http: //dx .doi .org/10.5852/ej t .2013.5 0
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2013 • De Meyer M. et al.
This work is licensed under a Creative Commons Attribution 3.0 License.
Research article
um:lsid:zoobank.org:pub:F550B61F-1152-47A3-8217-7AAA4AD6BCAB
Notes on the frugivorous fruit fly (Diptera: Tephritidae) fauna of
western Africa, with description of a new Dacus species
Marc DE MEYER'-^ *, Ian M. WHITE^'* & Kim F.M.GOODGER^-^
^ Royal Museum for Central Afriea, Entomology Seetion, Tervuren, Belgium
E-mail: [email protected]
^ Natural History Museum, Department of Entomology, Eondon, UK
E-mail: [email protected]
* Corresponding author: Mare De Meyer, Royal Museum for Central Afriea, Eeuvensesteenweg 13,
B-3080 Tervuren, Belgium
^ um:lsid:zoobank.org:author:29491E42-8F20-4711-B73C-55142EB2A744
" urn:lsid:zoobank.org:author:D7A0D6AC-77A3-4561-B7D5-2975DlCB3FAE
^ um:lsid:zoobank.org:author:2969C240-3148-40ED-8CFB-BClFB3AF0487
Abstract. The speeies riehness of the frugivorous fruit fly fauna of western Afriean (in partieular of
Ivory Coast, Ghana, Togo, Benin and Nigeria) is diseussed. The diversity is eompared at a national level
and between the eeoregions within the national boundaries of the study area. A new speeies, Dacus
goergeni sp. nov. is deseribed and additional taxonomie notes are presented.
Keywords. Daeini, Afrotropieal, fruit flies, Tephritidae, West Afriea.
De Meyer M., White l.M. & Goodger K.F.M. 2013. Notes on the frugivorous fruit fly (Diptera: Tephritidae) fauna
of western Africa, with description of a wq^n Dacus species. European Journal of Taxonomy 50: 1-17. http://dx.doi.
org/10.5852/eit.2013.50
Introduction
Tephritidae are pieture-winged flies of variable size and worldwide distribution. Although eommonly
named ‘fruit flies’, the larvae of some speeies develop in other parts of the host plant, ineluding flowers,
seeds and stems (White & Elson-Harris 1994). True frugivorous tephritids mainly belong to the Daeini
(sometimes referred to as subfamily Daeinae) and inelude the genera Bactrocera Maequart, Dacus
Fabrieius (Daeina), Ceratitis MeEeay, Capparimyia Bezzi, Carpophthoromyia Austen, Neoceratitis
Hendel, Perilampsis Bezzi and Trirhithrum Bezzi (Ceratitidina) (Thompson 1999), several of whieh
are of agrieultural importanee as pests of eommereial fruits and vegetables (White & Elson-Harris
1994). However, the majority of fruit fly speeies of these genera are reported from a limited number of
indigenous non-co mm ercial ifuits.
The West Afriean fauna has been extensively studied over more than a eentury. In partieular, the fauna
of Ivory Coast, Ghana, Togo, Benin and Nigeria has been more or less eontinuously researehed, with
1
European Journal of Taxonomy 50: 1-17 (2013)
expeditions dating baek to 1912 when F. Silvestrii was sent out by the Board of Agrieulture and Forestry
of the Territory of Hawaii in seareh of natural parasitoids of the Mediterranean fruit fly, Ceratitis capitata
(Wiedemann, 1824) (Silvestri 1913). In reeent deeades, researeh earried out in these eountries by staff
from the Cotonou station of the International Institute of Tropieal Agrieulture (IITA), has eontributed
eonsiderably to our knowledge of the regional tephritid fauna. This work has been eomplemented by
national fruit fly researeh aetivities in several eountries sueh as Benin (Vayssieres et al 2005), Ivory
Coast (N’depo et al. 2009), Togo (Amevoin et al. 2009) and Nigeria (Umeh et al. 2008). The seope of
our analysis is therefore restrieted to the area eonflned by the politieal boundaries of the above mentioned
eountries beeause of the relatively high number of sampling events and the sampling eontinuity in time.
Although ineomplete, it eovers a large proportion of the main terrestrial eeoregions found in this area,
as deflned by Burgess et al. (2004), in partieular the Western and Eastern Guinean Forests, the Guinean
Forest Savannah Mosaie and a representative proportion of the West Sudanian Savannah. It, therefore,
allows us to provide some preliminary data on the speeies riehness and faunal similarity between these
eeoregions. During the identifleation of material for this study, a hitherto undeseribed speeies belonging
to the genus Dacus was diseovered, whieh is also deseribed in this paper.
Material and methods
The study was based upon all available tephritid speeimens in eolleetions worldwide, plus those literature
reeords that eould be eheeked for aeeuraey. Study of the extensive eolleetion at IITA (Cotonou) proved
espeeially valuable. That eolleetion is predominantly the result of sampling aetivities by Dr G. Goergen
(IITA). Additional speeimens were available through sampling by Dr J.-F. Vayssieres of CIRAD and
attaehed to IITA. In total, 9814 speeimens, eolleeted during 1965 sampling events, were ineluded in
the study. Sampling events were not standardized or of equal intensity, but the result of oeeasional
eolleeting events within the framework of ongoing researeh aetivities. All these reeords are entered in
the fruit fly speeimen database at the Royal Museum for Central Alfiea and are also publiely available
through http://proieets.bebif be/fruitflv/index.htink ineluding information on sampling methods (lure
attraetion, sweeping, rearing) when known. All speeimens were (re-)identifled within the seope of reeent
taxonomie revisions (De Meyer 1996, 1998, 2000, 2006, 2009; De Meyer & Copeland 2001; De Meyer
Fig. 1. Country boundaries of Ivory Coast (1), Ghana (2), Togo (3), Benin (4) and Nigeria (5) with sites
of eolleeting events for Tephritidae (open eireles).
2
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
& Freidberg 2005, 2006; White 2006; White et al. 2003; White & Goodger 2009) or during subsequent
visits to collections using these recent revisions.
All collecting events were then plotted on a map representing the country boundaries (Fig. 1) and the
ecoregions (Fig. 2) using Arc View GIS (version 3.2). Geographical co-ordinates were based on GPS
readings indicated on the samples, or geo-references of particular localities checked on various GIS
databases, and are available through the above mentioned website I http://pro)ects.bebif be/fruitflv/index.
html F The respective collecting events per country and per ecoregion were calculated using selection
functions within the Arc View GIS software.
The data were analyzed, comparing the data sorted as country records and as ecoregion records both in
absolute figures and as percentages. Both methods were included because country richness estimates
allow governments to evaluate national diversity, while ecoregional estimates puts the observed richness
in a larger regional perspective with regard to biodiversity conservation. Because of differences in
sampling methodology, which does not allow to compare individuals obtained through the different
methods, only incidence based analyses were conducted. Compositional similarity was calculated by
means of the incidence based Jaccard and Sorensen indices using MSExcel according to the formulas
in Magurran & McGill (2011). Estimated species richness incidence parameters ICE and CHA02 were
calculated using Estimates (version 8.2) (Colwell 2012). For this, the different samples were divided
in 10 year periods (1891-1910, 1911-1920, etc.) resulting in 12 samples. This approach was suggested
by Meier & Dikow (2004) to use collection specimens for richness estimations, except that we used
10 year periods rather than 5 years because of the limited number of records for certain periods. As
species richness is known to have a positive relationship with area size (Burgess et al. 2004), this can
be corrected for by using a mathematical approach using the formula developed by Rosenzweig (1995):
BVd=BV/A^ with BV being the biological value in question (in our analysis the species richness), A
being the area (in square kilometers) and BVd the biological value corrected for area, and with ^ set at
0.2 as suggested by Burgess et al. (2004). This was done both for country and ecoregion richness data.
Fig. 2. Ecoregion boundaries of West Guinean Eowland Forest (1), East Guinean Forest (2), Nigerian
Eowland Forest (3), Cross-Nigerian Transition Forest (4), West Sudanian Savannah (5), Guinean Forest
Savannah Mosaic (6), Jos Plateau Forest Grassland Mosaic (7), Central African Mangroves (8), with
sites of collecting events for Tephritidae (open circles).
3
European Journal of Taxonomy 50: 1-17 (2013)
Abbreviations
IITA = International Institute for Tropieal Agrieulture, Dar es Salaam
NHM = Natural History Museum, London
RMCA = Royal Museum for Central Afriea, Tervuren
Results
Biodiversity
The total number of eolleeting events (1965) were divided over 285 point loealities (Fig. 1).
The total number of speeies found in the study area was 117 (Table 1), whieh is about 30 % of all
known Daeina and Ceratitidina speeies in the Afrotropieal region. For the genera Carpophthoromyia,
Ceratitis and Perilampsis, the pereentage was 40% or higher. The genus Neoceratitis Hendel was absent
from western Afriea, and the genera Capparimyia and Bactrocera were poorly represented. The speeies
Table 1. List of Tephritidae reeorded from the different eountries and eeoregions in the study area,
(abbreviations: WGLL = West Guinean Lowland Forest; EGF = East Guinean Forest; NEF = Nigerian
Eowland Forest; CNTF = Cross-Nigerian Transition Forest); WSS = West Sudanian Savannah; GFSM =
Guinean Forest Savannah Mosaie; JOS = Jos Plateau Forest Grassland Mosaie; CAM = Central Afriean
Mangroves).
Genus
Species
Ghana
Togo
Benin
Nigeria
Ivory Coast
WGLL
EGF
NLF
CNTF
WSS
GFSM
JOS
CAM
Bactrocera
cucurbitae
X
X
X
X
X
X
X
X
X
X
X
Bactrocera
invadens
X
X
X
X
X
X
X
X
X
X
X
X
Bactrocera
mesomelas
X
X
X
X
X
X
X
X
X
X
Capparimyia
melanaspis
X
X
Carpophthoromyia
dividua
X
X
X
X
Carpophthoromyia
interrupta
X
X
X
Carpophthoromyia
nigribasis
X
X
Carpophthoromyia
pseudotritea
X
X
X
X
X
X
X
X
X
X
Carpophthoromyia
scutellata
X
X
X
X
Carpophthoromyia
tritea
X
X
Carpophthoromyia
vittata
X
X
X
X
X
Ceratitis
acicularis
X
X
Ceratitis
anonae
X
X
X
X
X
X
X
X
X
X
Ceratitis
antistictica
X
X
X
X
X
X
X
Ceratitis
barbata
X
X
X
X
X
Ceratitis
bicincta
X
X
X
X
X
Ceratitis
bremii
X
X
X
X
X
X
X
X
X
X
Ceratitis
capitata
X
X
X
X
X
X
X
X
X
Ceratitis
colae
X
X
X
X
X
X
X
X
X
X
Ceratitis
connexa
X
X
Ceratitis
cosyra
X
X
X
X
X
X
X
X
X
X
Ceratitis
discussa
X
X
Ceratitis
ditissima
X
X
X
X
X
X
X
X
X
X
X
Ceratitis
dumeti
X
X
4
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
Genus
Species
Ghana
Togo
Benin
Nigeria
Ivory Coast
WGLL
EGF
NLF
CNTF
WSS
GFSM
JOS
CAM
Ceratitis
fasciventris
X
X
X
X
X
X
X
X
X
Ceratitis
flava
X
X
X
X
X
X
Ceratitis
flexuosa
X
X
X
X
X
X
X
X
X
Ceratitis
grahami
X
X
Ceratitis
guttiformis
X
X
X
Ceratitis
hamata
X
X
Ceratitis
lentigera
X
X
X
X
X
X
X
X
Ceratitis
lepida
X
X
Ceratitis
lineata
X
X
Ceratitis
lunata
X
X
Ceratitis
melanopus
X
X
Ceratitis
morstatti
X
X
X
X
X
X
Ceratitis
neostictica
X
X
Ceratitis
paracolae
X
X
Ceratitis
pedestris
X
X
X
X
Ceratitis
penicillata
X
X
X
X
X
X
X
X
X
X
Ceratitis
punctata
X
X
X
X
X
X
X
X
X
Ceratitis
quinaria
X
X
X
Ceratitis
semipunctata
X
X
Ceratitis
silvestrii
X
X
X
X
X
Ceratitis
stictica
X
X
X
X
Ceratitis
striatella
X
X
X
X
X
X
X
X
Ceratitis
tripteris
X
X
X
X
Ceratitis
whitei
X
X
Dacus
acutus
X
X
X
Dacus
albiseta
X
X
Dacus
annulatus
X
X
X
X
X
X
X
Dacus
armatus
X
X
X
X
X
X
X
X
X
X
X
Dacus
aspilus
X
X
Dacus
bakingiliensis
X
X
Dacus
binotatus
X
X
Dacus
bivittatus
X
X
X
X
X
X
X
X
X
X
X
X
Dacus
botianus
X
X
X
X
X
Dacus
earnest
X
X
X
X
X
X
X
X
X
Dacus
ceropegiae
X
X
Dacus
chapini
X
X
X
X
Dacus
ciliatus
X
X
X
X
X
X
X
X
X
X
Dacus
collarti
X
X
X
Dacus
congoensis
X
X
X
X
Dacus
croceus
X
X
Dacus
diastatus
X
X
X
X
X
X
X
X
X
X
Dacus
disjunctus
X
X
X
X
X
X
X
X
Dacus
elutissimus
X
X
Dacus
fasciolatus
X
X
Dacus
Jiavicrus
X
X
X
X
X
X
X
Dacus
frontalis
X
X
Dacus
fuscovitttatus
X
X
X
X
X
X
X
5
European Journal of Taxonomy 50: 1-17 (2013)
Genus
Species
Ghana
Togo
Benin
Nigeria
Ivory Coast
WGLL
EGF
NLF
CNTF
WSS
GFSM
JOS
CAM
Dacus
gabonensis
X
X
X
X
Dacus
goergeni sp. nov.
X
X
X
X
Dacus
guineensis
X
X
Dacus
gypsoides
X
X
Dacus
hamatus
X
X
X
X
X
X
Dacus
humeralis
X
X
X
X
X
X
X
X
X
X
Dacus
inflatus
X
X
Dacus
inornatus
X
X
X
X
Dacus
kurrensis
X
X
Dacus
langi
X
X
X
X
X
X
X
Dacus
limbipennis
X
X
X
X
X
X
X
X
X
Dacus
longistylus
X
X
X
X
Dacus
lounsburyii
X
X
Dacus
maynei
X
X
Dacus
mediovittatus
X
X
X
X
X
Dacus
parvimaculatus
X
X
X
X
Dacus
plagiatus
X
X
Dacus
pleuralis
X
X
X
X
X
X
X
Dacus
punctatifrons
X
X
X
X
X
X
X
X
X
X
Dacus
schoutedeni
X
X
Dacus
serratus
X
X
Dacus
setilatens
X
X
Dacus
goergeni sp.nov.
X
X
X
X
Dacus
theophrastus
X
X
X
X
X
X
X
X
X
X
Dacus
transitorius
X
X
X
X
X
Dacus
trigonus
X
X
X
X
Dacus
umehi
X
X
X
X
Dacus
vertebratus
X
X
X
X
X
X
X
X
X
X
X
Dacus
xanthinus
X
X
Perilampsis
atra
X
X
X
X
X
X
X
X
X
Perilampsis
decellei
X
X
X
X
Perilampsis
deemingi
X
X
Perilampsis
formosula
X
X
Perilampsis
furcata
X
X
X
X
X
X
X
Perilampsis
pulchella
X
X
X
X
X
Perilampsis
woodi
X
X
X
X
X
X
X
Trirhithrum
basale
X
X
Trirhithrum
brachypterum
X
X
Trirhithrum
coffeae
X
X
X
X
X
X
X
X
X
Trirhithrum
dimorphum
X
X
X
X
X
Trirhithrum
homogeneum
X
X
Trirhithrum
inscriptum
X
X
X
X
Trirhithrum
leonense
X
X
X
X
Trirhithrum
nigerrimum
X
X
X
X
X
X
X
X
X
Trirhithrum
nigrum
X
X
X
X
X
X
Trirhithrum
obscurum
X
X
X
X
X
X
X
X
6
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
Total/area
Total
Trirhithrum
Perilampsis
Dacus
Ceratitis
Carpophthoromyia
Capparimyia
Bactrocera
Ui
o
o\
Ui
-
o
-
WGLL
ooot
oo
o
o
o
64.0
o
o
-p
o
o
o
-p
o
WGLL%
On
On
Ui
oo
Ui
Ui
o
U)
EGF
ooot
to.7
On
33.3
36.0
OJ
o
o
-p
o
EGF%
On
1/1
On
O
On
o\
On
o
OJ
NEE
ooot
oot
oot
43.3
oo
OJ
U)
o
o
Ui
o
NEF%
p
1/1
U)
o
o
o
o
o
-
CNTF
ooot
o
o
o
o
66.7
o
o
o
o
o
o
33.3
CNTF%
p
Ui
Ui
-p
On
o
wss
ooot
OJ
On
U)
47.3
34.5
OJ
On
o
o
OJ
On
wss%
U)
Ui
CO
K)
o
-P
-
U)
GFSM
ooot
t2.1
OJ
■-P
34.5
36.2
On
Ui
GFSM%
oo
o
-
o
ui
O
o
JOS
ooot
o
o
Ui
On
55.6
p
bo
O
O
o
o
tl.t
J0S%
U)
bo
Ui
On
U)
O
o
CAM
ooot
t3.3
t3.3
40.0
o
o
O
O
o
o
t3.3
CAM%
o
ui
OJ
oo
-
OJ
AFF
ooot
oo
1/1
On
O
43.6
32.5
o\
o
o
On
AFF%
o
>
P.
7
European Journal of Taxonomy 50: 1-17 (2013)
accumulation curve, however, did not show any leveling off (Fig. 3), and both ICE and CHA02 indieated
a higher estimated speeies riehness (mean ICE 168.44; mean CHA02 153.3).
When the fauna of the different eeoregions was eompared (Table 2), the highest speeies riehness was
found in the East Guinean Forest (75 speeies) and the Nigerian Fowl and Forest (60 speeies), followed
by the Guinean Forest Savannah Mosaie (58) and the West Sudanian Savannah (55). The lowest speeies
riehness was found in the Cross-Nigerian Transition Forest (4). The same tendeneies were observed
after eorreetion for area size. The West Sudanian Savannah was espeeially rieh in Dacus speeies but
Table 3. Ineidenee based similarity indiees (a: Jaeeard; b: Sorensen) for Tephritidae between the
eeoregions in the study area (abbreviations: see Table 2) (range 0-1, with 0 being no similarity and 1
being eomplete similarity).
a) Jaeeard
JOS
WSS
GFSM
WGEE
NEF
EGF
CNTF
CAM
0.10
0.13
0.20
0.18
0.19
0.17
0.19
JOS
0.22
0.10
0
0.15
0.11
0.16
WSS
0.42
0.23
0.43
0.42
0.07
GFSM
0.26
0.48
0.60
0.07
WGEE
0.27
0.27
0.04
NEF
0.48
0.07
EGF
0.05
b) Sorensen
JOS
WSS
GFSM
WGEE
NEF
EGF
CNTF
CAM
0.18
0.23
0.33
0.3
0.32
0.29
0.32
JOS
0.36
0.18
0
0.26
0.19
0.27
WSS
0.59
0.38
0.60
0.59
0.14
GFSM
0.41
0.64
0.75
0.13
WGEE
0.42
0.42
0.07
NEF
0.65
0.13
EGF
0.10
Fig. 3. Speeies aeeumulation eurve for tephritid speeies reeorded from study area.
8
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
very poor in Trirhithrum species (in particular when expressed as a percentage of the total fruit fly
fauna). On the other hand, the Guinean Forest Savannah Mosaic and East Guinean Forest were very
rich in Trirhithrum. The Beta-diversity was very heterogenous (Table 3) with especially Central African
Mangroves, Jos Plateau Forest Grassland Mosaic, and Cross-Nigerian Transition Forest showing low
similarity with the other systems. Highest similarity was found between the East Guinean Forest and
the Guinean Forest Savannah Mosaic. For some of the ecoregions that are present within the political
boundaries of the area studied, no collecting records were available. This was the case for the Niger
Delta Swamp Forests, the Cross-Niger Transition Forests, the Cross-Sanaga-Bioko Coastal Forests, the
Cameroon Highlands Forests (all in Nigeria) and the Guinean Montane Forests (in Ivory Coast).
When the fauna within the different political boundaries was compared (Table 4), the highest species
richness was found in Nigeria (77 species) while all other countries had a similar richness (52-55
Table 4. Species richness by genus for Tephritidae in the different countries within the study area.
Ivory Coast
Ivory Coast%
Ghana
Ghana%
Togo
Togo%
Benin
Benin%
Nigeria
Nigeria%
AEE
AEE%
Bactrocera
3
5.8
3
5.8
3
5.8
3
5.5
3
3.9
3
2.6
Capparimyia
0
0.0
0
0.0
0
0.0
1
1.8
0
0.0
1
0.9
Carpophthoromyia
4
7.7
5
9.6
1
1.9
1
1.8
3
3.9
7
6.0
Ceratitis
20
38.5
23
44.2
16
30.8
20
36.4
21
27.3
38
32.5
Dacus
18
34.6
12
23.1
25
48.1
27
49.1
36
46.8
51
43.6
Perilampsis
4
7.7
1
1.9
2
3.8
2
3.6
7
9.1
7
6.0
Trirhithrum
3
5.8
8
15.4
5
9.6
1
1.8
7
9.1
10
8.5
Total
52
100.0
52
100.0
52
100.0
55
100.0
77
100.0
117
100.0
Total/area
4.1
4.4
5.8
5.3
4.9
Table 5. Incidence based similarity indices (a: Jaccard; b: Sorensen) for Tephritidae between the
countries in the study area. Range 0-1, with 0 being no similarity and 1 being complete similarity).
a) Jaccard
Benin
Togo
Ghana
Ivory Coast
Nigeria
0.43
0.47
0.40
0.39
Benin
0.53
0.37
0.41
Togo
0.51
0.49
Ghana
0.41
b) Sorensen
Benin
Togo
Ghana
Ivory Coast
Nigeria
0.61
0.64
0.57
0.56
Benin
0.69
0.54
0.58
Togo
0.67
0.65
Ghana
0.58
9
European Journal of Taxonomy 50: 1-17 (2013)
species). After correction for country area, the highest richness was observed in Togo and Benin. The
Beta-diversity was rather similar between all countries (Table 5) with Jaccard indices varying between
0.37 and 0.51 and Sorensen indices between 0.54 and 0.69. The highest similarity was between the fauna
of Benin and Togo while the lowest similarity was found between Benin and Ghana. Second lowest
similarity was between Nigeria and Ivory Coast.
Taxonomy
Phylum Arthropoda Siebold, 1848
Classis Insecta Linnaeus, 1758
Ordo Diptera Linnaeus, 1758
Familia Tephritidae Macquart, 1835
Genus Fabricius, 1805
Among the material, one Dacus species, new to science, was discovered. It is hereby described.
Dacus (Psilodacus) goergeni sp. nov.
um:lsid:zoobank.org:act:47FlAD2E-9183-4C0F-A65B-D9FDBB15DBlD
Fig. 4
Etymology
Named in honour of the collector. Dr Georg Goergen, who is also the founder and conservator of the
entomological collections at the International Institute of Agriculture.
Material
Holotype
(f, TOGO, Kioto, Jan. 2006, ‘on Solanum sp.’, leg. G. Goergen (deposited in collection of IITA).
Paratypes
TOGO: same locality as holotype, 1 S, Dec. 2005, 'on Acacia auriculiformis'; 2 SS, 5 $ $, Jan. 2006;
1 (?, 3 $$, Jan. 2006, 'on Solanum sp.’; 4 SS, 1 ?, Jan. 2008; 1 1 $, Feb. 2008. BENIN: Eokossa,
1 (?, 1 ?, Jan. 2006. All leg. G. Goergen. Paratypes deposited in collections of IITA, NHM and RMCA.
Description
Size. 6.2-7.5 mm, wing length 4.8-6.6 mm.
Head. Pedicel and 1st flagellomere not longer than ptilinal suture. Face (Fig. 4A): antennal furrow
without a dark spot; upper area with a dark marking, tending to an inverted V-shaped dark marking (in
some specimens this extends down each side of carina and may be mistaken for facial spots). Frons:
frontal setae 0, orbital seta 0.
Thorax. Scutum (Fig. 4B) predominantly fuscous, tending to red-brown antero-laterally; postpronotal
lobe entirely pale, yellowish; notopleural callus pale posteriorly, anteriorly concolorous with scutum;
notopleural xanthine probably isolated from notopleural callus but can appear almost joined (as in
wedge form); lateral and medial postsutural vitta absent. Scutellum without any dark patterning (except
for basal dark margin, which is very narrow). Anepistemum (Fig. 4C) with a stripe from notopleural
callus to (or almost to) katepistemum; stripe very broad (anteriorly extending almost to postpronotal
lobe); extended onto katepistemum. Eaterotergal xanthine confined to katatergite.
Thoracic setae. Anterior notopleural seta present; anterior supra-alar seta present.
10
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
Fig. 4. Dacus goergeni sp. nov. A. Head, frontal view. B. Thorax, dorsal view. C. Thorax, lateral view.
D. Abdomen, dorsal view. E. Wing.
11
European Journal of Taxonomy 50: 1-17 (2013)
Wing (Fig. 4E). Basal cell be without microtrichia; cell c with an almost complete (> 90%) covering
of microtrichia; cell bm without microtrichia. Narrow subbasal raised section of cell br with extensive
covering of microtrichia; partly bare in apical half Crossvein R-M beyond middle of cell dm. Costal band
complete; shallow, not or barely extending below vein R 2 ^ 3 , except in basal section (before crossvein
R-M) and at wing apex; expanded into a small spot at apex. Anal streak absent (but with a trace of colour
confined to cell bcu). Cells be and c coloured (not as deep as costal band). Without any crossbanding.
Legs. Forefemur pale, yellowish, sometimes indistinctly darkened apically; midfemur bicoloured (pale
basal half to two-thirds, red-brown apically); hindfemur pale, yellow, rarely distinctly darkened apically.
Abdomen. Predominantly fuscous; shape and patterning, see Fig. 4D. Tergites 1-V all fused.
Male
Tergite 111 with some very fine hairs (possible vestigial pecten); lacking hindtibia preapical “pad”. Basal
costal sections without specialised setae.
Female
Aculeus pointed, similar to B. stylifer.
Host
No host records known (some material is indicated as being found on Solanum sp. ox Acacia auriculiformis
but there is no indication that either of these plants is a host).
Distribution
Reported from Benin and Togo.
Remarks
The new species is very similar to Dacus stylifer which is an East African species. It differs in the wing
cell c having almost complete coverage of microtrichia in the males (50% in male stylifer)-, the mid¬
femur bicoloured (pale in typical stylifer)-, the notopleuron bicoloured and sutural xanthine distinct,
unlike typical stylifer. Dacus goergeni sp. nov. is placed in the ill-defined subgenus Psilodacus, based
upon a combination of characters, which typify the grouping, including the lack of facial spots: the dark,
almost inverted V-shaped, dark marking at the top of the face; lack of anal streak and male pectin; it
differs from most Psilodacus spp. in having anterior supra-alar setae. The type specimens were captured
in the Guinean Forest Savannah Mosaic and the Eastern Guinean Forest ecoregions. Label information
indicates that the specimens were collected in forested areas.
Other taxonomic notes
Dacus pleuralis Coll art, 1935
Among the material examined was a male specimen collected at Ibadan, Nigeria (4-8 Dec. 2003, cue
lure traps, leg. G. Goergen), in addition to the male specimen studied earlier (White & Goodger 2009). It
shows some morphological deviation from the earlier collected specimen and from the type material as
redescribed in White (2006): the xanthines (katatergite and anatergite) are fused, while in the typical D.
pleuralis the xanthines are clearly separated. The anterior supra-alar seta is absent or vestigial while well
developed in the typical D. pleuralis. These differences, however, appear to be intraspecific variation
and do not warrant separate description.
12
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
Dacus mocha Bqzzi, 1917
Dacus mocha was described from Eritrea, but the type specimens were lost. White (2006) placed it in
synonymy with D. annulatus Becker, based on the similarity of the original description. Subsequently,
White & Goodger (2009) reported a specimen from Ethiopia, which was clearly not D. annulatus, but
was a good match to the description of D. mocha, which they then removed from synonymy. Amongst
the material examined here, there was a male specimen from Kioto, Togo (Mar. 2006, leg. G. Goergen),
which is very similar to the D. mocha from Ethiopia, except that it has pallid face spots. Togo is a
considerable westward expansion of the known distribution of a species otherwise known only from
a restricted area of East Africa. Since it concerns a single specimen whose identity is uncertain, we
excluded it from the richness analysis.
Dacus blepharogaster Bqzzi, 1917
A male specimen from Serou, Benin (Dec. 2005, leg. G. Goergen) differs from D. blepharogaster, as
described by White (2006) in having some red pattern on the third abdominal tergite. However, since
this species belongs to a group that needs proper revision {Dacus (Lophodacus) brevis group as defined
by White 2006) and since only one specimen was found in the collections studied, it is not described as
a separate species. As the previous species, this is also predominantly an East African species (Kenya,
Eritrea and Ethiopia) but was not included in the richness analysis.
Discussion
In general, the frugivorous tephritid fauna of western Africa is very rich, with about 30% of all known
species found in Sub-Saharan Africa. However, the estimated species richness indices indicate that
the actual number of species can be considerably higher and, thus, the present knowledge should be
considered preliminary. A number of species appear to be endemic as they have until now only been
recorded from the study area: Ceratitis grahami Mumo, 1935, C. guttiformis Munro, 1935, C. lepida
(Munro, 1969), C. neostictica De Meyer, 1998 (all Ghana), Dacus albiseta White & Goodger, 2009
(Benin), D. acutus White & Goodger, 2009 and D. kurrensis White & Goodger, 2009 (both Nigeria). In
addition, a number of species that appear to be endemic for the larger area of West Africa, are present in
the study area: Carpophthoromyia tritea (Walker, 1849) (recorded from Ivory Coast and Sierra Eeone);
Ceratitis lunata Mumo, 1935 (Sierra Eeone and Benin); C. paracolae De Meyer & Freidberg, 2006
(Nigeria and Cameroon); C. tripteris (Munro, 1957) (Sierra Eeone, Ivory Coast and Nigeria); Dacus
carnesi (Mumo, 1984) (Benin, Ivory Coast, Ghana, Eiberia, Nigeria and Togo); D. elutissimus Bezzi,
1924 (Senegal and Togo); D.flavicrus Graham, 1910 (Ivory Coast, Eiberia, Nigeria, Sierra Eeone and
Togo); D. guineensis Hering, 1944 (Benin, Guinea and Senegal); D. umehi White, 2006 (Benin, Guinea
and Nigeria); D. xanthinus White & Goodger, 2009 (Nigeria and Senegal); Perilampsis atra Mumo,
1969 (Benin, Cameroon, Ghana, Ivory Coast, and Nigeria); P. decellei Munro, 1969 (Cameroon, Ivory
Coast and Nigeria); P. furcata Mumo, 1969 (Benin, Chad, Ivory Coast, and Nigeria); and Trirhithrum
dimorphum Mumo, 1934 (Ghana, Nigeria, and Sierra Eeone).
Remarkable is the presence of a number of species that was hitherto only known from eastern and/or
southern Africa but that are now reported for the first time from Western Africa, which results in a large
westward expansion of their known distribution range. Such cases are Capparimyia melanaspis (Bezzi,
1920) (the only representative of this genus which was only known from sub-Saharan eastern and southern
Africa, see De Meyer & Freidberg (2005)), Ceratitis discussa Mumo, 1935, and Dacus lounsburyii
Coquillett, 1901. This pattern had already been observed fox Dacus botianus (Mumo, 1984), D. ceropegiae
(Mumo, 1984), D. plagiatus Collart, 1935, D. serratus (Mumo, 1984) and Perilampsis woodi (Bezzi,
1917) (White 2006; De Meyer 2009) and the aberrant specimens of D. blepharogaster and D. mochii {cf.
supra) might also confirm this pattern once their identity is confirmed.
13
European Journal of Taxonomy 50: 1-17 (2013)
When looking at Beta-diversity indiees, the low overall similarity of the Central Afriean Mangroves,
the Jos Plateau Forest Grassland Mosaie and the Cross-Nigerian Transition Forest eeoregions is
probably a refleetion of the relatively low ric hn ess of these regions in eomparison with the others. A
plausible possibility for differenees between the other eeoregions eould be eaused by the different fruit
fly / host plant assoeiations. Ceratitis, Carpophthoromyia and Trirhithrum speeies, for example, are
mainly infesters of fleshy tree or shrub fruits (e.g., from Anaeardiaeeae, Annonaeeae, Euphorbiaeeae,
Rubiaeeae, Rutaeeae or Sapotaeeae, see De Meyer et al. 2002 and White et al. 2003), representatives
of whieh are predominantly found in forested areas. Dacus speeies, on the other hand, are infesters of
ereepers and elimbers of either Cueurbitaeeae, aselepiadApoeynaeeae, or Passifloraeeae (White 2006).
The West Sudanian Savannah is relatively rieh in Dacus speeies while low in Trirhithrum speeies. The
West Guinean Lowland Forest, Nigerian Lowland Forest, East Guinean Forest and Guinean Forest
Savannah Mosaie are relatively rieh in Trirhithrum, and the latter two are also low in Dacus and
Carpophthoromyia riehness. However, the speeies eomposition of the West Guinean Lowland Forest
(relatively rieh in Dacus speeies) does not eorrespond with this general tendeney. Haneoek (1989)
already indieated that eertain speeies groups within Dacus show a marked preferenee for forest and
moist woodland habitats in Southern Afriea, appearing only oeeasionally in drier woodlands, while
other groups do not oeeur in forests. It eould well be that a similar division is observed here, leading to
the diserepaney in some regions. However, this requires further investigation, in partieular with regard
to host plant data.
As for national differenees, it is not surprising that the highest similarity is found between neighbouring
eountries, (Benin and Togo) while the seeond lowest similarity is found between Nigeria and Ivory
Coast, whieh are the two eountries at the extremes of the study area. The lowest similarity, whieh is
found between Benin and Ghana, appears to be the result of Benin having a eonsiderably higher number
of Dacus speeies not present in Ghana, while Ghana is relatively rieh in Ceratitis, Carpophthoromyia
and Trirhithrum speeies. Again, this eould be the result of the different host plant preferenees and the
relative proportional representation of the different eeoregions in these two eountries, with the forested
areas in Be nin being very restrieted and mueh more widespread in Ghana while Benin is predominantly
oeeupied by the West Sudanian Savannah. It must be emphasized though, that this study is not based
upon a standardized sampling program with identieal sampling intensity and methodology. This ean
have an impaet on and presents a weakness of the Beta-diversity analysis both at eeoregional and eountry
level. Although a regular sampling protoeol at frequent intervals in all areas eould provide more reliable
information, given the magnitude of the study area, the held eonditions, and the life history of the flies,
this is eurrently not eonsidered a realistie approaeh.
The Conservation Status Index for all eeoregions in the study area for whieh fruit fly data are available
is eonsidered either Critieal (East Guinean Forest, Nigerian Lowland Forest, Cross-Nigerian Transition
Forest, Jos Plateau), Endangered (Guinean Forest Savannah Mosaie, Central Afriean Mangroves) or
Vulnerable (West Guinean Lowland Forest, West Sudanian Savannah) (Burgess et al 2004). The two
regions with highest speeies riehness are among the Critieal ones: East Guinean Forest and Nigerian
Lowland Forest. In the last deeade two invasive speeies of Asian origin have been reeorded tfom western
Afriea: Bactrocera invadens Drew, Tsuruta & White, 2005 (Drew et al. 2005; Goergen et al. 2011) and
B. cucurbitae Coquillett 1899 (Vayssieres et al. 2007). Both, partieularly the former, are known to show
eompetitive displaeement of indigenous speeies (Ekesi et al. 2009, Mwatawala et al. 2009), and are
now found in all eountries and eeoregions of the study area. B. cucurbitae is also known to be a strong
invader (Mwatawala et al. 2010) and is now found mainly in the Guinean Forest Savannah Mosaie and
West Sudanian Savannah. Although the majority of the existing data refer to eeonomieally signifleant
erops and speeies, it is likely that sueh invasive speeies will also have an impaet on the global indigenous
fruit fly diversity.
14
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
Acknowledgements
The first author would like to thank FWO Vlaanderen for a travel grant that enabled him to study the
collections at the International Institute for Tropical Agriculture. Many thanks to Dr G. Goergen for his
hospitality and assistance during this visit, and to Dr M. Virgilio for his assistance in producing the maps
for this paper.
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16
DE MEYER M., WHITE EM. & GOODGER KE.M., Emgivorous fruit flies of western Africa
Manuscript received: 18 January 2013
Manuscript accepted: 31 May 2013
Published on: 18 July 2013
Topic editor: Koen Martens
Desk editor: Kristiaan Hoedemakers
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