Resea ch A icle / A ículo de In es igación
Desc ip ion o he spe ma hecae and es icles o T ia oma melanocephala
Nei a & Pin o, 1923 (Hemip e a: Redu iidae: T ia ominae)
Desc ipción de las espe ma ecas y es ículos de T ia oma melanocephala Nei a y Pin o, 1923
(Hemip e a: Redu iidae: T ia ominae)
Jade de Oli ei a¹* , Kaio Cesa Chaboli Ale i² , Juliana Damieli Nascimen o³ ,
Vagne José Mendonça4 , Rena o F ei as de A aújo5 and João A is eu da Rosa6
1Labo a ó io de En omologia em Saúde Pública, Depa amen o de Epidemiologia, Faculdade de Saúde Pública,
Uni e sidade de São Paulo, A . D . A naldo 715, São Paulo, SP, B asil. 2Labo a ó io Nacional e In e nacional
de Re e ência em Taxonomia de T ia omíneos, Ins i u o Oswaldo C uz (FIOCRUZ), A . B asil 4365, Pa ilhão
Rocha Lima, sala 505, 21040-360 Rio de Janei o, RJ, B asil. 3Ins i u o de Biologia, Uni e sidade Es adual
de Campinas (UNICAMP), Campinas, SP, B asil. 4Depa amen o de Pa asi ologia e Mic obiologia/ CCS,
Uni e sidade Fede al do Piauí (UFPI), Te esina, PI, B asil. 5Sec e a ia Es adual de Saúde da Bahia. Sal ado ,
BA, B asil. 6Faculdade de Ciências Fa macêu icas, Uni e sidade Es adual Paulis a “Júlio de Mesqui a Filho”
(UNESP), A a aqua a, SP, B asil. *jd [email p o ec ed]
ZooBank: u n:lsid:zoobank.o g:pub:AA780686-F113-486C-825D-7ADC0E6602B0
h ps://doi.o g/10.35249/ che.49.2.23.07
Abs ac . T ia oma melanocephala is an endemic species o B azil. Pheno ypic (ex e nal mo phology)
and molecula s udies we e pe o med o con ibu e o he axonomy and phylogeny o ia omines,
since ew s udies on he in e nal mo phology o hese we e ound. In o de o con ibu e o he
biological, mo phological and axonomic knowledge o he sub amily T ia ominae, his wo k
desc ibes he in e nal mo phology o he ep oduc i e appa a us o adul males and emales
o T. melanocephala. Fo his wo k 30 specimens om Jequié, Bahia, B azil we e used. T ia oma
melanocephala’s ep oduc i e sys em included slim spe ma hecae inse ions, la ge/medium ounded
e minal bodies, and i egula ellipsoid-shaped es icles wi h a anspa en , i eous appea ance.
These esul s con ibu e o he knowledge o he ep oduc i e biology o his species o ec o ial
impo ance and show axonomic ele ance, as well as possible applica ion in sys ema ic s udies.
Key wo ds: Chagas disease; gonads; in e nal mo phology; ia omine; ec o s.
Resumen. T ia oma melanocephala es una especie endémica de B asil. Se ealiza on es udios
eno ípicos (mo ología ex e na) y molecula es pa a con ibui a la axonomía y ilogenia de los
ia ominos, ya que se encon a on pocos es udios sob e la mo ología in e na de es os. Con el in
de con ibui al conocimien o biológico, mo ológico y axonómico de la sub amilia T ia ominae,
es e abajo mues a la mo ología in e na del apa a o ep oduc o de adul os machos y hemb as de
T. melanocephala. Pa a es e abajo se u iliza on 30 especímenes ecolec ados en Jequié, Bahía, B asil.
El sis ema ep oduc i o de T. melanocephala incluye inse ciones delgadas de espe ma ecas, cue pos
g andes/medianos con e minaciones edondeadas, y es ículos de o ma elipsoidal i egula con
una apa iencia anspa en e y i ea. Es os esul ados con ibuyen al conocimien o de la biología
ep oduc i a de es a especie de impo ancia ec o ial y mues an ele ancia axonómica, así como su
posible aplicación en es udios sis emá icos.
www.bio axa.o g/ ce. ISSN 0718-8994 (online) Re is a Chilena de En omología (2023) 49 (2): 271-278.
Es e es un a ículo de acceso abie o dis ibuido bajo los é minos de la licencia C ea i e Commons License (CC BY NC 4.0)
Recei ed 13 Feb ua y 2023 / Accep ed 11 Ap il 2023 / Published online 30 Ap il 2023
Responsible Edi o : José Mondaca E.
Oli ei a e al.: Desc ip ion o he spe ma hecae and es icles o T ia oma melanocephala.
272
Palab as cla e: En e medad de Chagas; gónadas; mo ología in e na; ia omino; ec o es.
In oduc ion
T ia oma melanocephala Nei a & Pin o, 1923 (Hemip e a: T ia ominae) is a wild ia omine
ha occu s in he s a es o Bahia, Pa aíba, Pe nambuco, Rio G ande do No e and Se gipe
(Gal ão e al. 2003; Gal ão 2014). Ch omosomal (Ale i e al. 2012, 2014a), meio ic (Ale i e
al. 2013), mo phological (She lock & Gui on 1980) and, mainly, phylogene ic (Jus i e al.
2014, 2016) da a demons a e ha his species, oge he wi h T ia oma i iceps (S ål, 1859),
o m he T. i iceps subcomplex (Ale i e al. 2017). I is belie ed ha species om his
subcomplex eached he A lan ic coas by dispe sal and di e si ied be o e he No he n
Andean upli (23–10 Ma) (Jus i e al. 2016).
The p o ozoan T ypanosoma c uzi (Chagas, 1909) (Kine oplas ida: T ypanosoma idae)
was isola ed om a specimen o T. melanocephala om Bahia (Ribei o e al. 2014). This
in o ma ion and he epo s o cap u e o his ia omine in human dwellings (She lock
& Gui on 1980; Gal ão 2014) emphasize i s epidemiological impo ance in he ec o ial
ansmission o Chagas disease (CD). In addi ion o ha , She lock and Gui on (1980)
sugges ha T. melanocephala is p obably an impo an link be ween he cycles o wild and
domes ic ansmission, ansmi ing T. c uzi om domes ic ese oi s o humans.
The way o minimize he incidence o CD is he con ol o ec o s, which equi es he
knowledge o he di e en biological aspec s o hese hema ophagous insec s. The i s
biological in o ma ion desc ibed o T. melanocephala was mo phological cha ac e is ics
(She lock & Gui on 1980). Cu en ly, se e al o he analyses such as ch omosome (Ale i
e al. 2012, 2014a; Pi a e al. 2021), meio ic (Ale i e al. 2013), cell (Ale i e al. 2016a, b),
spe mio axonomy (Ale i e al. 2014b), biological (Oli ei a e al. 2015), molecula (Ga dim e
al. 2014), epidemiological (Ribei o e al. 2014), ep oduc i e (Ale i e al. 2014c), e olu iona y
(Jus i e al. 2014, 2016; Ale i e al. 2017), mo phological and mo phome ic s udies (Oli ei a
e al. 2015) we e pe o med in T. melanocephala.
S udies on he in e nal ana omy o ia omines a e ela ed o he wo ks o Snodg ass
(1935), Da ey (1958), Lacombe (1965) and Ba h (1980). Snodg ass (1935) desc ibes he
ep oduc i e ac o emales o species o he class Insec a as consis ing o wo o a ies,
o med by smalle cylind ical g oups called o a ioles, wo la e al o iduc s leading o he
common o iduc , whe e he pai s o spe ma hecae and he accesso y gland a e inse ed
( he common o iduc al eady has an opening o he coupling bag). Fo i s pa , he
male ep oduc i e sys em consis s o a pai o es icles, la e al duc s, as de e ens, he
seminal esicle, accesso y glands, and he ejacula o y duc , which leads o he phallus, he
copula o y o gan o a h opods.
The main way o mi iga e he incidence o CD is h ough he con ol o ec o s and,
o his, i is necessa y o know he di e en biological aspec s o hese hema ophagous
insec s. Thus, he p esen s udy sough o p o ide an ana omical and mo phological
cha ac e iza ion o he male and emale ep oduc i e sys ems o T. melanocephala, wi h he
pu pose o knowing he ep oduc i e biology o his species.
Ma e ial and Me hods
The specimens o T. melanocephala used a e main ained in he Insec a ium o T ia ominae
o he School o Pha maceu ical Sciences/UNESP since Oc obe 8, 2009 (CTA 219). The
specimens o T. melanocephala ha o igina ed he colony we e collec ed in he pe idomicile
o he municipali y o Jequié, s a e o Bahia, by a eam o he Sec e a y o Heal h o he S a e
o Bahia (Sesab) and sen by Eliane Góes Nascimen o and Rena o F ei as de A aújo.
Re is a Chilena de En omología 49 (2) 2023
273
Mo phological s udy by op ical mic oscopy (MO)
Fo he obse a ion o he ep oduc i e appa a us by means o op ical mic oscopy, he
p ocedu es es ablished by Lacombe (1965) and Nascimen o e al. (2017) we e adop ed.
The images we e cap u ed unde a s e eoscopic mic oscope Leica MZ APO and Mo ic
Ad anced 3.2 plus an image analysis sys em. The numbe o insec s used was 15 male and
15 emale.
Mo phological s udy by scanning elec on mic oscopy (SEM)
Fo SEM obse a ions, he spe ma hecae and es is o T. melanocephala we e p e iously
p epa ed ollowing he me hods desc ibed by Nascimen o e al. (2017). The images we e
subsequen ly examined and eco ded ollowing Rosa e al. (1992) and Oli ei a e al. (2015).
Resul s
Op ical mic oscopy
The comple e emale ep oduc i e sys em o T. melanocephala has: wo o a ies, wo
la e al o iduc s, a common o iduc , wo spe ma hecae, an accesso y gland, and a geni al
chambe . The s udy o o a ies o adul emales o T. melanocephala showed ha hey a e
double and consis o se e al o a ioles, as i can be seen in Fig. 1A. The o a ies a e di ided
in o h ee pa s: e minal ilamen , o a ian o o a y ubes, and peduncle o chalice. The
e minal ilamen ends on he ho acic adipose body, p omo ing he posi ioning o he
o a y. The o a y is co e ed by a unic o i s own, which is lined ex e nally by he pe i oneal
memb ane ha goes om he la e al o iduc o he e minal ilamen , ha ing li le con ac
wi h he o a ies.
The spe ma hecae o T. melanocephala p esen ed a a ia ion in he posi ioning (Fig. 1)
and di e ences in size and shape depending on whe he hey a e ull o emp y o spe m
(Figs. 1B, C). Slim inse ions we e ound in he common o iduc , and he e we e la ge and
medium bodies wi h ounded e minal po ions (Figs. 1B, C).
Figu e 1. Female ep oduc i e sys em o adul T. melanocephala by op ical mic oscopy. A. O e iew
o he (o ) o a ioles magni ica ion 10X. B. Full spe ma hecae. C. Nea ly emp y spe ma hecae. (sp)
spe ma hecae Magni ica ion 40x. / Sis ema ep oduc i o emenino del adul o de T. melanocephala is o
po mic oscopia óp ica. A. Visión gene al de los (o ) o a iolos con aumen o de 10X. B. Espe ma ecas
llenas. C. Espe ma ecas casi acías. (sp) espe ma ecas. Aumen o de 40x.
The comple e male ep oduc i e sys em has: wo es icles, wo as de e ens, semini e ous
esicle, wo accesso y glands and ejacula o y duc . The es icles o T. melanocephala ha e
se en es icula ollicles ha a e encased by a pe i oneal shea h (Figs. 2A, B, C). They
p esen ed an i egula con ou , app oxima ely ellipsoid shaped, and a anspa en
colo a ion wi h sligh i eous lus e (Fig. 2A).
Oli ei a e al.: Desc ip ion o he spe ma hecae and es icles o T ia oma melanocephala.
274
Scanning Elec on Mic oscopy
The spe ma hecae o T. melanocephala showed a ounded shape wi h a ia ions when
hey a e ull o spe m. When emp y, hey p esen ed he same o ma in all samples analyzed
(Fig. 3).
Figu e 2. Male ep oduc i e sys em o adul male T. melanocephala by op ical mic oscopy. A. O e iew
o he (Te) es icles; magni ica ion 20X. B. O e iew o he (F ) ollicles es icula ; magni ica ion
20x. C. O e iew o he (du) duc us de e ens 20x; (ag) accesso y glands 20x and (s ) semini e ous
esicle; magni ica ion 20x. / Sis ema ep oduc i o masculino del adul o de T. melanocephala is o po
mic oscopía óp ica. A. Visión gene al de los (Te) es ículos; aumen o de 20x. B. Visión gene al de los
(F ) olículos es icula es; aumen o de 20x. C. Visión gene al del (du) conduc o de e en e; 20x; (ag)
glándulas acceso ias 20x y (s ) esícula seminal; aumen o de 20x.
Figu e 3. Female ep oduc i e sys em by scanning elec on mic oscopy (SEM). A. Spe ma hecae
adul emale T. melanocephala: (sp) spe ma hecae; (R¹ and R²) egion whe e he cu s o assembling
he pa s we e made whe e wo o iduc s side and geni al second came a (Ramí ez Pé ez 1969). B.
De ail o spe ma hecae. / Sis ema ep oduc i o emenino obse ado po mic oscopía elec ónica de
ba ido (MEB). A. Espe ma ecas de la hemb a adul a de T. melanocephala: (sp) espe ma ecas; (R¹ y R²)
egión donde se ealiza on los co es pa a el ensamblaje de las pa es donde se encuen an los dos
o iduc os la e ales y la segunda cáma a geni al (Ramí ez Pé ez 1969). B. De alle de las espe ma ecas.
The es icles o T. melanocephala p esen ed a ounded shape, ha ing a di e en ia ion in
he egions o connec ion wi h an e io as de e ens and e minal ilamen (Fig. 4).
Re is a Chilena de En omología 49 (2) 2023
275
Discussion
Analyzing he emale ep oduc i e sys em o Rhodnius S ål, 1859, Ramí ez Pé ez (1969)
epo s ha he emale R. p olixus S ål, 1859 p esen s in i s adul phase a pai o o a ies
loca ed on ei he side o he diges i e ac and a common o iduc ha bi u ca es an e io ly
o ming he la e al o iduc s, he same being obse ed in T. melanocephala.
When compa ed o T. in es ans (Klug, 1834) (Nascimen o e al. 2017), T. melanocephala
p esen ed a di e ence in he numbe o o a ies and hei size. Al hough new species
should be s udied, we sugges ha his s uc u e o he emale ep oduc i e sys em may
be a axonomic ool o di e en ia e species, as well as i was obse ed ha he size o he
es icula ollicles ep esen s an impo an ool o di e en ia e he gene a o he sub amily
T ia ominae (Ale i e al. 2015).
Snodg ass (1935) men ions ha he o ma ion o he spe ma ic ubes has a simila o igin
o he o ma ion o he eggs in he o iduc s o he emales, hus hey a e denomina ed
es icula ollicles. The accesso y glands p oduce an opaque g anula sec e ion esponsible
o he mo emen o he spe ma ozoa in he emale, which oge he gi e ise o he
spe ma opho e.
All ia omine species ha e se en es icula ollicles (Gonçal es e al. 1987). This
synapomo phy is no sha ed wi h o he Hemip e a ha p esen a ia ion in he numbe o
ollicles (Pe ei a e al. 2015). The es icula ollicles consis o a hin anslucen memb ane
ha allows he isualiza ion o he in e nal cells (Gonçal es e al. 1987). Tes icula ollicle
lining cells a e known as cys ic cells and a e esponsible o nou ishing cells in cell di ision
du ing spe ma ogenesis (Schmid & Do n 2004). In ia omines, spe ma ogenesis is cys ic,
ha is, he cell di isions happen synch onized wi hin he spe ma ogonial cys s (Silis ino-
Souza e al. 2012; Ale i e al. 2015).
The colo ing o he pe i oneal shea h o he es es was cha ac e ized as a synapomo phy
o T ia ominae (Ale i e al. 2014c). The anspa en colo a ion obse ed o all he species
analyzed by he au ho s is ex emely di e en om o he Hemip e a ha ha e mul icolo ed
shea hs ( eddish, o ange, yellowish o anspa en ) (Gomes e al. 2013). The es icles o T.
melanocephala p esen ed an i egula ou line, wi h an app oxima ely ellipsoid shape and
a anspa en shea h, which co obo a es he desc ip ion p oposed by Ale i e al. (2014c).
Figu e 4. Tes icles o adul male T. melanocephala by (A) Tes icle do sal iew (B) Tes icle en al iew
by scanning elec on mic oscopy. / Tes ículos del adul o de T. melanocephala obse ados po (A) is a
do sal del es ículo (B) is a en al del es ículo median e mic oscopía elec ónica de ba ido.
Oli ei a e al.: Desc ip ion o he spe ma hecae and es icles o T ia oma melanocephala.
276
The spe ma hecae is a compa men esponsible o s o ing he semen eleased a he
ime o copula ion, un il he oocy e is de eloped. La e he oocy e descends h ough he
la e al o iduc , which has se e al olds na u ally o allow a good dila ion a he ime o
o ula ion, when he egg passes h ough he geni al chambe and he spe ma hecae injec s
he s o ed spe m, hus e ilizing he egg (Ba h 1973; Ramí ez Pé ez 1969).
Compa ing T. melanocephala wi h he species s udied by Nascimen o e al. (2017), i is
possible o obse e a simila i y in he mo phology o he spe ma hecae o his species wi h
T. in es ans and di e ences wi h T. b asiliensis Nei a, 1911, T. juazei ensis Cos a & Felix, 2007,
T. she locki Papa e al., 2002 and P. ibiamacula us (Pin o, 1926). These esul s a e impo an
because T. melanocephala was ini ially g ouped wi h T. b asiliensis, T. juazei ensis, T. she locki
and P. ibiamacula us in he T. b asiliensis subcomplex (Scho ield & Gal ão 2009). Howe e ,
i is cu en ly known ha hese species a e no phylogene ically ela ed (Ale i e al. 2012;
Ga dim e al. 2014; Jus i e al. 2014, 2016; Oli ei a e al. 2017), which indica es ha he
analysis o he spe ma heca in SEM has axonomic alue.
The e o e, he pa ame e s o he emale and male ep oduc i e sys ems analyzed in
adul s o T. melanocephala ha e con ibu ed o he knowledge o he ep oduc i e biology
o his species o ec o ial impo ance and ha e showed axonomic ele ance, as well as
possible applica ion in sys ema ic s udies.
Funding
This wo k was suppo ed by he Founda ion o Resea ch Suppo o he S a e o São
Paulo (FAPESP) (P ocess numbe 2010/15386-3).
Acknowledgmen s
Thanks o he Sec e a y o Heal h o he S a e o Bahia (SESAB) and Eliane Góes
Nascimen o o sending he specimens ha o igina e he colony.
Li e a u e Ci ed
Ale i, K.C., Mendonça, P.P., Pe ei a, N.P., Rosa, J.A. and Oli ei a, M.T. (2012) Ka yo ype
o T ia oma melanocephala Nei a and Pin o (1923). Does his species i in he B asiliensis
subcomplex?. In ec ion Gene ic and E olu ion, 12: 1652-1653.
Ale i, K.C.C., Mendonça, P.P., Pe ei a, N.P., Rosa, J.A. and Aze edo-Oli ei a, M.T.V.
(2013) Spe ma ogenesis in T ia oma melanocephala (Hemip e a: T ia ominae). Gene ics
and Molecula Resea ch, 12: 4944-4947.
Ale i, K.C.C., Rosa, J.A. and Aze edo-Oli ei a, M.T.V. (2014a) Dis ibu ion o cons i u i e
he e och oma in in T ia oma melanocephala (Hemip e a, T ia ominae). Gene ics and
Molecula Resea ch, 13: 7899-7903.
Ale i, K.C.C., Mendonça, P.P., Pe ei a, N.P., Rosa, J.A. and Aze edo-Oli ei a, M.T.V.
(2014b) He e opykno ic ilamen in spe ma ids o T ia oma melanocephala and T. i iceps
(Hemip e a, T ia ominae). In e eb a e Rep oduc ion & De elopmen , 58: 9-12.
Ale i, K.C.C., Oli ei a, J., Rosa, J.A. and Aze edo-Oli ei a, M.T.V. (2014c) Colo a ion
o he es icula pe i oneal shea h as a synapomo phy o ia omines (Hemip e a,
Redu iidae). Bio a Neo opical, 14: 1-3.
Ale i, K.C.C. (2015a) Mo phology o es icula ollicles as axonomic ool in he sub amily
T ia ominae. En omology O ni hology & He pe ology, 4: e100.
Ale i, K.C.C., Cas o, N.F.C., Oli ei a, J., Rosa, J.A. and Aze edo-Oli ei a, M.T.V.
(2015) Cys ic spe ma ogenesis in h ee species o he p olixus complex (Hemip e a:
T ia ominae). I alian Jou nal o Zoology, 82: 172-178.
Re is a Chilena de En omología 49 (2) 2023
277
Ale i, K.C.C., Nascimen o, J.G.O., Mo ei a, F.F.F., Ju be g, J. and Aze edo-Oli ei a, M.T.V.
(2016a) Analysis o me abolic ac i i y in cys ic cells o T ia oma ub o ascia a (Hemip e a:
T ia ominae) and i s capaci y o occupy di e en en i onmen s. A ican En omology, 24(1):
261-264.
Ale i, K.C.C., Nascimen o, J.G.O., Mo ei a, F.F.F., Ju be g, J. and Aze edo-Oli ei a, M.T.V.
(2016b) Cy ogene ic cha ac e isa ion o T ia oma ub o ascia a (De Gee ) (Hemip e a,
T ia ominae) spe ma ocy es and i s cy o axonomic applica ion. A ican En omology,
24(1): 257-260.
Ale i, K.C.C., Oli ei a, J., Aze edo-Oli ei a, M.T.V. and Rosa, J.A. (2017) T ia oma i iceps
subcomplex (Hemip e a, Redu iidae, T ia ominae): A new g ouping o Chagas disease
ec o s om Sou h Ame ica. Pa asi es & Vec o s, 10: 180.
Ba h, R. (1973) Es udo ana ômico e his ológico sob e a sub amilia T ia ominae (He e op e a,
Redu iidae). Pa e XXIII: O o á io de T ia oma in es ans. Memo ias do Ins i u o Oswaldo
C uz, 71: 123-147.
Ba h, R. (1980) A glândula ec ode mal do apa elho copulado do T ia oma in es ans
(He e op e a, T ia ominae). Memo ias do Ins i u o Oswaldo C uz, 75: 119-124.
Da ey, K.G. (1958) The mig a ion o spe ma ozoa in he emale o Rhodnius p olixus S al. J.
Expe imen al Biology, 35: 694-701.
Gal ão, C. (2014) Ve o es da Doença de Chagas no B asil. Scielo books: 289 ISBN 978-85-
98203-09-6.
Gal ão, C., Ca ca allo, R., Rocha, D.S. and Ju be g, J. (2003) A checklis o he cu en
alid species o he sub amily T ia ominae Jeannel, 1919 (Hemip e a, Redu iidae) and
hei geog aphical dis ibu ion, wi h nomencla u al and axonomic no es. Zoo axa, 202:
1-36.
Ga dim, S., Almeida, C.E., Takiya, D.M., Oli ei a, J., A aújo, R.F., Cica elli, R.M. and da
Rosa, J.A. (2014) Mul iple mi ochond ial genes o some syl a ic B azilian T ia oma: non-
monophyly o he T. b asiliensis subcomplex and he need o a gene ic e ision in he
T ia omini. In ec ion, Gene ics and E olu ion, 23: 74-79.
Gomes, M.O., Cas anhole, M.M.U., Souza, H.V., Mu akami, A.S., Fi mino, T.S.S., Sa an,
P.S., Bahno, C.A., Mon ei o, L.S., Sil a, J.C.P. and I oyama, M.M. (2013) Mo phological
aspec s o he es es o 18 species o e es ial He e op e a om No hwes e n São
Paulo (B azil). Bio a Neo opica, 13(3): 131-135.
Gonçal es, T.C.M., He man, L. and Almeida, J.R. (1987) Es udo ana ômico e mo omé ico
dos olículos es icula es de algumas espécies de T ia ominae (Hemip e a: Redu iidae).
Memo ias do Ins i u o Oswaldo C uz, 82(4): 543-550.
Jus i, S.A., Gal ão, C. and Sch ago, C.G. (2016) Geological changes o he Ame icas
and hei in luence on he di e si ica ion o he Neo opical kissing bugs (Hemip e a:
Redu iidae: T ia ominae). PLOS Neglec ed T opical Diseases, 10(4): e0004527.
Jus i, S.A., Russo, C.A.M., Malle , J.R.D.S., Oba a, M.T. and Gal ão, C. (2014) Molecula
phylogeny o T ia omini (Hemip e a: Redu iidae: T ia ominae). Pa asi es & Vec o s, 7:
149.
Lacombe, D. (1965) Es udos ana ômicos e his ológicos sob e a sub amília T ia ominae
(He e op e a, Redu iidae). Pa e XXI. Es udos compa ado do sis ema aqueal em
T ia oma, Pans ongylus e Rhodnius. Memo ias do Ins i u o Oswaldo C uz, 63: 75-105.
Nascimen o, J.D., Ribei o, A.R., Almeida, L.A., Oli ei a, J., Mendonça, V.J., Cilense, M.
and Rosa, J.A. (2017) Mo phology o he spe ma hecae o wel e species o T ia ominae
(Hemip e a, Redu iidae) ec o s o Chagas disease. Ac a T opica, 176: 440-445.
Oli ei a, J., Mendonça, V.J., De A aújo, R.F., Nascimen o, E.G. and Da Rosa, J.A. (2015)
Biological, mo phological and mo phome ic s udies o T ia oma melanocephala Nei a &
Pin o, 1923 (Hemip e a: Redu iidae: T ia ominae). Zoo axa, 4012(3): 514-524.
Oli ei a e al.: Desc ip ion o he spe ma hecae and es icles o T ia oma melanocephala.
278
Oli ei a, J., Ma ce , P.L., Takiya, D.M., Mendonça, V.J., Belin ani, T., Ba gues, M.D.,
Ma eo, L., Chagas, V., Ramos, P.F., Es ela, P.C., Gonçal es, R.G., Cos a, J. and Almeida,
C.E. (2017) Combined phylogene ic and mo phome ic in o ma ion o delimi and uni y
he T ia oma b asiliensis species complex and he B asiliensis subcomplex. Ac a T opica,
170: 140-148.
Pe ei a, L.L.V., Ale i, K.C.C., Cas anhole, M.M., Mo ei a, F.F.F., Ba bo sa, J.F. and
I oyama, M.M. (2015) Cy ogene ics analysis and es is mo phology o aqua ic species
o he amilies Belos oma idae, Gelas oco idae, Ge idae, No onec idae and Veliidae
(He e op e a). Jou nal o Insec Science, 15(21): 1-10.
Pi a, S., Lo i e, P., Cuad ado, A., Panze a, Y., Oli ei a, J., Ale i, K.C.C., Rosa, J.A.,
F ei as, S.P.C., Gómez-Palacio, A., Sola i, A., Mon oy, C., Do n, P.L., Cab e a-B a o,
M. and Panze a, F. (2022) High ch omosomal mobili y o DNA clus e s in holocen ic
ch omosomes o T ia ominae, ec o s o Chagas disease (Hemip e a-Redu iidae).
Medical and Ve e ina y En omology, 36: 66-80.
Ramí ez Pé ez, J. (1969) Es udio sob e la ana omía de Rhodnius p olixus. Re is a Venezolana
de Sanidad y Asis encia Social, 34: 9-98.
Ribei o, A.R., Mendonça, V.J., Al es, R.T., Ma inez, I., A aújo, R.F., Mello, F. and da
Rosa, J.A. (2014) T ypanosoma c uzi s ains om ia omine collec ed in Bahia and Rio
G ande do Sul, B azil. Re is a de Saúde Pública, 48: 295-302.
Rosa, J.A., Ba a a, J.M.S. and Ba elli, N. (1992) Spi acles o 5 h ins a nymphs in six species
o T ia ominae (Hemip e a, Redu iidae) using scanning elec on mic oscopy. Memo ias
do Ins i u o Oswaldo C uz, 87: 301-302.
Schmid , E.D. and Do n, A. (2004) S uc u al pola i y and dynamics o male ge mline s em
cells in he milkweed bug (Oncopel us ascia us). Cell and Tissue Resea ch, 318: 383-394.
Scho ield, C.J. and Gal ão, C. (2009) Classi ica ion, e olu ion, and species g oups wi hin
he T ia ominae. Ac a T opica, 110(2-3): 88-100.
She lock, Í.A. and Gui on, N. (1980) Fauna T i ominae do Es ado da Bahia, B asil: IV -
T ia oma melanocephala Nei a & Pin o, 1923. Memo ias do Ins i u o Oswaldo C uz, 75(3-4):
23-31.
Silis ino-Souza, R., Pe uque i, R.L., Taboga, S.R. and Aze edo-Oli ei a, M.T.V.
(2012) Ch oma oid body: Remnan s o nucleola p o eins du ing spe ma ogenesis in
T ia omine (He e op e a, T ia ominae). Mic on, 43: 954-960.
Snodg ass, R.E. (1935) P inciples o insec mo phology. Ed. McG aw-Hill, New Yo k and
London. 552 pp.
