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An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae) c yp ic
species complex: a e iew
De Meye , Ma c; Dela e, Hélène; Ekesi, Sunday; Jo daens, Ku ; Kalino a, Blanka;
Man akhan, A una; Mwa awala, Maulid; S eck, Ga y; Van Cann, Joannes; Vanciko a,
Lucie; B izo a, Radka; Vi gilio, Massimiliano
De Meye , M., Dela e, H., Ekesi, S., Jo daens, K., Kalino a, B., Man akhan, A.,
Mwa awala, M., S eck, G., Van Cann, J., Vanciko a, L., B izo a, R., & Vi gilio, M.
(2015). An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)
c yp ic species complex: a e iew. ZooKeys, 540, 405-427.
h ps://doi.o g/10.3897/zookeys.540.10046
2015
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 405
An in eg a i e app oach o un a el he Ce a i is FAR
(Dip e a, Teph i idae) c yp ic species complex: a e iew
Ma c De Meye 1, Hélène Dela e2, Sunday Ekesi3, Ku Jo daens1,8,
BlankaKalino á4, A una Man akhan5, Maulid Mwa awala6, Ga y S eck7,
JoannesVanCann8,9, Lucie Vaníčko á4,10, Radka Břízo á11, Massimiliano Vi gilio1
1 Royal Museum o Cen al A ica, In e eb a es Sec ion and JEMU, Leu enses eenweg 13, B3080 Te u en,
Belgium 2 CIRAD, UMR PVBMT, 7 ch de l’IRAT, 97410 Sain -Pie e, La Réunion, F ance 3 In e na ional
Cen e o Insec Physiology and Ecology (icipe), P.O. Box 30772 – 00100, Nai obi, Kenya 4 Ins i u o O ganic
Chemis y and Biochemis y, Academy o Sciences o he Czech Republic, P ague 5 Ci us Resea ch In e na io-
nal, PO Box 28, Nelsp ui 1200, Sou h A ica 6 Sokoine Uni e si y o Ag icul u e, Dep . o C op Science and
P oduc ion, Mo ogo o, Tanzania 7 Flo ida Depa men o Ag icul u e and Consume Se ices, Gaines ille FL,
USA 8 Uni e si y o An we p, G oenenbo ge laan 171, B-2020 An we p, Belgium 9 Uni e si y o Jy äskylä,
PO Box 35, 40014 Finland 10 Labo a ó io de Ecologia Química, Ins i u o de Química e Bio ecnologia,
Uni e sidade Fede al de Alagoas, A . Lou i al de Melo Mo a, s/n, Tabulei o, 57072-970, Maceió, AL, B azil
11Ins i u e o Chemical Technology in P ague, Technická 5, CZ-166 28 P ague 6, Czech Republic
Co esponding au ho : Ma c De Meye ([email p o ec ed])
Academic edi o : J. Hend ichs|Recei ed 26 May 2015|Accep ed 5 Augus 2015|Published 26 No embe 2015
h p://zoobank.o g/C072905C-89EE-4BE4-B5B7-33819CD86A3F
Ci a ion: De Meye M, Dela e H, Ekesi S, Jo daens K, Kalino á B, Man akhan A, Mwa awala M, S eck G, Van Cann
J, Vaníčko á L, Břízo á R, Vi gilio M (2015) An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)
c yp ic species complex: a e iew. In: De Meye M, Cla ke AR, Ve a MT, Hend ichs J (Eds) Resolu ion o C yp ic Species
Complexes o Teph i id Pes s o Enhance SIT Applica ion and Facili a e In e na ional T ade. ZooKeys 540: 405–427.
doi: 10.3897/zookeys.540.10046
Abs ac
This pape e iews all in o ma ion ga he ed om di e en disciplines and s udies o esol e he species
s a us wi hin he Ce a i is FAR (C. asci en is, C. anonae, C. osa) complex, a g oup o polyphagous ui
ly pes species (Dip e a, Teph i idae) om A ica. I includes in o ma ion on la al and adul mo phol-
ogy, wing mo phome ics, cu icula hyd oca bons, phe omones, mic osa elli es, de elopmen al physiol-
ogy and geog aphic dis ibu ion. The gene al consensus is ha he FAR complex comp ises C. anonae, wo
species wi hin C. osa (so-called R1 and R2) and wo pu a i e species unde C. asci en is. The in o ma-
ion ega ding he la e is, howe e , oo limi ed o d aw inal conclusions on speci ic s a us. E idence o
his ecogni ion is discussed wi h e e ence o publica ions p o iding u he de ails.
ZooKeys 540: 405–427 (2015)
doi: 10.3897/zookeys.540.10046
h p://zookeys.penso .ne
Copy igh Ma c De Meye e al. This is an open access a icle dis ibu ed unde he e ms o he C ea i e Commons A ibu ion License (CC
BY 4.0), which pe mi s un es ic ed use, dis ibu ion, and ep oduc ion in any medium, p o ided he o iginal au ho and sou ce a e c edi ed.
RESEARCH ARTICLE
Launched o accele a e biodi e si y esea ch
A pee - e iewed open-access jou nal
Ma c De Meye e al. / ZooKeys 540: 405–427 (2015)
406
Keywo ds
Taxonomy, Ce a i is osa, Ce a i is asci en is, Ce a i is anonae, A ica, ui ly
In oduc ion
His o ical backg ound and sys ema ic posi ion
Ce a i is MacLeay is an A o opical genus o eph i id ui lies comp ising close o
100 species ound in Sub-Saha an A ica and he islands o he Wes e n Indian Ocean.
Phylogene ically i belongs o he sub ibe Ce a i idina wi hin he ibe Dacini. The la e
ibe includes all main pes gene a occu ing na u ally in A ica, i.e. Bac oce a Macqua ,
Cappa imyia Bezzi, Ce a i is, Dacus Fab icius, Neoce a i is Hendel and T i hi h um Bezzi.
Howe e , he monophyly o he genus is no suppo ed and some species appea phylo-
gene ically mo e closely ela ed o T i hi h um, while o he Ce a i is species p obably do
no belong o he genus (Ba and McPhe on 2006, Vi gilio e al. 2015). Con a y o
o he indigenous A ican ui ly pes g oups ha ha e a pa icula hos niche wid h a -
acking pa icula plan amilies, species o he genus Ce a i is demons a e a e y a iable
hos speci ici y. Some species a e la gely monophagous, while o he s o m monophyle ic
s enophagous clus e s specialized on pa icula hos gene a. Ye , o he s a e polyphagous
species a acking a wide a ie y o un ela ed plan s (De Meye e al. 2002, E bou e al.
2011). Fo example, he Medi e anean ui ly, Ce a i is capi a a (Wiedemann), is he
mos ex eme case o polyphagy, wi h 304 di e en hos plan s belonging o 154 gene a
in 57 amilies conside ed o be sui able hos plan s (Liquido e al. 2014).
The Ce a i is FAR complex is a g oup o polyphagous species comp ising h ee
mo phologically simila species: C. asci en is (Bezzi), C. anonae G aham and C. osa
Ka sch (Figu e 1). Al hough he FAR complex is ecognized as a g oup o closely ela -
ed species, hei monophyly is no always suppo ed in phylogene ic s udies. Ba and
Wiegmann (2009) p esen ed a molecula analysis o a numbe o Ce a i is (P e an-
d us) species including one specimen o each Ce a i is FAR complex species. They
s a e ha CAD1 and ND6 ma ke s suppo ed he monophyly o he complex, while
ango and pe iod ma ke s placed he h ee species in di e en clus e s wi hin he so-
called ‘P e and us A’ g oup. A majo i y ule Bayesian ee o he combined molecula
da a suppo ed he monophyly o he complex. Ye , a cladis ics analysis based on adul
mo phological cha ac e s could no ecognize hem as a monophyle ic clus e wi hin
he subgenus P e and us (De Meye 2005).
Cu en axonomic s a us o ecognized species and sexual dimo phism
The Na al ui ly, C. osa, was o iginally desc ibed om ’Delagoabai’ (nowadays Baia
de Mapu o, Mozambique and p obably e e ing o a loca ion nea Mapu o) by Ka sch
in 1887, based on a single male specimen (Ka sch 1887). Bezzi (1920) desc ibed as-
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 407
c
b
a
Figu e 1. Habi us image o a Ce a i is asci en is b C. anonae c C. osa (a, c pho os R.S. Copeland, b
pho o G. Goe gen).
Ma c De Meye e al. / ZooKeys 540: 405–427 (2015)
408
ci en is as a a ie y o P e and us osa, based upon ma e ial om Uganda (unknown
locali y). De Meye (2001) conside ed i o be a sepa a e en i y wi h speci ic s a us.
Ce a i is anonae was desc ibed by G aham om ma e ial ea ed om an Annona ui
in Ashan i, Ghana (G aham 1908).
All h ee species show ema kable sexual dimo phism, wi h he males ha ing leg
o namen a ion ha is absen in emales. Ce a i is anonae males ha e he mid leg (Fig-
u e2) wi h a ow o long da k, la ened se ae (so-called ‘ ea he ing’) en ally along
he en i e leng h o he mid emu . The mid ibia is b oadened wi h ea he ing do sally
along dis al 0.9 and en ally along he dis al 0.8. The mid leg is la gely b ownish o
b ownish black in colou . In C. osa males (Figu e 3), he en al ea he ing on mid
emu is absen (a mos he e a e a ew hin and dispe sed se ulae en ally). The mid
ibiae is mode a ely b oadened, an e io ly black wi h a conspicuous sil e y e lec ion
seen when kep unde a ce ain angle and black ea he ing do sally along dis al 0.75
and en ally along dis al 0.66–0.75. O he wise, he leg is yellow. Ce a i is asci en is
males ha e he mid leg shaped simila ly o C. osa, excep ha he mid ibia is no dis-
inc ly b oadened and he black ea he ing is es ic ed o he dis al 0.5 a mos (Figu e
4). The leg is colou ed uni o mly yellow, excep in some specimens whe e he an e io
pa is pa ially b ownish in he dis al 0.3 (De Meye 2001). P elimina y cou ship
beha io s udies by Quilici e al. (2002) gi e some indica ion ha , o a leas in C.
osa, he mid legs a e used in p ecopula o y beha io .
Figu e 2. Mid leg o male Ce a i is anonae, an e io iew o emu and ibia (pho o I.M. Whi e, NHM).
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 409
Figu e 3. Mid leg o male Ce a i is osa, an e io iew (pho o I.M. Whi e, NHM).
Figu e 4. Mid leg o male Ce a i is asci en is, an e io iew (pho o I.M. Whi e, NHM).
Ma c De Meye e al. / ZooKeys 540: 405–427 (2015)
410
Unlike he males, emales, a e mo phologically almos indis inguishable. Ce a i is
anonae emales di e om he o he wo axa in he pilosi y o he anepis e num and
o e emu . In C. anonae he anepis e num has some ew da k se ulae medio en ally,
and he o e emu has dispe sed sho da k se ulae be ween he en al se ae and pos-
e io ow o se ae. In C. osa and C. asci en is, he anepis e nal pilosi y is comple ely
pale and he o e emu usually only has pale se ulae p esen be ween he en al se ae
and pos e io ow o se ae (De Meye and F eidbe g 2006). Females o C. osa and C.
asci en is emales canno be eliably di e en ia ed on mo phological cha ac e s.
Dis ibu ion pa e ns h oughou A ica
The occu ence o he h ee species h oughou A ica shows di e en dis ibu ion pa -
e ns wi h only a pa ial o e lap be ween some o he species. Al hough some species
do occu sympa ically o pa apa ically, nowhe e do all h ee species co-occu . Ce a i is
osa is ound h oughou Sou h ( om wes e n Cape onwa ds) o Eas A ica wi h he
no he nmos eco ds om he Cen al Highlands in Kenya (Figu e 5). The la e oc-
cu ence seems o be a ecen expansion (Copeland and Wha on 2006) wi h i s p e i-
ous ex ension eaching ill he Kenyan coas only. I has been in oduced o he Indian
Ocean Islands o La Réunion and Mau i ius in he 20 h Cen u y (Whi e e al. 2000).
The single eco d o C. osa (one male) om Came oon in Wes A ica is p obably due
o mislabeling, while he p esence o he species in I o y Coas (N’dépo e al. 2010,
2013) is an e oneous eco d. No eliable eco ds om wes e n A ica ha e been ound.
Ce a i is asci en is has a wide dis ibu ion and is ound h oughou wes e n A ica,
wi h isola ed eco ds om Cen al A ica, and ex ensi e dis ibu ion along he Albe -
ine and G ego y Ri s in eas e n A ica, as a no h as E hiopia (Figu e 5). Ce a i is
anonae has a p edominan ly equa o ial bel dis ibu ion, being widesp ead h ough
Wes ( om Senegal onwa ds) and Cen al A ica (Figu e 6). I s eas e nmos dis ibu-
ion seems o be con ined o he wes e n side o he G ego y Ri in Kenya (Copeland
e al. 2006) ( eco ds u he o he eas need con i ma ion).
Sympa ic and allopa ic occu ence is ound be ween C. anonae and C. asci en is
in wes e n A ica (Vayssiè es e al. 2004), and wes e n Kenya (Copeland and Wha on
2006), while C. osa and C. asci en is occu oge he in pa s o Kenya and Tanzania
(Figu e 5) (Copeland e al. 2006, Mwa awala e al. 2006). Ce a i is anonae and C. osa
we e no ound oge he .
Indica ions o inconsis encies
Despi e he mo phological di e ences in adul males and he pa ially disjunc dis i-
bu ion, he e a e indica ions ha c yp ic specia ion occu s wi hin he h ee cu en ly
ecognized mo phospecies.
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 411
Figu e 5. Geog aphical dis ibu ion o Ce a i is osa (blue iangles) and C. asci en is (yellow ci cles).
Figu e 6. Geog aphical dis ibu ion o Ce a i is anonae.
Ma c De Meye e al. / ZooKeys 540: 405–427 (2015)
412
When compa ing sequences o mi ochond ial and nuclea ma ke s, Vi gilio e al.
(2008) could no eco e he h ee mo phospecies as monophyle ic g oups, al hough
di e en ma ke s eco e ed well-suppo ed clades wi hin he ep esen a i es o C. as-
ci en is om Wes and Eas A ica. So, al hough he molecula da a did no con adic
o suppo he mo phological sepa a ion, i did sugges ha C. asci en is is i sel a
complex o c yp ic species. This s udy, as well as Douglas and Hayme (2001) and
Ba e al. (2006) indica ed mo eo e he exis ence o o he sepa a e clus e s wi hin he
complex (al hough no always wi h high suppo ).
Finally, co ela i e ecological niche modeling showed ha C. osa p e e s clima ic
condi ions wi h lowe empe a u es when compa ed o C. capi a a (De Meye e al.
2008). This was co obo a ed by indings on he island o La Réunion whe e C. osa
occupies a colde and mo e humid clima e niche han C. capi a a (Duyck e al. 2006).
Howe e , hese esul s we e con adic ed by G ou and S ol z (2007) who s udied he
de elopmen al h esholds o a numbe o Ce a i is species, based on Sou h A ican
popula ions and concluded ha he si ua ion in Sou h A ica is di e en om ha in
La Réunion. This led o he sugges ion ha C. osa could comp ise en i ies wi h di e -
en biological equi emen s.
Objec i e
The inclusion o he Ce a i is FAR complex in he Coo dina ed Resea ch P ojec
(CRP) on c yp ic species, he e o e, was o es ablish whe he he h ee mo phological
en i ies ac ually ep esen h ee dis inc species, o i he ela ionship can be esol ed
in a di e en manne . The au ho s op ed o an in eg a i e app oach, using di e -
en me hodologies, including popula ion gene ics, la al mo phology, wing mo pho-
me ics, cu icula hyd oca bons, de elopmen al physiology and p e- and pos zygo ic
ma ing compa ibili y. The esul s o hese di e en app oaches a e p esen ed in his
olume o ha e been published elsewhe e (Vi gilio e al. 2013, Vaníčko á e al. 2014).
We he ewi h summa ize he main esul s ob ained by each o hese app oaches and
p o ide a syn hesis on he cu en knowledge and s a us o he di e en en i ies wi hin
he Ce a i is FAR complex.
Resul s
Mic osa elli e analysis
Using mic osa elli es, Bali aine e al. (2004) e ealed signi ican di e en ia ion wi h
espec o geno ypic equencies be ween C. osa specimens om he A ican main-
land and he Indian Ocean islands, and be ween C. asci en is om Kenya and om
Uganda. Dela e e al. (2013) isola ed and cha ac e ized 16 mic osa elli e ma ke s us-
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 419
Many ola iles iden i ied in Ce a i is FAR male emana ion ha e been p e iously
iden i ied in o he eph i id phe omones (C uz-López e al. 2006, Hea h e al. 1991,
Lu and Teal 2001, Mile -Pinhei o e al. 2014, Robacke 1988, Rocca e al. 1993,
Vaníčko á 2012, Vaníčko á e al. 2012). O he s, speci ically isop enoid ge anyl ac-
e one, alipha ic ola ile (E)-non-2-enal, and me hyl (2E,6E)- a nesoa e, ha e no been
epo ed be o e. In e es ingly, he la e is a c us acean ep oduc i e ho mone, s uc-
u ally simila o insec ju enile ho mone, which is esponsible o enhancing ep o-
duc i e ma u a ion, main aining ju enile mo phology, and in luencing male sex de e -
mina ion (Olms ead and LeBlanc 2007, Naga aju and Bo s 2008). In insec s, me hyl
(2E,6E)- a nesoa e ep esen s he immedia e p ecu so o insec ju enile ho mone III
(Teal e al. 2014). As a semiochemical, me hyl (2E,6E)- a nesoa e was epo ed in
pen a omid bug phe omones (Milla e al. 2002).
The phe omone composi ion as well as elec oan ennog aphy may be used o spe-
cies iden i ica ion. Simila ly as o he composi ion o CHCs (Vaníčko á e al. 2014), he
phe omone composi ion and an ennal speci ici y sugges ha he h ee nominal species o
he Ce a i is FAR complex co espond o axonomically well-de ined en i ies. I is ecom-
mended, howe e , ha his wo k should be expanded by including phe omone s udies
o F1 and R1 and u he sampling and analysis o di e en popula ions o all ypes, o
de e mine in e -popula ion a iabili y.
De elopmen al physiology
De elopmen al physiology s udies can assis in de ec ing di e ences be ween species
wi h ega d o biological equi emen s. Only he de elopmen al physiology o he wo
C. osa ypes was s udied in de ail (Tanga e al. 2015), because o he lack o es ablished
colonies o he o he species and because i was he di e ences obse ed be ween de-
elopmen al s udies conduc ed in La Réunion (Duyck and Quilici 2002) and Sou h
A ica (G ou and S olz 2007) ha ini ia ed he idea ha C. osa may consis o di e en
bio ypes wi h di e en clima ic equi emen s. R1 and R2 popula ions we e, he e o e,
s udied simul aneously in Kenya and Sou h A ica (Tanga e al. 2015). Depending on
locali y and empe a u e, ma ked di e ences we e obse ed be ween R1 and R2 popula-
ions in he de elopmen al du a ion o imma u e li e s ages. In bo h Kenyan and Sou h
A ican popula ions, R2 appea s o be less adap ed o ho e en i onmen s han R1. In
Kenya, R2 appea ed o be be e adap ed o colde en i onmen s, while in Sou h A -
ica, bo h C. osa ypes we e able o ole a e lowe empe a u es. Despi e disc epancies
in empe a u e ela ed de elopmen al physiology o he wo ypes in he wo locali ies,
esul s om Kenya and Sou h A ica clea ly demons a e and suppo he exis ence o
wo gene ically dis inc popula ions o C. osa ha a e di e gen in hei physiological
esponse o empe a u e. Disc epancies be ween he locali ies wi h ega ds o obse ed
and es ima ed de elopmen al ime pa ame e s o he wo C. osa ypes can be due o ge-
og aphic a ia ion (popula ions om Sou h A ica o igina ed om empe a e clima es
Ma c De Meye e al. / ZooKeys 540: 405–427 (2015)
420
and popula ions om Kenya o igina ed om opical clima es). O he ac o s such as
ood quan i y and quali y, ea ing condi ions, acclima iza ion and gene a ion age a e
also men ioned as ac o s in luencing he de elopmen al physiology.
Geog aphical dis ibu ion and al i udinal ansec
The dis ibu ion o C. asci en is and C. osa was e-analyzed, aking in o accoun he
exis ence o wo ypes o each o hese species. As he known dis ibu ion was la gely
based upon museum specimens collec ed o e a pe iod o 130 yea s and he DNA
e ie al om olde specimens (i.e. >10y s) is cumbe some and wi h low success a e,
i was decided o e-assign specimens based only on mo phological cha ac e s which
excluded emale specimens. In o al, specimens om 218 locali ies we e e-examined
and assigned o one o he ou ypes (F1, F2, R1, R2). The obse ed dis ibu ions
a e gi en in Figu es 10–11. F1 is mainly ep esen ed in wes e n A ica bu ex ends i s
dis ibu ion h oughou sou he n and eas e n pa s o he con inen wi h eco ds om
Angola, Zambia, Malawi and Tanzania (Figu e 10). F2 is con ined o he Ri a eas o
E hiopia, Kenya and eas e n pa o he Democ a ic Republic o Congo, wi h sou he n
expansion in o Ka anga egion o he Democ a ic Republic o Congo and Zambia. So
a , no sympa ic o pa apa ic occu ence o he wo ypes has been obse ed. R1 and
R2 on he o he hand do no show a clea geog aphic isola ion (Figu e 11). Only in
he Cape and cen al pa s o Sou h A ica is a single ype (R2) p esen , as well as in
he ad en i e popula ions on he Indian Ocean islands. In he no he n pa o Sou h
A ica and no hwa ds he e a e eco ds showing o e lap o dis ibu ions.
A de ailed s udy was conduc ed in he Ulugu u Moun ains nea Mo ogo o (Tan-
zania) whe e bo h ypes a e known o occu (Mwa awala e al. 2015). Along an al-
i udinal ansec , anging om 540 o 1650 masl he occu ence o bo h ypes was
moni o ed using aps wi h EGO-lu e. A g adual shi was obse ed wi h bo h ypes
occu ing a lowe al i udes (wi h p edominance o R1) while only R2 was obse ed
a he highes ele a ions. Geu s e al. (2012) obse ed a empe a u e shi o 7–8 °C
along his ansec . When looking a he de elopmen al di e ences obse ed o , a
leas he Kenyan popula ions, empe a u e could well play a majo ole in he obse ed
pa e n, al hough i emains o be seen whe he o he aspec s such as di e en ial hos
ange and a ailabili y also ha e an in luence ega ding he di e ences obse ed.
P e- and pos zygo ic incompa ibili y
Al hough no de ailed da a a e p esen ed in his olume, p elimina y da a by S. Ekesi
(pe s. communica ion), indica e ha in ield cage s udies he e is a signi ican p e- and
pos -zygo ic incompa ibili y bo h be ween Ce a i is osa and C. asci en is (F2), as
well as be ween he wo geno ypic clus e s o C. osa: R1 and R2. None o he o he
geno ypic clus e s o mo phospecies we e included.
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 421
Figu e 10. Geog aphical dis ibu ion o Ce a i is asci en is, F1 (b own ci cles), F2 (yellow ci cles), un-
assigned (whi e ci cles).
Conclusion
The majo i y o he esea ch app oaches discussed he e indica e ha he Ce a i is FAR
complex consis s o a leas he h ee ecognized mo phospecies, bu possibly o i e
di e en species. All me hodologies (excep la al mo phology) used con i m ha Ce -
a i is anonae, C. asci en is and C. osa a e well ecognized g oups. Ce a i is anonae is
a uni o m g oup showing no appa en mo phological o gene ic a iabili y and has a
well-de ined dis ibu ion ange. The wo o he en i ies consis each o wo sepa a e
g oups. Fo C. osa, he wo en i ies (called ‘R1’, ‘lowland’ o ‘ho osa’ on one hand,
and ‘R2’, ‘highland’ o ‘cold osa’ on he o he hand) can be dis inguished mo pho-
logically in males, as well as by o he means and demons a e a di e en de elopmen-
al physiology. They occu sympa ically in some egions, bu also show a disjunc
dis ibu ion ha appea s o be co ela ed wi h ambien empe a u e. I is concluded
ha bo h ypes should be conside ed as wo di e en species. Taxonomically, he ype
Ma c De Meye e al. / ZooKeys 540: 405–427 (2015)
422
ma e ial o C. osa belongs o he R1 ype, which means ha he R2 ype should be
conside ed as a new species, and a o mal desc ip ion will be published in he nea
u u e. Simila ly, he e iew indica es ha also C. asci en is ends o be composed o
wo en i ies. Ye , because he da a a e cu en ly insu icien o es ablish clea ly whe he
hey should also be conside ed wo di e en species o no , we cu en ly sugges o
main ain he wo ypes unde one and he same species.
Acknowledgemen s
We would like o hank he Join FAO/IAEA P og amme o ini ia ing he Coo di-
na ed Resea ch P ojec on “Resolu ion o c yp ic species complexes o eph i id pes s
o o e come cons ain s o SIT applica ion and in e na ional ade”, and o p o iding
suppo o he au ho s o conduc ing pa o he esea ch and o a end he esea ch
Figu e 11. Geog aphical dis ibu ion o Ce a i is osa, R1 ( ed ci cles), R2 (blue ci cles), sympa ic
occu ence (b own ci cles), un-assigned (whi e ci cles).
An in eg a i e app oach o un a el he Ce a i is FAR (Dip e a, Teph i idae)... 423
coo dina ion mee ings. Some o he da a p esen ed he e a e also he esul o inancial
suppo by he Belgian De elopmen Coope a ion ( h ough he F amewo k Ag ee-
men wi h he Royal Museum o Cen al A ica, RMCA) o Sokoine Uni e si y o
Ag icul u e, and by he Belgian Science Policy o he Join Expe imen al Molecula
Uni (JEMU) o he RMCA and he Royal Belgian Ins i u e o Na u al Sciences. The
de elopmen al physiology wo k on he wo C. osa ypes in Sou h A ica was unded
by Ci us Resea ch In e na ional.
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