he jou nal o
biodi e si y da a
NOTES ON GEOGRAPHIC DISTRIBUTION
1143
Academic edi o : Robe o Leonan No aes
Recei ed: 8 Sep embe 2025
Accep ed: 13 No embe 2025
Published: 24 No embe 2025
Copy igh © The au ho s. This is an open‑access
a icle dis ibu ed unde e ms o he C ea i e Commons
A ibu ion License (A ibu ion 4.0 In e na ional – CC
BY 4.0)
Abs ac . We epo he i s eco d o His io us diaphanop e us Feijó, Rocha & Al ho , 2015 in he s a e
o Tocan ins, B azil and he hi d eco d o he species in he Ce ado biome, ein o cing i s associa ion
wi h seasonally d y en i onmen s o he Sou h Ame ican diagonal o open o ma ions. Mo phological ai s
con i m specimen iden i y, wi h anspa en wings as he mos dis inc i e diagnos ic cha ac e . Ou indings
highligh he species’ ecological speci ici y and exposu e o an h opogenic h ea s, including habi a loss
and clima e change, unde sco ing he need o u he s udies and in eg a ed conse a ion s a egies.
Key wo ds. Ce ado biome, global biodi e si y ho spo s, Neo opical Sa anna, ange ex ension, Sou h
Ame ica, anspa en wings
Gen il MA, Ho mann GS, Coelho IP, Co dei o JLP, de Oli ei a MB (2025) Fi s eco d o His io us diaphan-
op e us Feijó, Rocha & Al ho , 2015 (Chi op e a, Vespe ilionidae) in he s a e o Tocan ins, B azil. Check
Lis 21 (6): 1143–1152. h ps://doi.o g/10.15560/21.6.1143
In oduc ion
The amily Vespe ilionidae cu en ly comp ises 545 species a anged in o 62 gene a and ou sub am-
ilies: Vespe ilioninae, Ke i oulinae, Mu ininae, and Myo inae (Simmons and Ci anello 2025). Nine
espe ilionid gene a occu in he Neo opics: Baue us Van Gelde , 1959, Co yno hinus H. Allen, 1865,
Ep esicus Ra inesque, 1820, His io us Ge ais, 1856, Lasiu us G ay, 1831, Myo is Kaup, 1829, Neoep esicus
Cláudio e al., 2023, Pe imyo is Menu, 1984, and Rhogeessa H. Allen, 1866 (Wilson and Reede 2005;
Ga dne 2007; Cláudio e al. 2023).
His io us is cu en ly ep esen ed by 11 species (Simmons and Ci anello 2025), all endemic o
Sou h Ame ica and b oadly dis ibu ed ac oss he con inen , om no he n Venezuela o he sou he n
egions o Chile and A gen ina. I s ange ex ends om wes e n Venezuela and Colombia sou hwa d
h ough Ecuado , Pe u, and Boli ia o Pa aguay and no hwes e n A gen ina (Wilson and Reede 2005;
Ga dne 2007; Cláudio e al. 2023).
Fi e species a e known o occu in B azil (Ga bino e al. 2024): His io us alienus Thomas, 1916, H.
diaphanop e us Feijó, Rocha & Al ho , 2015, H. laepho is Thomas, 1916, H. mon anus (Philippi & Land-
beck, 1861), and H. ela us (I. Geo oy, 1824). These species a e b oadly dis ibu ed excep o he
Amazon, wi h eco ds om he No heas e n, ac oss he Cen al-Wes e n, o he Sou heas e n and
Sou he n egions, eaching as a as he s a e o Rio G ande do Sul (Ga dne 2007; Cláudio e al. 2023).
Species o his genus a e medium-sized o la ge ba s cha ac e ized by a dis inc i e combina ion
o mo phological ai s, including long ea s connec ed by a band o skin wi h a iable de elopmen ;
skull is delica e, na ow, and long; sagi al and lambdoidal c es s weakly de eloped; pos o bi al p o-
cess o he jugal well de eloped; and usually long do sal u (Reis e al. 2017; Cláudio e al. 2023).
21 (6) · h ps://doi.o g/10.15560/21.6.1143
21 (6): 1143–1152. h ps://doi.o g/10.15560/21.6.1143
Fi s eco d o His io us diaphanop e us Feijó, Rocha & Al ho , 2015
(Chi op e a, Vespe ilionidae) in he s a e o Tocan ins, B azil
Ma iane de Almeida Gen il1, 2
, Gab iel Selbach Ho mann3, Igo P ei e Coelho4, José Luís Passos Co dei o3, 5
,
Ma cioneB i ode Oli ei a2, 6
1 Ins i u o de Biociências, Uni e sidade Fede al do Es ado do Rio de Janei o, Rio de Janei o, RJ, B azil
2 Se o de Mas ozoologia, Depa amen o de Ve eb ados, Museu Nacional, Uni e sidade Fede al do Rio de Janei o, Rio de Janei o, RJ, B azil
3 In e na ional Pla o m o Science, Technology and Inno a ion in Heal h (PICTIS), A ei o, Po ugal
4 Independen Resea che , Flo ianópolis, SC, B azil
5 Ins i u o Oswaldo C uz, Fundação Oswaldo C uz, Eusébio, CE, B azil
6 Labo a ó io de Biologia e Pa asi ologia de Mamí e os Sil es es Rese a ó ios, Ins i u o Oswaldo C uz, Fundação Oswaldo C uz, Rio de
Janei o, RJ, B azil
Co esponding au ho : Ma cione B. de Oli ei a (oli ei a01ma [email p o ec ed])
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1144
His io us species a e a ely cap u ed in he ield and a e poo ly ep esen ed in scien i ic collec ions.
Fo ins ance, in he Mammal Collec ion o he Museu Nacional a Uni e sidade Fede al do Rio de
Janei o, B azil, he e a e only 32 specimens o he genus, o which 28 a e ca aloged, including a single
specimen o H. diaphanop e us documen ed in he p esen s udy. Consequen ly, His io us emains one
o he leas -known gene a o Neo opical ba s, al hough ecen s udies ha e e ealed he exis ence
o new species wi hin he genus (Feijó e al. 2015; Rod íguez-Posada e al. 2021; Velazco e al. 2021).
His io us diaphanop e us was desc ibed by Feijó e al. (2015), wi h he ype locali y in Boquei ão da
Onça, municipali y o Sen o Sé, s a e o Bahia, wi hin he B azilian Caa inga. In he same s udy, he
species was also epo ed om ou addi ional locali ies, h ee in he Caa inga and Ce ado biomes o
B azil, including he s a es o Pa aíba (São José dos Co dei os), Cea á (Fo aleza), and Ma anhão (Tas-
so F agoso); and one in he Chaco eco egion o Boli ia, depa men o San a C uz (Pampag ande). The
dis ibu ion o he species was la e expanded wi h new eco ds in he Ce ado biome om he s a e o
Ma o G osso (Cuiabá) and wi h addi ional eco ds in he Chaco eco egion o San a C uz, Boli ia (Valle
G ande; Semedo and Feijó 2016). Recen ly, he species was epo ed om he s a e o Pe nambuco,
wi h eco ds om wo municipali ies, Buíque and Tupana inga, wi hin he Caa inga biome (Be na d
e al. 2023).
He e, we epo a ange ex ension o H. diaphanop e us o Tocan ins, ep esen ing he hi d eco d
o he species in he Ce ado biome o B azil. We also p o ide an upda ed dis ibu ion map and a able
summa izing ex e nal and c anial measu emen s o all known specimens.
Figu e 1. Reco ds o His io us diaphanop e us in
B azil and Boli ia (see also Table 1). This s udy:
1 = Tocan ins, Liza da; Li e a u e eco ds: 2 =
Bahia, Sen o Sé, Boquei ão da Onça ( ype lo-
cali y), 3 = Cea á, Fo aleza, Faculdade Ve e i-
ná ia do Cea á, 4 = Ma anhão, Tasso F agoso,
T anquei a, 5 = Ma o G osso, Cuiabá, Chapada
dos Guima ães, Escola E angélica Bu i , 6 =
Pa aíba, São José dos Co dei os, Rese a Pa -
icula do Pa imônio Na u al Fazenda Almas,
7 = Pe nambuco, Buíque, Pa que Nacional
do Ca imbau, 8 = Pe nambuco, Tupana inga,
Pa que Nacional do Ca imbau, 9 = San a C uz,
Valle G ande, Po e illos, and 10 = San a C uz,
Pampag ande, Flo ida.
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1145
S udy a ea
The new eco d o H. diaphanop e us was made in he Liza da municipali y, Tocan ins, B azil (Figu e
1). This egion has a opical clima e wi h a d y win e (Aw ype on he Köppen classi ica ion), cha -
ac e is ic o opical sa annas (Peel e al. 2007). The ege a ion co esponds o Ce ado upes e, a
phy ophysiognomy associa ed wi h ocky ou c ops o qua zi e–a eni e (Ribei o and Wal e 1998).
This en i onmen is cha ac e ized by shallow, s ony soils wi h low nu ien a ailabili y and high acid-
i y, esul ing in es ic i e edaphic condi ions o ege a ion de elopmen . The ege a ion co e is
p edominan ly he baceous-sh ubby, spa se and discon inuous, in e spe sed wi h small woody spe-
cies adap ed o nu ien -poo subs a es and seasonal d ough . Such a eas ha bo a specialized lo a,
wi h high le els o endemism and unc ional ai s ha enhance esilience o i e and soil limi a ions
(Ribei o and Wal e 1998).
Me hods
In Ap il 2019, we ins alled wo sampling se s o i e mis ne s: one wi hin he ege a ion in on o
a qua z–a eni e ocky o ma ion, app oxima ely 120 m om he ou c op, and he o he along an
abandoned oad abou 100 m om he i s se . Ne s we e opened a sunse and emained ac i e o
six hou s. Inspec ions we e ca ied ou e e y 20 minu es. All cap u ed specimens we e placed in indi-
idual holding bags and ield anno a ions included o ea m leng h, body mass, sex, and ep oduc i e
condi ion. P elimina y iden i ica ion was conduc ed in he ield using axonomic keys (Reis e al. 2017).
We measu ed he specimens in he labo a o y using digi al calipe s (0.01 mm o p ecision) acco d-
ing o Feijó e al. (2015) and Semedo and Feijó (2016; Table 2). Ex e nal measu emen s: weigh (W),
o al leng h (TL), head–body leng h (HBL), ail leng h (T), hind- oo leng h (HF), ea leng h (E), heigh
o skin be ween ea s (HSE), o ea m leng h (FA), agus leng h (T ), calca leng h (CaL), ea b ead h
(EB). C anial measu emen s: g ea es leng h o skull (GLS), condyloincisi e leng h (CIL), pos o bi al
cons ic ion (PC), zygoma ic b ead h (ZB), b aincase b ead h (BB), mas oid b ead h (MB), heigh o
pos o bi al p ocess (HPP), pala al leng h (PL), maxilla y oo h ow leng h (C-M³), pala al b ead h (PB),
b ead h ac oss M3–M3 (M³-M³), b ead h ac oss canines (C-C), mandible leng h (ML), mandibula oo-
h ow leng h (C-M₃), mandible heigh (MH).
Resul s
His io us diaphanop e us Feijó, Rocha & Al ho , 2015
Figu es 2, 3
New eco ds. BRAZIL – Tocan ins • Liza da municipali y; −9.5733, −46.8104; 362 m a.s.l.; 22.IV.2019;
de Oli ei a MB; 1 adul ♀, MN84764.
Table 1. Geog aphic eco ds o His io us diaphanop e us in B azil and Boli ia, including coun y, s a e/depa men , locali y, coo -
dina es, associa ed ege a ion domain, and e e ences. The new eco d om Tocan ins (p esen s udy) ep esen s he no he n-
mos occu ence o he species in he Ce ado biome. No. = numbe ela i e o he loca ions in Figu e 1.
No. Coun y S a e /
depa men Locali y La i ude Longi ude Vege-
a ion Re e ence
1B azil Tocan ins Liza da −9.5733 −46.8104 Ce ado P esen s udy
2B azil Bahia Sen o Sé, Boquei ão da
Onça
−9.8803 −41.0709 Caa inga Feijó e al. 2015; ICMBio 2025
3B azil Cea á Fo aleza, Faculdade Ve e -
iná ia do Cea á
−3.7860 −38.5525 Caa inga Feijó e al. 2015; ICMBio 2025
4B azil Ma anhão Tasso F agoso, T anquei a −8.4725 −45.7428 Ce ado Feijó e al. 2015; ICMBio 2025
5B azil Ma o G osso Cuiabá, Chapada dos Gui-
ma ães, Escola E angélica
Bu i i
−15.4156 −55.8071 Ce ado Semedo and Feijó 2016;
ICMBio 2025
6 B azil Pa aíba São José dos Co dei os, Res-
e a Pa icula Pa imônio
Na u al Fazenda Almas
−7.4708 −36.8975 Caa inga Feijó e al. 2015; ICMBio 2025
7 B azil Pe nambuco Buíque, Pa que Nacional do
Ca imbau
−8.4791 −37.2369 Caa inga Be na d e al. 2023; ICMBio
2025
8B azil Pe nambuco Tupana inga, Pa que
Nacional do Ca imbau
−8.4282 −37.3096 Caa inga Be na d e al. 2023; ICMBio
2025
9 Boli ia San a C uz Valle G ande, Po e illos −18.5500 −64.9333 Chaco Semedo and Feijó 2016;
Poma-U ey e al. 2023
10 Boli ia San a C uz Pampag ande, Flo ida −17.9297 −64.1633 Chaco Feijó e al. 2015; Poma−U ey
e al. 2023
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1146
Iden i ica ion. The specimen weighs 8 g, wi h a head–body leng h o 110.86 mm, ail leng h 50.95
mm, o ea m leng h 46.74 mm, and ea leng h 28.01 mm (Table 2). The ea s a e conspicuously la ge,
anslucen , connec ed by a high band ac oss he o ehead (4.62 mm), and wi h he agus ex ending
o abou hal hei heigh (Figu e 3A, C). Wings and u opa agium a e anslucen and well de eloped,
wi h he ail sligh ly exceeding he dis al ma gin o he u opa agium (Figu e 3B, C). The do sal u is
long (9.5 mm) and dis inc ly bicolo ed, wi h do sal da k b own hai s a he base and golden a he
ips, and en al da k b own u a he base wi h whi ish ips (Figu e 3B, C). C anially, he inne uppe
inciso s a e widely spaced; he ou e inciso s a e small, in con ac wi h he inne inciso s and sligh ly
sepa a ed om he canines. The uppe M³ is na ow, abou hal he wid h o M¹ and M² (Figu e 2).
Den al o mula: i: 2/3; c: 1/1; pm: 1/2; m: 3/3 = 32.
Compa ison. Compa ison wi h H. ela us, which occu s in B azil and may also be p esen in he s udy
egion. His io us diaphanop e us and H. ela us a e he mos simila species wi hin he genus, sha ing a
iangula ea ( ine and acu e) wi h a p ominen inne lobe connec ed by a band ac oss he o ehead.
Howe e , hey di e in a combina ion o ex e nal and c anial ai s. His io us diaphanop e us exhibi s
well-de ined bicolo hai s on he do sum (wi h hai s ipped in pale), whe eas H. ela us ends o ha e a
sub ly bicolo ed do sum. The wing memb anes o H. diaphanop e us a e pale and anslucen (simila
o hose o H. laepho is) in con as o he da ke memb anes o H. ela us. C anial and body measu e-
men s u he suppo hei dis inc ion. Al hough he o e all skull shape is simila , H. diaphanop e us
Figu e 2. Do sal, en al, and la e al iews o
he skull and mandible o His io us diaphan-
op e us (MN84764).
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1147
is gene ally smalle han H. ela us in se e al dimensions, including o al body and ail leng h, skull
leng h, b aincase wid h, mas oid wid h, wid h ac oss M³, and mandibula leng h. In he examined
specimen, he combina ion o anslucen wings, conspicuously bicolo ed do sum, p opo ionally na -
owe M³, and longe en al u suppo s i s iden i ica ion as H. diaphanop e us. When compa ed wi h
o he species occu ing in B azil bu ou side he s udy a ea, H. laepho is also has anslucen wings
bu di e s in pelage pa e n (sub ly bicolo ed do sum) and in au icula (o al in shape) and c anial
ea u es. His io us mon anus and H. alienus gene ally ha e da ke pelage and wings, whe eas he ana-
lyzed specimen is dis inguished by i s bicolo ed do sum and anslucen memb anes. Mo eo e , hese
species end o ha e a mo e o al ea mo phology compa ed o H. diaphanop e us. The combina ion o
anspa en wings, bicolo ed u , la ge ea s connec ed by a high band, and educed M³ wid h dis in-
guishes he specimen om all o he His io us.
Figu e 3. His io us diaphanop e us (MN84764).
A. La e al iew showing conspicuously la ge
ea s. B. Gene al golden-b own u wi h dis-
inc ly bicolo ed do sal hai s (da k b own a
he base and golden a he ips) and ans-
lucen u opa agium. C. T anslucen ea s and
wings.
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1148
Li e a u e eco ds. The dis ibu ion o H. diaphanop e us includes B azil and Boli ia. In B azil, he e
a e eco ds om he no heas (Bahia, Cea á, Ma anhão, Pa aíba, and Pe nambuco) and he wes
(Ma o G osso). The ype locali y is in B azil, in he s a e o Bahia, in Sen o Sé municipali y, a Boquei ão
da Onça. In Boli ia, he depa men o San a C uz (Table 1). Mo eo e , he species occu s in he Caa -
inga, Ce ado, and Chaco eco egions.
Ecological no e on oos use and in e ac ions. Roos sha ing by H. diaphanop e us and Molossus sp.,
and a po en ial in e ac ion be ween hese ba species and an insec o he amily Redu iidae, since an
indi idual was obse ed in he same oos . Bo h ba species we e obse ed in his e ical ca i y wi hin
a qua z–a eni e ocky o ma ion (Figu e 4), app oxima ely 2 m abo e he g ound, he oos measu ed
app oxima ely 6 cm in wid h, 25 cm in leng h, and 20 cm in dep h.
Discussion
His io us diaphanop e us has been eco ded om only 10 locali ies o da e, occu ing ac oss he Caa in-
ga, Ce ado, and Chaco eco egions (Feijó e al. 2015; Semedo and Feijó 2016; Be na d e al. 2023). The
eco d epo ed he e ex ends he known geog aphic ange o he species by app oxima ely 170 km
om he nea es locali y in he S a e o Ma anhão, wi hin he Ce ado biome, now o aling h ee known
locali ies in his biome (Figu e 1, locali y 4), and abou 630 km om he ype locali y in he Caa inga
biome (Figu e 1, locali y 2). Addi ionally, we inc ease he numbe o ba species eco ded o he s a e o
Tocan ins o 87 (Souza e al. 2021). When compa ed wi h he 2024 mammal checklis published by he
Sociedade B asilei a de Mas ozoologia (Ab eu e al. 2024), Tocan ins anks six h in ba species ichness,
ied wi h Amapá. Howe e , some o he alues in ha checklis may be ou da ed, since he species ich-
ness epo ed o Tocan ins is conside ably lowe han ha documen ed by Souza e al. (2021).
The specimen epo ed in his s udy, as well as hose p e iously documen ed in he li e a u e, was
ound in a egion cha ac e ized by al e na ing we and d y seasons, a clima ic pa e n ypical o he
Ce ado, Caa inga, and Chaco (Peel e al. 2007). These h ee eco egions, whe e he species occu s,
sha e ecological and clima ic simila i ies and a e o en desc ibed as pa o he Sou h Ame ican diag-
onal o open o ma ions (We neck 2011). All a e de ined by p onounced clima ic seasonali y, wi h p o-
longed d y pe iods ha shape ege a ion adap ed o wa e sca ci y, he e ogeneous habi a mosaics,
Table 2. Ex e nal and c anial measu emen s (in mm) and body mass (g) o His io us diaphanop e us specimens examined in his s udy and om he li e a u e. Specimens a e e-
p esen ed by ouche MN84764 om Liza da, Tocan ins; UPFB 9490–9500 om Sen o Sé, Bahia; UFPB 6014 and UFPB 6015 om São José dos Co dei os, Pa aíba; UFMT 2751 om
Chapada dos Guima ães, Ma o G osso, B azil; MNKM 3714 and AMNH 264086 om Valle G ande, San a C uz, Boli ia. Sex: emale (♀) and male (♂). Museu Nacional, Uni e sidade
Fede al do Rio de Janei o (MN), B azil; Uni e sidade Fede al da Pa aíba (UFPB), B azil; Uni e sidade Fede al de Ma o G osso (UFMT), B azil; Museo Noel Kemp Me cado (MNKM),
Boli ia; Ame ican Museum o Na u al His o y (AMNH), USA. Abb e ia ions o measu emen s a e explained in Me hods.
MN
84764
UPFB
9490*
UPFB
9491
UPFB
9492
UPFB
9493
UPFB
9494
UPFB
9495
UPFB
9496
UPFB
9497
UPFB
9498
UPFB
9499
UPFB
9500
UFPB
6015
UFPB
6014
MNKM
3714
UFMT
2751
AMNH
264086
Sex ♀♀♂♀♀♂♀♂♀♀♂♀♀♀♀♀♀
W 8.00 10.00 8.50 9.50 9.00 8.00 − 8.00 11.00 10.00 8.00 8.50 10.50 9.00 9.00 − 10.90
TL 110.86 104.60 100.60 111.80 111.00 108.30 105.00 104.00 114.80 103.60 105.50 106.60 113.00 104.00 − − 121.00
HBL 59.91 53.20 52.60 56.50 60.00 54.80 58.80 52.00 60.20 52.40 53.20 56.20 − 52.90 60.00 − −
T50.95 51.40 48.00 55.3 51.00 53.50 46.20 52.00 54.60 51.20 52.20 50.40 43.00 52.00 57.00 − 57
HF 9.64 10.10 10.00 7.90 9.30 8.00 10.90 8.10 8.40 10.50 9.30 10.10 8.00 7.60 8.00 − 8.00
E 28.01 31.40 33.10 32.10 30.40 30.30 31.40 28.80 31.70 29.90 30.90 30.60 29.30 28.20 27.00 − 31.00
HSE 4.62 4.00 − − − 3.90 4.50 − 4.40 3.40 3.80 − − − − − −
FA 46.74 44.90 44.60 45.40 46.80 43.90 44.70 41.80 47.20 45.30 44.40 45.70 46.60 45.30 46.00 45.40
T 13.62 13.50 12.90 13.50 13.80 13.30 13.00 13.20 13.20 13.00 14.10 13.50 14.70 13.10 − − 17.00
CaL 17.86 17.70 18.90 17.20 17.20 23.30 17.50 17.90 17.70 20.30 17.80 20.10 − − − − −
EB 19.92 21.30 21.30 − 21.70 22.80 20.80 18.50 20.50 20.10 21.40 20.40 − − − − −
GLS 17.53 18.63 18.08 18.12 18.31 17.80 17.90 17.67 18.30 − 17.50 17.50 18.00 − − 17.44 −
CIL 17.22 17.13 16.79 17.01 17.11 16.44 16.95 16.34 17.10 16.90 16.30 16.30 16.60 − − 16.91 17.17
PC 3.73 4.05 4.05 4.06 3.87 4.00 3.73 3.86 4.00 3.90 4.10 3.85 4.25 − − 3.89 3.79
ZB − 10.65 10.00 10.29 10.34 10.04 9.90 10.06 10.26 9.90 10.10 10.00 10.41 − − 10.48 10.43
BB 7.50 7.85 8.05 7.73 7.64 7.64 7.48 7.68 7.84 7.70 7.60 7.46 8.00 − − 7.96 7.67
MB 8.20 6.96 7.04 6.95 6.86 6.60 6.90 6.62 7.10 6.90 6.40 6.55 7.22 − − 8.11 6.35
HPP 2.19 2.05 1.76 02.01 2.00 1.96 1.90 1.76 1.82 2.02 2.40 1.90 2.38 − − 1.89 1.76
PL 7.61 7.64 7.61 7.65 7.70 7.22 7.55 7.26 7.56 7.60 7.10 7.40 8.33 − − 7.37 7.68
C-M³ 5.95 6.21 5.84 6.10 6.27 5.90 6.05 5.82 6.08 5.80 5.80 5.75 5.93 − − 6.19 6.16
PB 3.77 3.61 3.33 3.26 3.31 3.36 2.96 3.28 3.16 3.10 3.00 3.42 3.09 − − 3.70 3.41
M³-M³ 6.38 6.89 6.23 6.4 6.18 6.30 6.22 6.32 6.62 6.40 6.10 6.18 6.46 − − 6.77 6.52
C-C 4.98 5.20 4.81 4.92 − 4.60 4.67 4.73 5.14 4.70 4.80 4.68 4.78 − − 5.16 4.91
ML 12.12 12.70 12.30 12.35 12.55 11.85 12.54 11.72 12.36 12.70 12.00 12.05 12.57 − − 12.28 12.42
C-M₃ 6.29 6.63 5.95 6.18 6.18 6.16 6.35 6.31 6.57 6.20 6.10 6.23 6.39 − − 6.53 6.64
MH 3.97 4.46 3.85 4.20 4.20 4.16 4.36 4.08 4.32 4.10 4.00 4.04 4.20 − − 5.45 4.40
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1149
and high di e si y o species specialized in a id en i onmen s (We neck 2011; Fe nandes e al. 2022).
Wi hin his sys em, he Ce ado biome plays a cen al ole as a majo sou ce o lo is ic exchanges
wi h neighbo ing eco egions, no ably he Caa inga, which ep esen s he mos species- ich nucleus o
he Seasonally D y T opical Fo es s and has ecei ed subs an ial con ibu ions om Ce ado lineages
(Fe nandes e al. 2022). Al hough connec i i y wi h he Chaco has his o ically been mo e limi ed, all
h ee eco egions sha e di e si ica ion p ocesses linked o global a idi ica ion since he Miocene, which
os e ed he e olu ion o d ough -adap ed aunas and lo as (Ca migno o e al. 2012; Fe nandes e al.
2022). F om a aunal pe spec i e, he Ce ado biome exhibi s he highes mammalian ichness and a
high p opo ion o endemic species, ollowed by he Caa inga and hen he Chaco, while bo h Caa inga
and Chaco display ela i ely low le els o endemism, suppo ing hei inclusion wi hin a b oade com-
plex o in e connec ed open landscapes (Klink and Machado 2005; de Albuque que e al. 2012; Ab eu
e al. 2024). Based on cu en eco ds, H. diaphanop e us is he only species o he genus es ic ed o
opical egions wi h d y and sh ubby ege a ion (Feijó e al. 2015; Semedo and Feijó 2016; Be na d
e al. 2023; Poma-U ey e al. 2023). The o he species ha occu in B azil, H. alienus, H. laepho is, H.
mon anus, and H. ela us, a e associa ed wi h he A lan ic Fo es , whe eas H. ela us is also eco ded in
he Caa inga, Ce ado, and Pampa biomes (ICMBio 2025).
Ul ima ely, mo phological a ia ion in H. diaphanop e us appea s o be sub le, and he species
can be eadily dis inguished om i s congene s based on he ai s desc ibed by Feijó e al. (2015),
mos no ably i s anspa en wings. Mos eco ded measu emen s a e consis en wi h he expec ed
ange o he species, wi h only mino a ia ions. The heigh o skin be ween he ea s and he mas oid
b ead h we e sligh ly highe han he a e age obse ed in o he indi iduals o he same species. In
addi ion, ou obse a ion o H. diaphanop e us and Molossus sp. sha ing a oos wi h an insec o he
amily Redu iidae sugges s a po en ial ecological in e ac ion be ween ba s and edu iids. Al hough
ou obse a ion is occasional, simila associa ions ha e been epo ed in he li e a u e. Some edu i-
Figu e 4. A ea whe e ouche specimen
MN84764 was collec ed. A. Rupes ian ce -
ado wi h qua zi e–sands one ou c ops in
he backg ound. B. Colony o His io us diapha-
nop e us in a qua zi e–sands one ou c op a
Liza da, Tocan ins, B azil. The a ows black in-
dica es (A) he oos loca ion and (B) he insec
belonging o he amily Redu iidae.
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1150
ids ound in ba oos s a e species ha may ac as p eda o s associa ed wi h guano deposi s, con ib-
u ing o he egula ion o in e eb a e popula ions and highligh ing he complexi y o guano-based
ood webs (Fe ei a and Ma ins 1999). Mo e ecen s udies based on ossil eco ds p ese ed in
Dominican ambe ha e e ealed one o he oldes known associa ions be ween a ia omine bug and
a ypanosome, as well as he p esence o ba hai s wi hin he same specimens, indica ing ha ba s
may ha e se ed as e eb a e hos s o T ypanosoma pa asi es millions o yea s ago (Poina 2005).
Toge he , hese indings unde sco e he ecological complexi y o ba oos communi ies and sugges
ha in e ac ions be ween ba s and edu iids may ange om ancien pa asi ic ela ionships o con-
empo a y ophic associa ions.
Howe e , as a ecen ly desc ibed species wi h only sca e ed occu ence eco ds, consis en in o -
ma ion emains lacking ega ding key aspec s o i s biology, including die and popula ion s uc u e.
Mo eo e , simila o he eco egions i inhabi s, he Caa inga, Ce ado, and Chaco, H. diaphanop e us
aces h ea s om human ac i i ies, such as ag icul u al expansion, de o es a ion, and i e, which lead
o habi a agmen a ion and biodi e si y loss (We neck 2011; de Albuque que e al. 2012). His o ically,
hese egions ha e ecei ed less scien i ic and conse a ion a en ion compa ed o humid biomes
such as he Amazon and he A lan ic Fo es , u he inc easing he ulne abili y o hei na i e species
(We neck 2011; de Albuque que e al. 2012). Clima e change exace ba es hese h ea s h ough ising
empe a u es, educed ain all, and he expansion and in ensi ica ion o he d y season (Ho mann
e al. 2021, 2023), exe ing addi ional p essu e on popula ions. In addi ion, he species is lis ed as
Leas Conce n in he B azilian Lis o Th ea ened Species (ICMBio 2025). Howe e , he e a e ew known
eco ds, and i has no ye been assessed by he In e na ional Union o Conse a ion o Na u e (IUCN
2025). Taken oge he , hese ac o s highligh he u gen need o comp ehensi e s udies on H. diapha-
nop e us and o in eg a ed conse a ion s a egies ha conside bo h he sha ed e olu iona y his o y
o hese open biomes and he c i ical ole o hei species in main aining con inen al biodi e si y.
Acknowledgemen s
We hank D . Luiz Flama ion B. de Oli ei a o his con ibu ions o he de elopmen o he manusc ip
and Rod igo C. Du a o his assis ance in edi ing he skull igu e. We hank he e iewe s and edi o s
o hei con ibu ions, which imp o ed his wo k.
Addi ional in o ma ion
Con lic o in e es
The au ho s decla e ha no compe ing in e es s exis .
E hical s a emen
All p ocedu es we e conduc ed unde au ho iza ion om he ‘Sis ema de Au o ização e In o mação
em Biodi e sidade’ (SISBIO/ICMBio), pe mi No. 60058, and app o ed by he Animal E hics Commi -
ee (CEUA/UFRJ), p o ocol No. 016/21.
Funding
Field sampling was unded by ‘Fundação de Ampa o à Pesquisa do Es ado de São Paulo’ (FAPESP;
g an 2017/22269-2). Ma cione B. de Oli ei a ecei ed suppo om he ‘Fundação de Ampa o à Pes-
quisa do Es ado do Rio de Janei o’ (FAPERJ; g an E-26/200.022/2024).
Au ho con ibu ions
Concep ualiza ion: MBO, MAG. Da a cu a ion: MBO, MAG. In es iga ion: MBO, MAG, GSH, IPC, JLPC.
Me hodology: MBO, MAG. Supe ision: MBO. Visualiza ion: MBO, MAG. W i ing – o iginal d a : MBO,
MAG. W i ing – e iew and edi ing: MBO, MAG, GSH, IPC, JLPC.
Au ho ORCID iDs
Ma iane de Almeida Gen il h ps://o cid.o g/0009-0007-2850-0014
Gab iel Selbach Ho mann h ps://o cid.o g/0000-0003-2525-8537
Igo P ei e Coelho h ps://o cid.o g/0000-0003-3645-9094
José Luís Passos Co dei o h ps://o cid.o g/0000-0001-5821-8764
Ma cione B i o de Oli ei a h ps://o cid.o g/0000-0003-1628-3458
Da a a ailabili y
All da a ha suppo he indings o his s udy a e a ailable in he main ex .
Check Lis 21 (6) · h ps://doi.o g/10.15560/21.6.1143
Gen il e al. · His io us diaphanop e us in Tocan ins 1151
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