from

Matonguine were damaged.

females being larger than the males, as was reported by Koopman (1971) for other species of the genus

Glauconycteris.

Discussion

Currently known variation in pelage patterning and morphometric data of G. superba does not indicate a

clear geographic signal and might suggest individual variation as found in other Glauconycteris-spQciQS.

We concur with Rosevear (1965) that the characters differentiating G. s. sheila from the nominate form

might represent the range of variation within populations, although the currently available sample size

is probably too limited for a definite conclusion regarding the taxonomic status of G. s. sheila. As such,

we tentatively consider G. s. sheila a synonym of the nominate form.

The four localities of G. superba are located in the tropical forest biome of West and Central Africa at

elevations between 137 m and 775 m (Tab. 4, Fig. 7). Within this region, Matonguine, Oda and Pawa

are characterized by semi-deciduous forest at the periphery of evergreen lowland forest while Mbiye is

located in the central Congo Basin. Today, land cover around the historic collection sites at Pawa, Oda

and Matonguine is a highly fragmented mosaic of secondary forests, farmland and human settlements

as assessed with Google Earth (Oct. 2012), while Mbiye Island is covered by a mosaic of closed and

secondary forest. Thus, contemporary satellite data on land cover show a mixed picture of both closed

forests and mosaics of forest and croplands around the four localities (Tab. 4). However, more data are

needed to evaluate whether G. superba critically depends on closed forests or to what extent this species

tolerates habitat disturbance. The climate at the Kisangani weather station - taken as a proxy for Mbiye

Island - is characterized by fairly high annual precipitation (mean: 1728 mm, range: 1418-1915 mm),

12

GEMBU TUNGALUNA G.-C. et al. , Rediscovery of Glauconycteris superba Hayman, 1939

Table 4. Geographic and climatic data for the localities where Glauconycteris superba Hayman, 1939

has been collected. GLC2000 = Global Land Cover 2000 (Mayaux et al. 2004); Precipitation = annual

precipitation; Tmin = mean annual m inim um temperature; Tmax = mean annual maximum temperature

(Hijmans et al. 2005).

Locality

Coordinates

GLC2000

Elevation

[m]

Precipitation

[mm/year]

Tmin

[°C]

Tmax

[°C]

Pawa

2°31’36”N, 27°42’10”E

Closed evergreen lowland forest

775

2116

18.6

29.6

Mbiye island

0°27’50”N, 25°17’14”E

Closed evergreen lowland forest

413

1786

20.5

30.2

Oda

5°55’23”N, 00°59’42”W

Mosaic Forest / Croplands

137

1573

22.0

30.7

Matonguine

7°17’20”N, 08°02’17”W

Mosaic Forest / Croplands

356

1986

18.3

30.9

with two short dry seasons (December to February and June to July), and an average annual temperature

varying between 23.5 and 25.3°C (Nshimba 2008). The other localities are also characterized by a

humid, tropical climate and only Matonguine differs in having one dry and one wet season as well as

higher temperature fluctuations during the dry season (Fig. 8). Mean annual precipitation at the various

localities is fairly high to high and ranges between 1573 mm and 2166 mm. Compared to the two

localities in DRC, the West African localities are more seasonal, and Matonguine in particular has much

lower monthly m inim um temperatures during the dry season than the other sites (Fig. 8).

Many authors suggest that striped or spotted colouration patterns in bats improve their camouflage

when roosting in foliage, e.g. Yalden & Morris (1975) for Glauconycteris , Hall & Richards (1979) for

500

1000 km

o

Fig. 7. Distribution map of G. superba showing the four currently known records from Ivory Coast,

Ghana, and DRC; red dot: Mbiye Island. Land cover data from GLC2000 (Mayaux et al. 2004).

13

European Journal of Taxonomy 42: 1-18 (2013)

Nyctimene Borkhausen, 1797, Hill & Smith (1984) for Euderma H. Allen, 1892, Lim & Dunlop (2008)

for New World emballonurids, and Santana et al. (2011) for bats in general. Such colouration is likely

to provide protection from predators in their day roosts and probably also lowers the chances of being

detected by researchers, but this would not influence the capture success of techniques such as shooting,

mist netting or harp trapping. The low number of G. superba in museum collections might thus indicate

that it is indeed a very rare species, or that we simply do not know yet where to look for it.

In the IUCN Red List of Threatened Species, G. superba is currently ranked as “Least Concern” (LC;

Fahr et al. 2008). The species had been ranked as “Vulnerable” (VU) during the initial workshop of

the Global Mammal Assessment in 2004 but was later downgraded to “Least Concern” due to its wide

distribution, presumed large population, and because it was considered unlikely to be declining fast

enough to qualify for listing in a more threatened category. However, G. superba is either very rare or

unusually difficult to record as evidenced by the four currently known specimens, which were collected

over a period of 74 years between 1938 and 2012, with collection intervals of eight, 26 and 40 years,

respectively.

Fig. 8. Climate graphs for Pawa (upper left), Mbiye (upper right), Oda (lower left), and Matonguine

(lower right). Upper line of squares: mean monthly maximum temperature, lower line of squares: mean

monthly m inim um temperature, bars: mean monthly precipitation (plotted using DIVA-GIS, data from

Hijmans et al. 2005).

14

GEMBU TUNGALUNA G.-C. et al. , Rediscovery of Glauconycteris superba Hayman, 1939

In fact, the large intervals between previous captures and the long time elapsed since the last record

(1972) could have led to the erroneous conclusion that G. superba had become extinct. Fisher &

Blomberg (2012), who examined the extinction probability of ma mm als with a restricted number of

sighting records, reported that the interval between last sightings and estimated extinction for most

mammals was between 1 and 31 years, with a mean of 8.8 years. This interval was clearly exceeded

in G. superba prior to our rediscovery, which stresses the need for more surveys and research effort to

determine the status of elusive small mammals such as G. superba.

Although the maximum distance of ca. 4000 km between the most distant records (Matonguine and

Pawa) indicates a large extent of occurrence, the extremely low number of known localities recorded

over a long period might indicate very specific habitat requirements and possibly a very restricted area

of occupancy. Moreover, the specimens from Pawa, Oda and Matonguine originated from forest areas

that are today heavily degraded and fragmented, especially those in West Africa (Chatelain et al. 2004;

Duveiller et al. 2008), and it is currently unknown whether G. superba still occurs at, or near these

localities. Given these uncertainties, we believe that ranking this species as “Data Deficient” (DD)

is more appropriate under a precautionary principle, and that more data are required for an informed

assessment of the threat status of this species. Furthermore, available material should be sequenced to

assess whether the taxon, as currently conceived, comprises cryptic species despite the morphological

agreement of the currently known specimens that were collected over a large distribution range.

Acknowledgements

We would like to thank W im Wendelen (RMCA) for extracting the skull of the new specimen and for

taking the skull photos in the Tervuren collection, and Rene Schaerlaeken for the drawing of the head

of the Matonguine specimen. The efforts by Judith Eger (Royal Ontario Museum, Toronto) and Wim

Bergmans (Zoological Museum Amsterdam) that helped to locate the “lost” specimen from Matonguine

at the University of Antwerp are highly appreciated. We further thank Teresa Kearney and Ernest

Seamark (Ditsong National Museum of Natural History [formerly Transvaal Museum]) as well as the

Natural History Museum, London for providing the photos of the type specimen of G. superba sheila.

Thanks to Charles Balekage, Marc Bolcetshu, Prosper Kakule, Japhet Kasereka, Joseph Kussa, John

Mbusa and Charles Andabathi, students at the University of Kisangani, for their contribution to regular

sampling of bats. The LEGERA field work was supported by a CIFOR scholarship from the European

Union (REAFOR program: GCP/DRC/036/EC), for which we thank their respective directors: Dr. Nasi,

and Prof. Mate and Ndjele. Emmanuel Gilissen is thanked for his hospitality during GCGT’s stay at the

RMCA, which was supported by a travel grant from VLIR-UOS and ABIC (DGD).

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GEMBU TUNGALUNA G.-C. et al. , Rediscovery of Glauconycteris superba Hayman, 1939

Manuscript received: 5 November 2012

Manuscript accepted: 24 January 2013

Published on: 10 April 2013

Topic editor: Rudy Jocque

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of

the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National Botanic Garden

of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium; Natural History

Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium;

Natural History Museum of Denmark, Copenhagen, Denmark.

17

European Journal of Taxonomy 42: 1-18 (2013)

Appendix: Details of available Glauconycteris superba specimens

DEMOCRATIC REPUBLIC OF THE CONGO: Pawa, 30 mi ENE Medje (Hayman 1939, Hayman et

al. 1966; RMCA 14765, leg. Dr. Albert Dubois [received 8 Aug. 1938], subad ?, ale & skull, holotype

G. s. superba ); Mbiye Island (this study; RMCA al.097-M-3153, leg. Guy-Crispin Gembu Tungaluna,

collected 19 Feb. 2012, original number CHIM 012, ad S, ale & skull).

GHANA: Oda (Hayman 1947, Rosevear 1965; BMNH 47.10, leg. George Soper Cansdale, collected 6

Dec. 1946, original number 807, ad §, skin & skull, holotype G. s. sheila).

IVORY COAST: Matonguine (Peterson & Smith 1973; RMCA 97.077-M-7719, leg. Louis Bellier,

collected 24 Jan. 1972, original number A9396, ad c?, ale & skull).

18