European Journal of Taxonomy 233: 1-23
http://dx.doi.org/10.5852/ejt.2016.233
BY
This work is licensed under a Creative Commons Attribution 3.0 License.
ISSN 2118-9773
www. europeanj ournaloftaxonomy. eu
2016 • De Meyer M. et at.
Research article
urn:lsid:zoobank.org:pub:150B9DC4-DlA2-49BB-A608-B7D89950FD65
Description of new Ceratitis species (Diptera: Tephritidae) from Africa, or
how morphological and DNA data are complementary in discovering
unknown species and matching sexes
Marc DE MEYER 1 *, Maulid MWATAWALA 2 , Robert S. COPELAND 3 &
Massimiliano VIRGILIO 4
14 Royal Museum for Central Africa, Department Biology & JEMU, Tervuren, Belgium.
2 Sokoine University of Agriculture, Department of Crop Science and Production, Morogoro, Tanzania.
3 International Centre of Insect Physiology and Ecology, Nairobi, Kenya.
* Corresponding author: [email protected]
2 Email: [email protected]
3 Email: rcopeland@ icipe.org
4 Email: [email protected]
1 urn:lsid:zoobank.org:author:2F07B9E7-3545-49DE-8C72-55E8ED95A455
2 urn:lsid:zoobank.org:author:F61E74AB-FABD-4347-A125-64C5C5F7F241
3 urn:lsid:zoobank.org:author:F2F7C761-2E36-4DAB-9EDF-Q91220DA8A88
4 urn: lsid: zoobank, org: author: A4C7D3FF -E127-4645-9BFD-78222520FE26
Abstract. This paper describes five new Ceratitis species from the eastern and southern parts of the
Alf otropical Region: C. (Pterandrus ) quilicii De Meyer, Mwatawala & Virgilio sp. nov.; C. ( Ceratalaspis )
pallidula De Meyer, Mwatawala & Virgilio sp. nov.; C. ( Ceratalaspis ) taitaensis De Meyer & Copeland
sp. nov.; C. ( Ceratalaspis ) sawahilensis De Meyer & Virgilio sp. nov.; and C. ( Ceratalaspis ) fiavipennata
De Meyer & Virgilio sp. nov. Their relationships with closely allied species within their respective
subgenera are discussed where appropriate, and diagnostic characters are given. DNA barcodes are
provided for all new species. In addition, the hitherto unknown male of C. (Pardalaspis) serrata De
Meyer, 1996 is described, based on material collected in the Democratic Republic of Congo. Recognition
of these new species and sexes is the result of an integrative approach using morphological characters
and DNA data.
Keywords. Fruit flies, Afrotropical, DNA barcoding.
De Meyer M., Mwatawala M., Copeland R.S. & Virgilio M. 2016. Description of new Ceratitis species (Diptera:
Tephritidae) from Africa, or how morphological and DNA data are complementary in discovering unknown species
and matching sexes. European Journal of Taxonomy 233: 1-23. http://dx.doi.org/10.5852/ejt.2016.233
Introduction
Tephritidae are picture-winged flies of variable size and worldwide distribution. Although commonly
named “fruit flies”, larval development can also take place in other parts of the host plants, including
1
European Journal of Taxonomy 233 : 1-23 ( 2016 )
flowers, seeds, leaves and stems. Fruit flies of economic significance were treated by White & Elson-
Harris (1994), and the currently acknowledged classification was presented by Norrbom etal. (1999). The
genus Ceratitis MacLeay, 1829 belongs to the tribe Ceratitidini, which is predominantly an Afrotropical
group and comprises 95 described species. The larvae of Ceratitis develop in fruit, and several species
of agricultural importance are known from this genus including the Mediterranean fruit fly ( Ceratitis
capitata (Wiedemann, 1824)), and the Natal fruit fly ( Ceratitis rosa Karsch, 1887). The taxonomy of the
different subgenera within the genus Ceratitis has been revised in the last two decades (De Meyer 1996,
1998, 2000; De Meyer & Copeland 2001; De Meyer & Freidberg 2006), and additional descriptions
of species were presented after the publication of these revisions (De Meyer & Copeland 2005, 2009).
Virgilio et al. (2014) provided an electronic identification key for all known species, as well as for
representatives of related genera. The key was recently updated to include the newly described Ceratitis
species (available from http://fruitflvkevs.africamuseum .be ).
Material and methods
The material included in this paper has been studied within the framework of a number of recent
fruit fly related projects, including surveying activities in different parts of Africa and integrated pest
management (IPM) related collaborative projects in Tanzania. During these studies a number of hitherto
unknown species were encountered. A case in particular is the description of C. quilicii sp. nov. This
is the result of an integrative taxonomic study on a number of cryptic species complexes of economic
significance (Hendrichs et al. 2015), including the Ceratitis FAR complex (Barr & Wiegmann 2009;
Virgilio et al 2008).
Material from the following collections was studied:
ICIPE
RMCA
SANC
sus
TAU
ZMUC
International Centre of Insect Physiology and Ecology, Nairobi, Kenya
Royal Museum for Central Africa, Tervuren, Belgium
South African National Collections, Pretoria, South Africa
Stellenbosch University, Stellenbosch, South Africa
Tel Aviv University, Tel Aviv, Israel
Zoological Museum, University of Copenhagen, Copenhagen, Denmark
In the species descriptions, terminology follows the glossary of White et al. (1999). Wing and body
(without oviscape for females) length were taken for ten specimens unless less were available, and are
given in mm (average and range). Measurements of both sexes were pooled (where applicable) since no
sexual difference was noted. Images of body parts were taken using a digital camera mounted on a Leica
MZ12 dissecting microscope and using Automontage software (Syncroscopy, UK). Female terminalia
were dissected, cleared in potassium hydroxide solution and temporarily mounted in glycerine-
gelatine for drawings made by camera lucida. A set of DNA sequences were produced for part of the
mitochondrial Cytochrome c oxidase subunit I (COI) and deposited in the Barcode of Life Data Systems
(BOLD, http://www.boldsvstems.org ) as reference DNA barcodes for the molecular identification of the
new species (see Virgilio et al. 2012 for laboratory protocols and discussion). Kimura’s two parameter
genetic distances (Kimura 1980) between the newly generated Ceratitis DNA barcodes and a selection
of 367 reference DNA barcodes deposited in BOLD (corresponding to 51 species from all six Ceratitis
subgenera) were visualized in a neighbor joining tree. In addition to the above mentioned depositories,
the following acronyms are used to indicate institutions where type material is deposited:
BMNH = Natural History Museum (London, UK)
NMK = National Museums of Kenya (Nairobi, Kenya)
SUA = Sokoine University of Agriculture, Department of Crop Science and Production
(Morogoro,Tanzania)
USNM = National Museum of Natural History, Smithsonian Institution (Washington DC, USA)
2
De Meyer M. et al ., New Ceratitis species (Diptera: Tephritidae) from Africa
Results
Phylum Arthropoda Siebold, 1884
Class Insecta Linnaeus, 1758
Order Diptera Linnaeus, 1758
Family Tephritidae Macquart, 1835
Subfamily Dacinae Schiner, 1864
Tribe Ceratitidini Bezzi, 1910
Genus Ceratitis MacLeay, 1829
Ceratitis (Pterandrus) quilicii De Meyer, Mwatawala & Virgilio sp. nov.
um:lsid:zoobank.org:act:FE85EC42-12F6-4C76-80C2-CA8A7C4E7D5D
Fig. 1
Etymology
Named in honour of the late Dr. Serge Quilici (Centre de Cooperation Internationale en Recherche
Agronomique pour le Developpement, CIRAD, La Reunion) who passed away in 2015. The species
name should be treated as a noun in the genitive case.
Material examined
Holotype
TANZANIA: 3, Nyandira, EGOlure trap, 4 May 2013, M. Mwatawala (RMCA coll Nr T19315).
Paratypes (deposited in BMNH, NMK, RMCA, SANC, SUA, and USNM)
TANZANIA: same locality as holotype: 2 33, 4 May 2013 (coll Nr T19315); l 3,1 Jun. 2013, coll Nr
T19471; 2 33, 1 Jun. 2013 (coll Nr T19474); 16 33 (1 $ barcoded RMCA, AB42864782, see Table 1),
combined lures, 2006, M. Mwatawala; 33 33, Mgeta Visada, EGOlure trap, 4 May-29 Jun. 2013, M.
Mwatawala.
KENYA: 2 S3, Kirimiri Forest, reared ex fruits Englerophytum natalense, 23 Jan. 2002, coll Nrl683;
1 3, Kirimiri Forest, reared ex fruits Englerophytum natalense , 21 Jan. 2003, coll Nr 2380, all R.S.
Copeland.
Non-type material
BOTSWANA: Gaborone.
KENYA: Embu-Runyenjes; Kirimiri Forest; Taita Hills, near Ngangao Forest.
MALAWI: Bvumbwe Research Station; Kumbali; Zomba.
REUNION ISLAND: St. Pierre.
SOUTH AFRICA: Addo; Amoldton; Baltimore; Bavaria; Bloemfontein; Bonza Bay; Britstown;
Burgershall; Cato Manor; Cedara; Citrusdal; Clanwilliam; Doreen Clark Nature Reserve; Duivelskloof;
Dukudulcu; Durban; East London; Enon Farm, near Richmond; Eshowe; Ferncilff Nature Reserve;
Fort Beaufort; Gariepdam; Grahamstown; Haenertsburg; Wyllie’s Poort, Ingwe Motel; Jan Kempdorp;
King William’s Town; Kirkwood; Knysna; Komatipoort; Kruger National Park; Kwambonambi; Louis
Trichardt; Lynnwood; Malipsdrift; Marble Hall; Nelspruit; Nlcandla; Nkwalini; Olifantshoek; Onrus
River; Paarl; Pienaarspoort; Pietermaritzburg; Piketberg; Port Elizabeth; Port Shepstone; Porterville;
Pretoria; Riebeelc Kasteel; Roodeplaat; Rustenburg; Somerset West; Stellenbosch; Tshipise; Tzaneen;
Uitenhage; Vollcsrust; Vryburg; Warner Beach; Wynberg.
3
Table 1 . Accession numbers and details of the reference DNA barcodes deposited in the Barcode of Life Data Systems (BOLD, http ://www.boldsv stems.
org). PT = paratype; SP = non-type (supplementary) material; F = female; M = male. Negative latitudes are S. Longitudes are E.
European Journal of Taxonomy 233 : 1-23 ( 2016 )
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De Meyer M. et a! . New Ceratitis species (Diptera: Tephritidae) from Africa
Fig. 1. — a-f. Ceratitis quilicii De Meyer, Mwatawala & Virgilio sp. nov. $. a. Head and thorax,
anterodorsal view. b. Head and thorax, lateral view. c. Thorax, dorsal view. d. Midleg, anterior view,
e. Wing. f. Midtibia, anterior view. — g. C. rosa Karsch, 1887 s.str. midtibia, anterior view. Scale bars
= 1 mm .
5
European Journal of Taxonomy 233: 1-23 (2016)
SWAZILAND: Ngonini Estates.
TANZANIA: Amani; Arusha Municipality; Kibundi; Kidiwa; Langali; Lushoto; Mikese; Mlali;
Morogoro Municipality; Moshi Municipality; Mgeta Msikitini; Nyandira; Pinde; Tengeru; Visada.
ZIMBABWE: Harare; Vumba.
Description
Male
Body length. 4.81 (3.68-5.68) mm ; wing length: 5.55 (4.48-6.08) mm
Head (Fig. la). Antenna yellow. First flagellomere in lateral view 2-3 times as long as wide, obtuse
apically. Arista short to medium pubescent, ventral proximal rays at most twice width of arista at base.
Two frontal setae, thinner than, and subequal in length to, anterior orbital seta; two orbital setae, anterior
orbital longer than posterior one; ocellar seta at least 4 times as long as ocellar triangle; postocellar
seta black, shorter than lateral vertical seta. Frons convex, not protruding in lateral view, yellow to
yellowish-white. Genal seta and setulae black. Face and occiput yellowish-white, the latter somewhat
darker dorsally.
Thorax (Fig. lb-c). Postpronotal lobe white to yellowish-white, without black middle spot around
base of postpronotal seta. Scutum ground color greyish to greyish-brown, sometimes with orange tinge;
with streaks and darker markings but without distinct spots except pair of separate prescutellar white
markings, usually with pale yellowish-white area in between. Setae black. Anepisternum on ventral half
darker yellowish-brown to brown; with pale pilosity, one anepisternal seta. Anatergite and lcatatergite
white. Scutellum yellowish-white, usually with two narrow separate dark brown spots basally, sometimes
less distinct; apically with three separate black spots, extending anteriorly to level of or just anterior to
basal scutellar seta. Subscutellum black.
Fegs (Fig. Id, f). Slender; yellow or yellowish-white except where otherwise noted; setation mixed
pale and black. Forefemur with dispersed rows of long black setulae posterodorsally, posteroventrally
shorter and pale; ventral spine-like setae black. Midfemur with few dispersed pale setulae ventrally;
midtibia thin at base, moderately and gradually broadened; anteriorly black with conspicuous silvery
shine when viewed from certain angle on distal 0.66 to 0.75 (black color sometimes inconspicuous in
teneral specimens but silvery shine is always present), black color usually not reaching ventral and
dorsal margins, especially on basal part; with black feathering dorsally along distal 0.75 and ventrally
along distal 0.66, occasionally to distal 0.75. Hindfemur at distal 0.25 with longer setulae dorsally and
ventrally.
Wing (Fig. le). Markings yellowish-brown. Anterior apical band, subapical band and discal band
present, posterior apical band absent; anterior apical band not touching discal band; subapical band
isolated. Cross-vein R-M situated at or just before midlength of cell dm. Brown streaks and spots present
in basal cells.
Abdomen. Ground colour ma inl y yellow. Tergites 2 and 4 on posterior half with greyish microtrichosity;
anterior margin sometimes narrowly brownish colored, especially laterally. Tergite 3 with posterior half
patchily brownish, anterior half yellowish-brown, both parts not clearly demarcated; sometimes more
extensively brown. Tergite 5 with basal half brownish, sometimes divided medially by paler spot.
Female
Unknown (see remarks).
6
De Meyer M. et al. , New Ceratitis species (Diptera: Tephritidae) from Africa
Distribution
Based upon the above listing, C. quilicii sp. nov. is widely distributed throughout southern and eastern
Africa with confirmed records from Botswana, Kenya, Malawi, South Africa, Swaziland, Tanzania and
Zimbabwe. It is also known from Reunion Island.
Host records
The recent recognition that C. rosa s.lat. actually comprises two distinct species (see Remarks below)
requires a re-examination of all material previously reported under ‘C. rosa ’ , including records on
host use. Reared material that could be studied and identified confirmed the following plants as hosts
for C. quilicii sp. nov.: Myrtaceae: Psidium cattleianum Sabine, P. guajava T., Syzygium jambos (L.)
Alston; Rosaceae: Eriobotrya japonica (Thunb.) Lindley, Malus domestica Borkh., Prunus persica
(L.) Batsch, Pyrus communis T., Rubus sp.; Rubiaceae: Coffea arabica L.; Sapotaceae: Chrysophyllum
magalismontanum Sond., Englerophytum natalense (Sond.) T.D. Penn.
Remarks
Ceratitis quilicii sp. nov. belongs to the subgenus Pterandrus and in particular to the Pterandrus
section A as defined by Barr & Wiegmann (2009). Within this section, it belongs to the Ceratitis FAR
complex as defined by Barr & McPheron (2006) and Virgilio et al. (2008). A recent study by Virgilio
et al. (2013) recognized five microsatellite genotypic clusters within the complex, two of which
correspond with Ceratitis rosa. Further studies including morphometries, developmental physiology,
cuticular hydrocarbons, pheromones and mating incompatibility (De Meyer et al. 2015b and references
therein) provided evidence that these two genotypic clusters represent two distinct entities that should
be considered separate species. Mwatawala et al. (2015) furthermore presented evidence that the two
entities appear to have different environmental requirements, a case that was supported by some of
the differences observed by Tanga et al. (2015). These studies also confirmed the earlier proposed
hypothesis by Grout & Stoltz (2007) that C. rosa could actually include two separate entities with
different ecological requirements. However, a study along an altitudinal transect in central Tanzania
(Mwatawala et al. 2015), as well as re-examinations of material housed in natural history collections,
have shown that the two species can co-occur in particular areas.
This species is largely identical to C. rosa. Only the males can be distinguished by minor differences
of the midtibia, C. rosa having a broader midtibia with black coloration reaching the ventral and dorsal
margins of the tibia throughout (Fig. lg), while C. quilicii sp. nov. has a more slender tibia, gradually
tapering towards the base, and with the black coloration not reaching the ventral and dorsal margins
throughout the full length (Fig. If). Females cannot be differentiated currently and, therefore, no female
specimens are included in the type series. The description of the female of C. rosa as given in De Meyer
& Freidberg (2006) applies to C. quilicii sp. nov. as well.
Ceratitis rosa has been referred to as “Rl”, “the hot type” or “lowland type” in previous literature
regarding the two species, while C. quilicii sp. nov. has been referred to as “R2”, “the cold type” or
“highland type” (see De Meyer et al. 2015b and Hendrichs et al. 2015, and references therein). The DNA
barcodes produced for five specimens of C. quilicii sp. nov. from Tanzania, South Africa and Ta Reunion
(see Table 1 for voucher details and accessions) have a mean p-distance = 0.6% (Tamura et al. 2013). As
already observed (Virgilio et al. 2008), DNA barcoding does not allow the unambiguous identification
of species within the Ceratitis FAR complex. C. quilicii sp. nov. is no exception, as it clusters together
with vouchers of C. rosa from Kenya and Mozambique (the morphological identification of these latter
was verified and confirmed as C. rosa) (Supplementary file).
7
European Journal of Taxonomy 233: 1-23 (2016)
Ceratitis (Ceratalaspis) pallidula De Meyer, Mwatawala & Virgilio sp. nov.
urn:lsid:zoobank.org:act:A06A2BlF-6C0F-4AB5-80AD-5216E863C6A8
Fig. 2
Etymology
After the Latin word ‘pallidulus’, meaning somewhat pale (as diminutive of pallidus). The species name
should be treated as an adjective.
Material examined
Holotype
TANZANIA: S, Morogoro, Solcoine University of Agriculture, Horticulture Unit, EGOlure trap,
T19570, 15 Jun. 2013, M. Mwatawala & J. Kusolwa (RMCA).
Paratypes (deposited in RMCA, SUA, BMNH, SANC, and USNM)
TANZANIA: same locality as holotype: 16 $$, 1 Jun. 2013, T19480; 14 1 Jun. 2013, T19486;
8 15 Jun. 2013, T19570; 3 S3, 15 Jun. 2013, T19576; 10 S3, 29 Jun. 2013, T19660; 3 33, 29
Jun. 2013, T19663; 5 33, 29 Jun. 2013, T19666, all EGOlure trap, M. Mwatawala & J. Kusolwa; 18
33, 1 Jun. 2013, T19482; 10 33, 1 Jun. 2013, T19485, both Terpinyl acetate trap, M. Mwatawala &
J. Joseph; 34 33 (4 33 DNAbarcoded, see Table 1), 5-28 Jun. 2011, M. Mwatawala, EGOlure trap.
Non-type material
BURUNDI: 16 33, Rumonge, 24 Jul.-ll Nov. 2013, terpinyl acetate trap, B. Nzigidahera (RMCA).
ETHIOPIA: 2 33 (1 3 DNAbarcoded, see Table 1), Badano, Oct. 2010, terpinyl acetate trap, M. Mansell
(RMCA).
MALAWI: 2 33 (1 $ DNA barcoded, see Table 1), Zomba, 31 Oct. 2010-7 Mar. 2011, EGOlure trap,
M. Hendrickx (RMCA).
TANZANIA: 1 3, Kimani, RtA104, 29 Aug. 1996, A. Freidberg (TAU).
Description
Male
Body length. 3.0 (2.72-3.36); wing length: 3.10 (2.80-3.36) mm.
Head (Fig. 2a). Antenna yellow, first flagellomere in lateral view twice as long as wide; with short acute
tip apically. Arista short pubescent, ventral proximal rays at most equal to width of arista at base. Two
frontal setae, thinner than, and subequal in length to, anterior orbital seta; two orbital setae, anterior
orbital longer than posterior one; ocellar seta about 3 times as long as ocellar triangle; postocellar seta
black, shorter than lateral vertical seta. Frons convex, not protruding in lateral view; yellow-white.
Genal seta pale, genal setulae mixed yellow and dark. Face and occiput white.
Thorax (Fig. 2b-c). Postpronotal lobe yellowish-white, without black middle spot around base of
postpronotal seta. Scutum ground colour shining pale yellow to yellow-orange, with pale pilosity/
microtrichosity; distinct paired black spot situated between dorsocentral and prescutellar acrostichal
setae, (almost) touching, but not engulfing prescutellar acrostichal seta, posteriorly with prescutellar
white semi-circular or similar marking; indistinct to distinct black spot near mesal end of transverse
suture; occasionally additional black medial spot at level of transverse suture; lateral margins of scutum
with black spots reduced, only paired spot on posterior lateral corner well developed. Setae black,
except scapulars white. Pleura yellow-white. Anepisternum dorsal margin narrowly white, ventral
8
De Meyer M. et al ., New Ceratitis species (Diptera: Tephritidae) from Africa
half sometimes darker yellow, one anepistemal seta. Anatergite white, rarely with irregular black spot.
Scutellum yellowish-white, apical margin with three large separate black spots, anteriorly extending to
level of basal scutellar setae; with two brown to blackish, round submedial spots basally. Subscutellum
black, sometimes more yellow submedially on ventral part.
Fig. 2. Ceratitis pallidula De Meyer, Mwatawala & Virgilio sp. nov. a. Head and thorax, anterodorsal
view. b. Head and thorax, lateral view. c. Thorax, dorsal view. d. Abdomen, dorsal view. e. Wing. Scale
bars = 1 mm .
9
European Journal of Taxonomy 233: 1-23 (2016)
Legs. Slender; yellow; with dispersed pale pilosity. Forefemur with pale ventral spine-like setae.
Wing (Fig. 2e). Markings yellow to yellowish-brown. Anterior apical band, subapical band and discal
band present, posterior apical band absent; anterior apical band touching discal band; subapical band
isolated. Cross-vein R-M at about basal third of cell dm. Brown streaks and spots present in basal cells.
Abdomen (Fig. 2d). Ground colour yellow to orange; setulae and setae dark brown. Tergites 2 and 4 on
posterior half to two-thirds with greyish microtrichosity.
Female
Unknown.
Distribution
Burundi, Ethiopia, Tanzania, Malawi.
Host plants
Unknown.
Remarks
While this species was until recently unknown, it has been found in relatively high numbers (up to
400 flies/trap/week) through trapping activities using enriched ginger oil (EGOlure, Insect Science,
Tzaneen South Africa) in central Tanzania (Mwatawala et al. 2013; referred to as ‘C. near cosyra ’). The
first identification placed this as small specimens of C. cosyra (Walker, 1849) but the morphological
differences in mesonotal pattern and genetic differentiation of DNA barcodes (Table 1; Supplementary
file) indicated this was not the case. Afterwards, additional specimens were trapped in other countries
and it appears to be present throughout a wide range. The species is similar to C. cosyra but can be
readily differentiated by the absence of a black spot on the postpronotal lobe and the much smaller size.
It also resembles C. quinaria (Bezzi, 1918) and C. silvestrii Bezzi, 1912 in general appearance but can
be differentiated from the former by the three apical spots on the scutellum (five spots in C. quinaria )
and from both by the presence of a black spot near the prescutellar acrostichal seta (absent in both
C. quinaria and C. silvestrii).
Ceratitis (Ceratalaspis) taitaensis De Meyer & Copeland sp. nov.
urn:lsid:zoobank.org:act: 1BFD784E-BA50-4D89-899A-707DD7D6A3B6
Fig. 3
Etymology
The name is considered as an adjective derived from the geographical name ‘Taka’ referring to the
Taita Hills, located in southeastern Kenya. The Taita Hills are the northernmost block of the Eastern Arc
Mountain chain.
Material examined
Holotype
KENYA: S, Vuria, reared from fruits of Lepidotrichilia volkensii, 16 May 2012, R.S. Copeland (NMK)
(CHIESA coll. Nr 228).
Paratypes (deposited in NMK, ICIPE, RMCA, BMNH, and NMNH)
KENYA: same locality as holotype: 3 $$, 2 $ §, 6 Jun. 2012, CHIESA coll. Nr 170; 2 6 Jun. 2012,
CHIESA coll. Nr 171; 10 10 16 May 2012, CHIESA coll. Nr 228 (2 2 barcoded
10
De Meyer M. et a! . New Ceratitis species (Diptera: Tephritidae) from Africa
Fig. 3. Ceratitis taitaensis De Meyer & Copeland sp. nov. a. Head and thorax, anterodorsal view,
b. Thorax, lateral view. c. Thorax, dorsal view. d. Abdomen, dorsal view. e. Male wing. f. Female wing,
g. Female aculeus. h. Aculeus tip. Scale bars: A-F = 1 mm; G-H = 0.1mm.
11
European Journal of Taxonomy 233: 1-23 (2016)
RMCA, see Supplementary file); 1 8, 1 ?, 10 Jul. 2012, CHIESAcoll. Nr 261; 33 88, 37 22 Aug.
2012, CHIESA coll. Nr 269, all R.S. Copeland, reared from Lepidotrichilia volkensii (Gurke) Leroy.
Description
Male
Body length. 5.61 (5.04-6.08) mm; wing length: 6.20 (5.68-6.88) mm.
Head (Fig. 3a). Antenna yellow. First flagellomere in lateral view twice as long as wide; obtuse apically.
Arista short to medium pubescent, ventral proximal rays at most twice width of arista at base. One
frontal seta, thinner than, and subequal in length to, anterior orbital seta; two orbital setae, anterior
seta longer than posterior one; ocellar seta about 3^1 times as long as ocellar triangle; postocellar seta
black, shorter than lateral vertical seta. Frons convex, not protruding in lateral view; yellow to orange,
with greyish microtrichosity on posterior half. Gena broader than in other Ceratitis species (maximum
diameter of eye versus height of gena 2.5-3.0), genal seta and genal setulae yellow, latter sometimes
blackish.
Thorax (Fig. 3b-c). Postpronotal lobe yellowish-white to white; with black middle spot around
base of postpronotal seta. Scutum ground colour shining yellow-brown to dark brown, with silvery
microtrichosity covering most of dorsum, except circular area posterior to mesal end of transverse suture
and in trapezoid area extending posteriorly from dorsocentral setae to anterior margin of scutellum, and
narrowly along lateral margins; sublaterally the microtrichosity extends posteriorly to the intralar seta;
narrow area along anterior margin, extending posteriorly along midline and posterior of postpronotal
lobe, with less dense microtrichosity. Anepisternum yellowish-white, lower margin darker; with pale
pilosity, one anepisternal seta. Scutellum yellowish-white, with three yellow-brown spots restricted to
apical margin and ventral side; area between spots darker yellowish coloured. Subscutellum entirely
brown to black.
Legs. Slender; yellow to yellowish-orange, tarsi sometimes slightly paler than rest of leg; with dispersed
pale pilosity. Forefemur with dark brown ventral spine-like setae. Hindfemur at distal 0.25 with dark
brown setae dorsally.
Wing (Fig. 3e). Markings brownish to yellowish-brown. Of typical bands, only anterior apical band
distinct, including pterostigma and area posterior of pterostigma to vein R4+5; furthermore with
brownish spot covering area surrounding cross-vein R-M (i.e., apical margin of cell br) and basal third
of cell r4+5, continued in apical half of cell dm and anterior third of cell m, also broadly fused with
anterior apical band; additional small marks in middle of cell cUj and basal part of cell m. Cross-vein
R-M at or just beyond midlength of cell dm. Brown streaks and spots present in basal cells but poorly
developed.
Abdomen (Fig. 3d). Ground colour yellow to orange-brown. Tergites 2 and 4 on posterior half to two-
thirds with greyish microtrichosity.
Female
As male except for the following characters: gena less broad (maximum eye diameter to gena height
ratio less than 2.5). Wing with well developed bands as in other Ceratitis species (Fig. 3f): discal band
interrupted in cell dm; anterior apical band and discal band separated or only narrowly touching; subapical
band narrowly touching anterior part of discal band; posterior apical band isolated. Oviscape orange,
with dispersed dark brown to black pilosity. Tergal-oviscapal ratio (= length of abdominal tergites 1-5
versus length of oviscape): 1.5-2. Aculeus (Fig. 3g-h) flattened, 7-8 times longer than broad, apex
bifurcated, and with pair of subapical protuberances.
12
De Meyer M. et al ., New Ceratitis species (Diptera: Tephritidae) from Africa
Distribution
Kenya.
Host plants
Reared from fruits of Lepidotrichilia volkensii (Gtirke) Leroy (Meliaceae).
Remarks
Ceratitis whartoni was described by De Meyer & Copeland (2009) from forested areas in the Central
Highlands of Kenya (Gatamayu Forest, Nyanduma Forest, Mt. Kenya Forest), all reared from fruits of
Lepidotrichilia volkensii (Meliaceae). It was an enigmatic species because of the dimorphic wing pattern,
with that of the male largely differing from the standard wing banding found in most other Ceratitis
species. A series of specimens reared from the same host plant collected in Taita Hills (approximately 400
km southeast of the nearest site of C. whartoni) are morphologically almost identical, with only slight
differences in wing pattern and female aculeus (male wing with dark marking in cell dm occupying more
than half of cell in C. whartoni , less than half in C. taitaensis sp. nov.; female wing with discal band
complete in C. whartoni , partially interrupted in cell dm in C. taitaensis sp. nov.; aculeus with bifurcated
apex more slender in C. taitaensis sp. nov. (less than one-third of entire width) and invagination less
deep than in C. whartoni). However, the analysis of the available DNA barcodes of C. whartoni (Table 1;
Supplementary file) revealed remarkable genetic differentiation from C. taitaensis sp. nov. (p-distance
= 8.5%). It was, therefore, decided to recognize this as a separate species. The Taita Hills are a chain
of forested mountain tops surrounded by the flat and dry Tsavo Plains. They form the northernmost
mountainous massif of the Eastern Arc Mountains, a chain of ancient crystalline mountains (Burgess et
al. 2007) and are considered a major biodiversity hotspot (Meyers et al. 2000). We place this species in
the subgenus Ceratalaspis for the same reasons as outlined in De Meyer & Copeland (2009) regarding
the placement of C. whartoni.
Ceratitis (Ceratalaspis) sawahilensis De Meyer & Virgilio sp. nov.
urn:lsid:zoobank.org:act:F57EF53D-53E8-4EBF-A268-8FB819DBEEB4
Fig. 4
Etymology
The name is considered as an adjective derived from a geographical name, the Arabic word ‘sawahiF
being the plural of ‘sahiF, which means border or coast and refers to the Swahili Coast, which was
recognized as a historical cultural and geographic entity comprising the localities of Zanzibar and
Mkuranga, where the type series was collected.
Material examined
Holotype
TANZANIA: c?, Mkuranga, Mwanambaya, 21 Jan. 2001, Z. Seguni (RMCA).
Paratypes
TANZANIA: 1 same date and locality as holotype; 1 S, 1 $, Zanzibar, 7 Feb. 2000, Z. Seguni (1
$ barcoded, see Table 1; Supplementary file), (RMCA); 1 $ (Fig. 4f), Magombero Forest, 14-15 Sep.
2009, T. Pape & S.A. Marshall (ZMUC).
Description
Male
Body length. 5.01 (4.80-5.28); wing length: 5.04 (4.80-5.20) mm
13
European Journal of Taxonomy 233: 1-23 (2016)
g
I-1
Fig. 4. Ceratitis sawahilensis De Meyer & Virgilio sp. nov. a. Head and thorax, anterodorsal view,
b. Thorax, lateral view. c. Thorax, dorsal view. d. Abdomen, dorsal view. e. Wing. f. Habitus image
(credit: S.A. Marshall), g. Female aculeus. h. Aculeus tip. Scale bars: A-F = 1 mm; G-H = 0.1 mm.
14
De Meyer M. et at.. New Ceratitis species (Diptera: Tephritidae) from Africa
Head (Fig. 4a). Antenna yellow-orange, first flagellomere in lateral view 3 times as long as wide, obtuse
or with short acute tip dorsoapically. Arista medium pubescent, ventral proximal rays at most 3 times
width of arista at base. Two frontal setae, thinner than, and subequal in length to, anterior orbital seta;
two orbital setae, anterior orbital longer than posterior one; ocellar seta at least 4 times as long as ocellar
triangle; postocellar seta black, shorter than lateral vertical seta. Frons convex, not protruding in lateral
view; yellow-white, in between frontal setae more yellowish. Genal seta and genal setulae dark brown
to reddish-pale. Face and occiput yellowish-white, latter slightly darker dorsally.
Thorax (Fig. 4b-c). Postpronotal lobe white, without black middle spot around base of postpronotal
seta. Scutum ground colour shining yellow-brown to brown, sometimes with more orange tinge, with
greyish pilosity and microtrichosity; with slightly darker streaks but no distinct black spots, except
paired prescutellar spot extending anteriorly to dorsocentral seta, posteriorly with paired white to
yellow prescutellar semi-circular marking, separated by yellowish-white area in between; at mesal end
of transverse suture with shining brownish to blackish spot. Setae black. Pleura yellow to brownish-
yellow. Anepisternum dorsal half white, ventral half yellow; completely covered with pale pilosity,
one anepisternal seta. Anatergite and lcatatergite white. Scutellum yellowish-white to white, sometimes
with yellowish tinge; apical margin with three separate black spots, extending to level of basal scutellar
setae; with distinct brown-black spots basally, sometimes less pronounced and only darker yellow-
brown colour. Subscutellum black.
Legs. Slender; yellow; with dispersed mixed pale and dark pilosity. Forefemur ventral spine-like setae
black along distal 0.50, basally more pale.
Wing (Fig. 4e). Markings yellow to yellow-brown. Anterior apical band, subapical band, discal band and
posterior apical band present, latter sometimes only as a trace; anterior apical band touching discal band;
subapical band and posterior apical band isolated. Cross-vein R-M situated at or basal to midlength of
anterior margin of cell dm. Brown streaks and spots present in basal cells.
Abdomen (Fig. 4d). Ground colour yellow. Tergites 2 and 4 more greyish along posterior half; tergites
3 and 5 with, respectively, posterior and anterior margin narrowly more darkly yellow. With mixed pale
and dark pilosity.
Female
As male. Oviscape shorter than preabdomen. Aculeus (Fig. 4g-h) about 8-9 times longer than wide;
apical part sinuous, tip blunt and with distinct indentation.
Distribution
Tanzania.
Host plants
Unknown.
Remarks
The relationship of C. sawahilensis sp. nov. to any of the other described Ceratitis species is not easily
established. Although there is a slight partial darkening of the abdominal tergites 3 and 5, the male
legs are without modifications. The neighbor-joining tree (Supplementary file) does not place it in any
resolved cluster. We tentatively place it in the subgenus Ceratalaspis pending further clarifications on
its subgeneric position.
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European Journal of Taxonomy 233: 1-23 (2016)
Ceratitis (Ceratalaspis) flcivipennata De Meyer & Virgilio sp. nov.
um:lsid:zoobank.org:act:72C8DD57-0FDl-4B48-96FA-881BFDDBA205
Fig. 5
Etymology
After the Latin favus ’ meaning yellow and ‘ pennatus ’ meaning feathered, and referring to the yellow
feathering on the midtibia. The species name should be treated as an adjective.
Material examined
Holotype
KENYA: Karara Forest, 11 Aug. 2006, N. Erbout (RMCA).
Paratype
KENYA: 1 S, same locality and data as holotype (barcoded, see Table 1; Supplementary file).
Description
Male
Body length. 4.08 (3.92—4.16) mm; wing length: 4.88 (4.80-4.96) mm
Head (Fig. 5a). Antenna yellow, first flagellomere in lateral view 3 times as long as wide, obtuse apically.
Arista short to medium pubescent, ventral proximal rays at most twice width of arista at base. Two
frontal setae, thinner than, and subequal in length to, anterior orbital seta; two orbital setae, anterior
orbital longer than posterior one; ocellar seta about 3 times as long as ocellar triangle; postocellar seta
black, shorter than lateral vertical seta. Frons convex, not protruding in lateral view; yellow. Genal seta
and genal setulae pale to dark orange. Face yellow-white, occiput yellow.
Thorax (Fig. 5b-c). Postpronotal lobe yellow-orange, slightly paler than scutum; without distinct black
middle spot around base of postpronotal seta but with darker irregular marking near seta. Scutum ground
colour shining orange to orange-brown, with pale pilosity/microtrichosity; with distinct small black
spot near mesal end of transverse suture; otherwise no distinct black marking on central part of scutum;
paired separate prescutellar semi-circular white markings posterior of prescutellar acrostichal seta. Setae
black, except scapulars red brown. Pleura yellow-orange to yellow-brown. Anepisternum dorsal margin
narrowly white, ventral third to half more brownish; one anepisternal seta. Anatergite yellow-white,
with irregular black marking; katatergite yellow-white. Scutellum yellowish-white, apical margin with
three separate black spots, extending anteriorly beyond basal scutellar seta; without distinct black spots
basally, at most with slightly darker marking. Subscutellum yellow with three black spots of which the
upper margin is confluent, thereby forming single black band.
Legs (Fig. 5e). Slender; yellow to yellow-orange; with dispersed yellow-orange pilosity. Forefemur
with orange ventral spine-like setae, posterodorsally and posteriorly with several rows of longer yellow-
orange setae, producing bushy appearance. Midtibia with yellow-orange feathering dorsally along distal
0.45 and ventrally along distal 0.30.
Wing (Fig. 5f). Markings yellow to yellow-brown. Anterior apical band, subapical band, posterior apical
band and discal band present; anterior apical band not touching discal band; subapical band touching
discal band; posterior apical band touching anterior apical band. Cross-vein R-M situated just beyond
midlength of cell dm. Brown streaks and spots present in basal cells.
Abdomen (Fig. 5d). Ground colour yellow to orange, pale and brown setulae. Tergites 2 and 4 along
posterior margin slightly more greyish.
16
De Meyer M. et at.. New Ceratitis species (Diptera: Tephritidae) from Africa
Female
Unknown.
Distribution
Kenya.
Fig. 5. Ceratitis flavipennata De Meyer & Virgilio sp. nov. a. Head and thorax, anterodorsal view,
b. Thorax, lateral view. c. Thorax, dorsal view. d. Abdomen, dorsal view. E. Midleg, anterior view.
F. Wing. Scale bars = 1 mm
17
European Journal of Taxonomy 233: 1-23 (2016)
Host plants
Unknown.
Remarks
Ceratitis flavipennata sp. nov. resembles Ceratitis neostictica De Meyer, 1998 in the pilosity of the
forefemur and feathering on the midtibia. It can be differentiated from the latter by the differences in
wing banding (posterior apical band touching anterior apical band in flavipennata sp. nov., separate in
neostictica ; subapical band touching discal band in flavipennata sp. nov., separate in neostictica) and
the pattern of the scutum (extensive black markings in neostictica , largely absent in flavipennata sp.
nov.). Like C. neostictica , C. flavipennata sp. nov. is placed in the subgenus Ceratalaspis based on the
colouration of the abdominal tergites (predominantly yellow to orange, no distinct black transverse
bands). Based on its DNA barcode it does not cluster with any other species within the genus Ceratitis
(see Supplementary file).
Ceratitis (Pardalaspis) serrata De Meyer, 1996
Fig. 6
Material examined
Holotype
DEMOCRATIC REPUBLIC OF CONGO: $, Yangambi, 17 Nov. 1960, J.M. McGough (TAU).
Non type material
DEMOCRATIC REPUBLIC OF CONGO: Masako, near Kisangani: 16 33 , 21 Mar. 2008; 11 33 , 28
Mar. 2008; 4 33, 25 Apr. 2008; 5 33, 4 Apr. 2008; 1 3, 2 May 2008, all methyl eugenol trap, J.-L.
Juakaly (RMCA); 1 §, Congo River Expedition, Bomane, 19-24 May 2010, cue lure trap, R. Emeleme
& M. Virgilio (RMCA).
Description
Male
Head (Fig. 6a-b). Antenna orange. First flagellomere in lateral view 2-3 times as long as wide, obtuse
apically. Arista short pubescent, ventral proximal rays at most equal to width of arista at base. Two
frontal setae, thinner than, and equal in length, to anterior orbital seta; two orbital setae, anterior orbital
longer than posterior one; ocellar seta 3M times as long as ocellar triangle; postocellar seta black,
shorter than lateral vertical seta. Frons flattened, slightly protruding in lateral view, completely covered
with silvery shine. Genal seta and setulae black. Face orange, occiput yellowish.
Thorax (Fig. 6c-d). Postpronotal lobe greyish to greyish-yellow, without black middle spot around
base of postpronotal seta. Scutum ground color greyish-brown, sometimes with golden orange tinge;
with streaks and darker markings but without distinct spots except for darkish spot around prescutellar
acrostichal seta, and pale prescutellar semi-circular marking along posterior margin near prescutellar
acrostichal seta. Setae black; setulae mainly pale; black setulae restricted to area at mesal end of tranverse
suture extending posteriorly to prescutellar acrostical and dorsocentral setae. Anepisternum ventral half
brownish, dorsal half more greyish, completely covered with black pilosity, except for horizontal stripe
below dorsal margin with white pilosity; three anepisternal setae. Anatergite and katatergite brownish.
Scutellum dark yellowish, apical margin with three separate black spots, anteriorly extending anteriorly
beyond basal scutellar setae; with two large roundish black spots basally. Subscutellum black.
Legs. Slender; yellow-orange, midfemur more brownish; with dispersed and mainly black pilosity.
Fore femur with ventral setae black.
18
De Meyer M. et al ., New Ceratitis species (Diptera: Tephritidae) from Africa
Wing. Markings dark brown. Anterior apical band, subapical band and discal band present, posterior
apical band absent; anterior apical band touching discal band; subapical band isolated. Cross-vein R-M
situated at midlength of cell dm. Brown streaks and spots present in basal cells.
Abdomen (Fig. 6e). Ground colour mainly greyish to pale orange; with darker spots on all tergites. With
mixed pale and black pilosity.
Fig. 6. Ceratitis serrata De Meyer, 1996. a. Head, frontal view. b. Head and thorax, anterodorsal view,
c. Thorax, lateral view. d. Thorax, dorsal view. e. Abdomen, dorsal view. Scale bars: A = 0.5 mm; B-D =
1 mm
19
European Journal of Taxonomy 233: 1-23 (2016)
Distribution
Congo (Democratic Republic).
Host plants
Unknown.
Remarks
Ceratitis serrata was originally described from a female collected in Yangambi in the Democratic
Republic of Congo. Trapping with methyl eugenol at Masako (near Kisangani and approx. 100 1cm east
of Yangambi) in 2008 collected male specimens that did not match any of the known species within the
subgenus Pardalaspis. Virgilio et al. (2011) reported a female specimen of C. serrata from Bomane
along the Congo River, further west of Yangambi. DNA barcoding revealed that the COI sequences
obtained from the male specimen from Masako (series of 21.III.2008 AccessID 13954, AB33598909)
and the female specimen from Bomane (AccessID 15755, AB40159308) differed by a p-distance of
only 0.5% (see Table 1; Supplementary file) with a large barcoding gap (corresponding to 7% similarity)
separating C. serrata from the second closest match. It was therefore considered that the material from
Masako represents the hitherto unknown male of C. serrata. Male specimens of C. serrata can be
readily differentiated from other species within the subgenus Pardalaspis by the combination of the
following characters: frons completely silvery shining; face uniform orange coloured; anterior margin of
scutum same colour as middle part; anepisternum largely covered with black pilosity, base of scutellum
with a pair of distinct black spots.
Discussion
As indicated earlier, the taxonomy of the different subgenera within the genus Ceratitis has been
revised in the last two decades (De Meyer 1996, 1998, 2000; De Meyer & Copeland 2001; De Meyer
& Freidberg 2006). For the most part, these revisions were based on material already deposited in
different collections. However, because of more intensive surveying activities in different parts of Africa,
additional undescribed species were encountered, and some have already been described (De Meyer &
Copeland 2005, 2009). Moreover, the multidisciplinary approach used in integrative taxonomy revealed
the presence of cryptic species (see De Meyer et al. 2015a for a recent treatise of this in tephritid fruit
flies). In groups of economic significance, such as Tephritidae, a proper identification is not only a
prerequisite for any further research, be it fundamental or applied, but can also have serious political
and economic consequences in aspects like control programs, quarantine regulations, and trade barriers
(Hendrichs et al. 2015). Taxonomic recognition, preferably through different independent approaches,
is a necessity and may facilitate accurate recognition of biological species (Clarke & Schutze 2014).
The description of Ceratitis quilicii sp. nov. in this paper is a case in point. Additional alpha taxonomic
descriptions such as these contribute to the general knowledge of the group, and will allow more
accurate identifications. It is, however, a requisite that the newly acquired information is also transferred
to proper identification and reference tools such as DNA sequence libraries like BOFD or identification
tools such as those presented by Virgilio et al. (2014).
Acknowledgments
The authors acknowledge the financial support by the following funding agencies and projects: the
Belgian Development Cooperation through the framework program of the Royal Museum for Central
Africa (project S1TNZIPM: Integrated Pest Management (IPM) for Fruit Flies in Tanzania); the
joint Food and Agriculture Organisation / International Atomic Energy Agency Programme on Nuclear
Techniques in Food and Agriculture (Co-ordinated Research Project on Resolution of Cryptic Species
Complexes of Tephritid Pests to Overcome Constraints to the SIT (Sterile Insect Technique) Application
20
De Meyer M. et al ., New Ceratitis species (Diptera: Tephritidae) from Africa
and International Trade); Fonds voor Wetenschappelijk Onderzoek Vlaanderen (travel grant to MDM
to visit the collections of the Stellenbosch University); Government of Finland (CHIESA Project at the
International Centre for Insect Physiology and Ecology to RSC: Climate Change Impacts on Ecosystem
Services and Food Security in Eastern Africa). We thank the curators of the natural history collections
who allowed us to study material and assisted us in our visits: R. Urban and M. Mansell (SANC), P
Addison (SUS), A. Freidberg (TAU), T. Pape (ZMUC). Thanks also to the director of Kenya Forest
Service for permission to sample in the forests of Taita Hills; to S. Marshall for allowing us to use the
habitus photograph of C. sawahilensis sp. nov., to W. Fannes for his assistance in taking images, and to
Allen Norrbom and Amn on Freidberg for providing valuable co mm ents to an earlier draft of this article.
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Manuscript received: 7 March 2016
Manuscript accepted: 3 May 2016
Published on: 26 September 2016
Topic editor: Gavin Broad
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; Botanic Garden 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; Naturalis Biodiversity Center, Leiden, the Netherlands.
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