A taxonomic revision of the South American species of Pseudocanthon Bates, 1887 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini)
Author
Nazaré-Silva, Everton E.
Universidade Federal do Pará, Instituto de Ciências Biológicas. Programa de Pós-graduação em Zoologia da Universidade Federal do Pará e Museu Paraense Emílio Goeldi. Campus Belém. Rua Augusto Corrêa, 01. Guamá. Belém PA 66075 - 110, Brazil.
Author
Silva, Fernando A. B.
Universidade Federal Rural de Pernambuco, Unidade Acadêmica de Serra Talhada-UAST. Av. Gregório Ferraz Nogueira. Mailbox 063. Serra Talhada, PE 56909 - 535, Brazil.
text
Zootaxa
2021
2021-08-30
5027
1
61
86
journal article
10.11646/zootaxa.5027.1.3
1175-5326
5448088
D8FE51C2-C34F-47CE-87FC-80BAB16C54BD
Pseudocanthon xanthurus
(
Blanchard, 1847
)
(
Figs. 1E
,
2B, 2F
,
3B–C
,
4B
,
5B–C
,
6B–C
,
7A
,
8
,
9
)
urn:lsid:zoobank.org:act:
AD91B620-1E60-4091-886B-34D9D56EE278
Canthon xanthurum
Blanchard, 1847: 166
(original description);
Blackwelder 1944: 202
(checklist and distribution).
Canthon xanthurus
:
Harold 1868: 140
(checklist);
Harold 1869: 995
(catalogue and distribution);
Bruch 1911: 185
(catalogue and distribution);
Gillet 1911: 34
(catalogue and distribution);
Arrow 1913: 456
(taxonomic remarks);
Schmidt 1922: 64
, 82 (description and distribution);
Balthasar 1939: 189–190
(description and distribution).
Canthon
(
Pseudocanthon
)
xanthurus
:
Krajcik 2006: 32
(checklist);
Krajcik 2012: 64
(checklist).
Pseudocanthon xanthurum
:
Martínez 1947: 268
(catalogue, distribution, and taxonomic remarks);
Halffter 1961: 232–233
(taxonomic remarks);
Vaz-de-Mello 2000: 194
(checklist);
Ratcliffe
et al
. 2015: 196
(checklist).
Pseudocanthon xanthurus
:
Pereira & Martínez 1956: 109
(taxonomic remarks);
Martínez 1959: 57
(catalogue and distribution);
Pereira & Martínez 1960: 37–38
(taxonomic remarks);
Vulcano & Pereira 1964: 592
(catalogue);
Vulcano & Pereira 1967: 551
(identification key);
Halffter & Martínez 1977: 60
(checklist);
Medina
et al
. 2001: 137
(checklist); Hamel-Leige
et al
. 2006: 14 (checklist);
Medina & Pulido 2009: 59
(checklist);
Carvajal
et al
. 2011: 119
(distribution);
Schoolmeesters 2020
(catalogue);
Vaz-de-Mello 2020
(checklist).
Canthon felix
Arrow, 1913: 456
(original description and taxonomic remarks);
Boucomont 1928: 1
(checklist);
Balthasar 1939: 224
(description and distribution);
Blackwelder 1944: 199
(checklist and distribution);
Pereira & Martínez 1956: 109
(taxonomic remarks);
Pereira & Martínez 1960: 37–38
(synonymy of
Canthon felix
with
Pseudocanthon xanthurus
);
Halffter 1961: 232–233
(cited as junior synonym of
Pseudocanthon xanthurus
);
Bacchus 1978: 102
(
lectotype
designation of
C
.
felix
);
Krajcik 2006: 32
(cited as junior synonym of
Pseudocanthon xanthurus
);
Ratcliffe
et al
. 2015: 196
(checklist).
Diagnosis
.
Pseudocanthon xanthurus
is similar to
P
.
perplexus
in having inconspicuous microsculpture at central portion of pronotum (
Fig. 2B
). However,
P
.
xanthurus
can be distinguished from
P
.
perplexus
by the following characters: surface of clypeus, lateral edges of pronotum, humeral angle, lateral and apex of elytra with orange spots (
Fig. 3B–C
); internal edge of protibia substraight (
Fig. 2F
); apex of parameres rounded, curved outward (
Fig. 5B–C
); frontolateral peripheral (FLP) endophallite elongate, elliptical shaped (
Fig. 6B
); and accessory endophallite (AE) absent (
Fig. 6B
). In contrast, in
P
.
perplexus
dorsal surface of body almost completely black or metallic blue, orange portion restricted to clypeal surface (
Fig. 3A
); internal edge of protibia strongly arched (
Fig. 2E
); apex of parameres curved inward, with small cylindrical projection at apex of ventral edge (
Fig. 5A
); FLP elongate, bifurcate at inferior portion (
Fig. 6A
); and AE present (
Fig. 6A
).
Type material
.
Canthon xanthurus
Blanchard
lectotype
♂
(
here designated
;
MNHN
): “
5664
” // “TYPE” // “MUSEUM PARIS / PROVINCE DE /
CORRIENTES
/ D’ORBIGNY 1834” // “TYPE” // “
Canthon
/
xanthurus
/
Blanch
” [unknown handwriting; Blanchard’s?] // “
LECTOTYPE
♂
/
Canthon
/
xanthurus
/ Blanchard / des.
E.E. Nazaré-Silva
&
F.A.B. Silva
, 2021”
.
Paralectotypes
:
Unknown.
Canthon felix
Arrow
Lectotype
♂
(designated by
Bacchus 1978: 102
;
BMNH
): “
Santar
/
em
” // “LECTO / TYPE” // “
Limbatus
/
Lacordre
” // “
Canthon
/
felix, Arrow
/
type
” [
Arrow’s
handwriting] // “
Canthon
♂
/
felix Arrow
/
M.E. Bacchus
det 197
5
/
LECTOTYPE
”
.
Paralectotypes
♀
(
MNHN
): “
A. Fry
” // “
Pernamo
.
” // “
Pseudocanthon
/
felix
/
(Arrow)
/
GHyAM det 76
” [Halffter’s handwriting] // “Fry Coll. / 1905-100.” // “TYPE /
co
” // “
Canthon
/
felix, Arrow
/
co-type
” [Arrow’s handwriting] // “
PARALECTOTYPE
♀
/
Canthon
/
felix
/ Arrow / des. E.E. Nazaré- Silva &
F.A.B. Silva
, 2021”
.
Non-type material examined
.
277♀
and
163♂
.
Intraspecific
patterns listed by us in morphological variation section are indicated as follows: pattern 1 or 2 for body colour; pattern A or B for male genitalia.
NO DATA
: 2 unsexed specimens (
CERPE
; pattern 1).
FRENCH GUIANA
:
CAYENNE
:
Kourou
, [
05°09’40.07’’S
;
52°38’57.6’’W
]
—
1♀
(
CMNC
; pattern 1);
Montagne des Chevaux
, [
04°44’56’’N
;
52°26’28’’W
],
75 m
,
04.II.2013
,
Seag
—
1♀
1♂
(
CEMT
; pattern 1);
Paracou
field station, [
05°02’N
;
52°00’W
],
55 m
,
X.2003
, F. Feer—
3♀
5♂
(
CEMT
; pattern 1,
one ♂
pattern A).
SURINAME
:
PARAMARIBO
:
Zanderij
, [
05°53’N
;
55°05’W
],
17–18.IV.2010
, A. Hielkema—
1♀
(
CEMT
; pattern 1).
VENEZUELA
:
BOLÍVAR
:
10 km
northwest of
Corocito
[rainforest in Caura River], [
07°32’12.27’’N
;
65°00’23.23’’W
],
26 m
,
18.VI–03.VIII.1987
, J. Peck & S. Peck—
2♀
(
CMNC
; pattern 1);
15 km
northwest of
Corocito
, [
07°32’12.27’’N
;
65°00’23.23’’W
],
26 m
,
18.VI.1987
, J. Peck & S. Peck—
2♀
2♂
(
CMNC
; pattern 2,
one ♂
pattern B);
25 km
southwest of
Puerto Ordaz
, [
08°06’21.74’’N
;
62°35’49.62’’W
],
260 m
,
21.VI.1987
, J. Peck & S. Peck—
1♀
(
CMNC
; pattern 2);
20 km
east of
El Palmar
, [
08°00’33.16’’N
;
62°05’29.64’’W
],
380 m
,
18.VI.1996
, H. Howden & A. Howden—
1♂
(
CMNC
; pattern 1);
Caicara del Orinoco
,
15 km
east of
Caicara del Orinoco
, [
07°35’52.63’’N
;
66°00’38.47’’W
],
43 m
,
12.VI.1996
, H. Howden & A. Howden—
2♀
1♂
(
CMNC
; pattern 2,
one ♂
pattern A);
Caroní
,
35 km
southwest of
Puerto Ordaz
, [
08°06’34.97’’N
;
62°29’04.76’’W
],
388 m
,
13.VII–02.VIII.1987
, J. Peck & S. Peck—
8♀
7♂
(
CMNC
; both body colour patterns are represented, and
two ♂
pattern B);
Heres
,
20 km
southwest of
Bolívar
, [
07°59’27.13’’N
;
63°42’14.13’’W
],
79 m
,
19.VI.1987
, J. Peck & S. Peck—
2♀
(
CMNC
; both body colour patterns are represented);
Parupá
, [
05°41’28.21’’N
;
61°34’20.64’’W
],
1500 m
,
27–30.VI.1987
, J. Peck & S. Peck—
1♀
(
CMNC
; pattern 2);
Yaracuy
,
Aroa
, [
10°00’00’’N
;
68°00’00’’W
],
459 m
,
19.VII.2009
, pitfall with human feces, Asmussen—
1♀
(
CEMT
; pattern 1).
COLOMBIA
:
CASANARE
: [
05°45’32.14’’N
;
72°10’20.62’’W
],
184 m
,
I.2004
, J. Noriega—
1♀
2♂
(
CEMT
; pattern 2,
one ♂
pattern B);
VICHADA
:
Cumaribo
,
Santa Rita
,
Parque Nacional Natural El Tuparro
, [
05°21’03’’N
;
67°52’15’’W
],
135 m
,
03–05.II.2004
, I. Quintero & E. González—
1♀
(
CEMT
; pattern 2);
VAUPÉS
:
Mitú
,
Querari Pelotão
, [
01°05’N
;
69°51’W
],
07–23.IV.1993
, C. Motta—
2♂
(
INPA
; pattern 1);
AMAZONAS
:
Leticia
, [
04°16’54.39’’N
;
69°56’09.32’’W
],
76 m
,
23.II–02.III.1974
, Howden & Nealis—
2♀
(
CMNC
; pattern 1).
BRAZIL
:
No
locality,
20.V.1967
,
no collector—1 unsexed specimen (
DZUP
, pattern 1);
AMAPÁ
:
Porto Baton
, [no coordinates],
139 m
,
05.VIII.1983
, J. Lacerda—
1♂
(
DZUP
; pattern 1, A);
RORAIMA
:
Boa Vista
,
Serra da Moça
, [
02°16’13.82’’N
;
60°40’28.51’’W
],
105 m
,
VII.1997
, F. Vaz-de-Mello—
2♂
(
CEMT
; pattern 1, A);
Cantá
,
Lavrado
, [
02°47’N
;
60°39’W
],
70 m
,
IX.1996
, Ribeiro & Vaz-de-Mello—
1♀
1♂
(
CEMT
; pattern 1, A);
RIO GRANDE DO NORTE
:
Natal
, [
05°46’45.32’’S
;
35°12’03.3’’W
],
31 m
,
V.1951
—
1♀
(
DZUP
; pattern 2);
Parnamarim
, [
05°54’41.6’’S
;
35°16’16.74’’W
],
56 m
,
IV.1950
, M.Alvarenga—
2♂
(
DZUP
; pattern 2, B);
CEARÁ
:
Maranguape
,
Santuário Nossa Senhora de Penha
[shrubs], [
03°52’29’’S
;
38°40’48’’W
],
96 m
,
09.IV.2008
,
Gillet
—
1♀
1♂
(
CEMT
;
♂
with pattern 1, B and
♀
with pattern 2);
Ubajara
,
Parque Nacional de Ubajara
, [
03°50’19.68’’S
;
40°53’53.25’’W
],
797 m
,
14.II.2013
,P.Grossi&F.Vaz-de-Mello—
1♂
(
CERPE
;pattern2);
Quixeramobim
,[
05°29’54.23’’S
;
39°19’14.25’’W
],
212 m
, 1926,
M. Rocha
—
1♀
(
MZSP
; pattern 2);
PIAUÍ
:
Canto de Buruti
, [
08°07’56’’S
;
43°00’23’’W
],
366 m
,
05.IV.2008
,
Gillet
—
1♀
(
CEMT
; pattern 2);
Piracuruca
,
Parque Nacional de Sete Cidades
, [
04°06’02.71’’S
;
41°42’44.63’’W
],
184 m
,
12.II.2013
, P. Grossi & F. Vaz-de-Mello—
1♀
(
CERPE
; pattern 2);
Piracuruca
,
Parque Nacional de Sete Cidades
, [
04°06’02.71’’S
;
41°42’44.63’’W
],
184 m
,
07.III.2013
, P. Grossi & F. Vaz-de-Mello—
1♀
(
CERPE
; pattern 2);
Teresina
, [
05°02’40.87’’S
;
42°45’58.44’’W
],
85 m
,
VIII.1952
,
Oliveira
—
2♀
2♂
(
DZUP
; pattern 2,
two ♂
pattern B);
Teresina
, [
05°02’40.87’’S
;
42°45’58.44’’W
],
85 m
,
I.1953
,
Oliveira
—
3♀
(
CMNC
;
two ♀
pattern 2 and
one ♀
pattern 1);
Teresina
, [
05°02’40.87’’S
;
42°45’58.44’’W
],
85 m
,
I.1953
,
Oliveira
4♀
2♂
(
MZSP
;
two ♂
pattern 2, A,
two ♀
pattern 2, and
two ♀
pattern 1);
MARANHÃO
:
Chapadinha
, [
04°51’50’’S
;
43°20’51’’W
],
17.III.2012
, C. Silva—
1♂
(
CEMT
; pattern 1, B);
Chapadinha
,
Fazenda Unha de Gato
, [
03°41’42.3’’S
;
43°11’48.1’’W
], 2016,
pitfall
,
R. Silva
&
G. Machado
—
1♂
(
CEMT
; pattern 1, A);
Itapecuru Mirim
[pasture], [
03°23’S
;
44°32’W
],
31.VIII.2010
, R. Matavelli—
3♀
2♂
(
MZUFPA
; pattern 2,
two ♂
pattern B);
Loreto
,
Pedrinhas
, [
06°57’18.95’’S
;
45°09’27.58’’W
],
Mangrove
,
05.X.1987
, light trap, E. Bergmann—
1♀
(
CEMT
; pattern 2);
Ilha de São Luís
, [
02°35’11’’S
;
44°13’32’’W
],
54 m
,
04.VI.2001
, L. Costa Júnior—
1♂
(
CEMT
; pattern 2);
Ilha de São Luís
, [
02°35’11’’S
;
44°13’32’’W
],
54 m
,
08.VI.2001
, L. Costa Júnior—
1♀
(
CEMT
; pattern 1);
Ilha de São Luís
, [
02°35’11’’S
;
44°13’32’’W
],
54 m
,
12.VI.2001
, L. Costa Júnior—
1♀
(
MZUFPA
; pattern 2);
Santa Quitéria do Maranhão
,
Fazenda Rodiador
, [
03°24’50’’S
;
42°40’46’’W
],
47 m
,
18.V.2008
,
Gillet
—
1♀
(
CEMT
; pattern 2);
Santa Quitéria do Maranhão
,
Fazenda Rodiador
, [
03°24’50’’S
;
42°40’46’’W
],
47 m
,
23.II.2008
,
Gillet
—
1♀
(
CEMT
; pattern 1);
PARÁ
:
Belém
,
Universidade Federal Rural da Amazônia
, [
01°27’22’’S
;
48°26’15’’W
],
04–13.V.2019
, M. Silva—
7♀
5♂
(
MZUFPA
; pattern 1,
one ♂
pattern A);
Jacareacanga
, [
06°13’25.88’’S
;
57°45’38.73’’W
],
190 m
,
XII.1968
, M. Alvarenga—
2♀
(
DZUP
; pattern 1);
Jacareacanga
, [
06°13’25.88’’S
;
57°45’38.73’’W
],
190 m
,
XII.1969
, F. Barbosa—
1♀
(
DZUP
; pattern 1);
Monte Dourado
,
Jari
[eucalyptus area], [
00°52’04.14’’S
;
52°32’05.98’’W
],
28.IV.2009
,
Schiffler
—
3♀
(
MZUFPA
; pattern 1);
Monte Dourado
,
Jari
[eucalyptus area], [
00°52’04.14’’S
;
52°32’05.98’’W
],
01.V.2009
,
Schiffler
—
3♀
(
MZUFPA
; pattern 1);
Monte Dourado
,
Jari
[eucalyptus area], [
00°52’04.14’’S
;
52°32’05.98’’W
],
02.V.2009
,
Schiffler
—
1♀
(
MZUFPA
; pattern 1);
Novo Progresso
,
Fazenda Florentino
[Amazon rainforest], [
07°06’54’’S
;
55°24’23’’W
],
207 m
,
I.2011
, pitfall with human feces, Pelissari—
5♀
3♂
(
CEMT
; pattern 1,
two ♂
pattern A);
Paragominas
, [
2.9372°S
;
47.2102°W
],
II.2011
, R. Solar—
64♀
34♂
(
CEMT
; pattern 1,
five ♂
pattern A);
Paragominas
, Canindé
Rio Gurupi
, [
02°34’00’’S
;
46°31’00’’W
],
65 m
,
27.II.1966
,
Malkin
—
1♀
(
MZSP
; pattern 1);
Santarém
, [
02°30’15.35’’S
;
54°57’16.71’’W
],
23 m
,
VII–VIII.2003
, R. Matavelli—
3♂
(
CEMT
;
two ♂
pattern 2, B and
one ♂
pattern 1);
AMAZONAS
:
Benjamin Constant
, [
04°31’13’’S
;
69°57’18’’W
],
III.2008
, V. Korasaki—
1♀
(
MZUFPA
; pattern 2);
Benjamin Constant
,
Comunidade Nova Aliança
, [
04°21’12’’S
;
69°36’16’’W
],
20.III.2008
, pitfall with human feces, V. Korasaki—
3♀
2♂
(
CEMT
; pattern 1,
one ♂
pattern A);
Benjamin Constant
,
Comunidade Nova Aliança
, [
04°22’33.57’’S
;
70°01’48.59’’W
],
71 m
,
II.2004
, P. Silva—
2♂
(
CEMT
; pattern 1);
Manaus
,
Fazenda Dimona
, [
02°25’S
;
59°50’W
],
23–26.VI.2000
, I. Quintero—
23♀
16♂
(
INPA
; pattern 1);
Manaus
,
Fazenda Dimona
, [
02°25’S
;
59°50’W
],
05– 08.VII.2000
, I. Quintero—
14♀
6♂
(
INPA
; pattern 1);
Manaus
,
Fazenda Esteio
, [
02°35’26.58’’S
;
60°01’36.73’’W
],
61 m
, no date or collector
—
1♂
(
INPA
; pattern 1);
Manaus
,
Reserva Florestal Adolpho Ducke
, [
02°55’56’’S
;
59°58’22’’W
],
28.IV–03.V.2013
, F. Vaz-de-Mello—
1♀
2♂
(
CEMT
; pattern 1,
one ♂
pattern A);
Nhamundá
, [
02°13’36.68’’S
;
57°11’15.48’’W
],
47 m
, 1959,
Capelleto
—
1♂
(
DZUP
; pattern 1, B);
Santa Isabel do Rio Negro
,
Tapuruquara
,
Rio Negro
, [
00°22’47.59’’S
;
64°59’46.68’’W
],
45 m
,
01.XII.1962
, J. Bechyne—
1♀
(
MZSP
; pattern 1);
São Gabriel da Cachoeira
,
Uapés
, [
00°07’05.33’’S
;
67°04’19.03’’W
],
69 m
,
28.III.1964
, J. Bechyne & B. Bechyne—
1♂
(
CMNC
; pattern 1);
PARAÍBA
:
Coremas
, [
07°01’03.75’’S
;
37°56’48.42’’W
],
216 m
,
VI.1957
—
1♀
(
MZSP
; pattern 2);
PERNAMBUCO
:
Araçoiaba
,
Campo de Instrução Marechal Newton Cavalcanti
[open habitat], [
07°49’53.06’’S
;
35°06’13.31’’W
],
126 m
,
22.I.2010
, pitfall with human feces, F. Costa—
1♀
1♂
(
CEUPE
; pattern 1,
♂
pattern A);
Caruaru
,
Brejo Novo
, [
08°21’00’’S
;
35°58’59.88’’W
],
599 m
,
18.I.2004
, pitfall with human feces, no collector—
1♂
(
CEUPE
; pattern 2);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°49’42.76’’S
;
34°52’26.82’’W
],
25.V.2007
, pitfall, M. Costa—
1♀
(
CEMT
; pattern 2);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
25.V.2007
, pitfall, Costa—
2♂
(
CEUPE
; pattern 2, B);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
10.VI.2007
, pitfall, Costa—
1♀
(
CEUPE
; pattern 2);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
31.I.2006
, pitfall with human feces, Costa—
1♀
(
CEUPE
; pattern 2);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
30.IX.2006
, pitfall with rotten meat, Costa—
1♀
1♂
(
CEUPE
;
♂
pattern 2, B and
♀
pattern 1);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
30.III.2006
, pitfall with human feces, Silva—
1♀
(
CEUPE
; pattern 1);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
02.VI.2006
, pitfall with rotten meat, Silva—
2♀
(
CEUPE
; pattern 1);
Igarassu
,
Refúgio Ecológico Charles Darwin
[Atlantic rainforest], [
07°48’37’’S
;
34°57’25’’W
],
29.XII.2006
, pitfall with rotten meat, Silva—
1♀
(
CEUPE
; pattern 2);
Recife
, [
08°03’08.07’’S
;
34°55’42.99’’W
],
30 m
,
V.1949
, no collector—
3♀
(
MZSP
; pattern 2);
Recife
, [
08°03’08.07’’S
;
34°55’42.99’’W
],
30m
,
M.Alvarenga
—
2♀
(
DZUP
;pattern2);
ACRE
:
Rio Branco
,[
09°58’31.36’’S
;
67°49’29.63’’W
],
156 m
,
29.X.1954
, M. Alvarenga—
1♂
(
MZSP
; pattern 1, A);
BAHIA
:
Cruz
das
Almas
, [
12°40’24.8’’S
;
39°06’06.7’’W
],
I.2005
, A. Brito & G. Ribeiro—
3♀
2♂
(
CEMT
; pattern 1,
one ♂
pattern A);
Porto Seguro
[pasture], [
16°23’35’’S
;
39°09’45’’W
],
74 m
,
15.V.2012
, L. Audino—
1♀
(
CEMT
; pattern 1);
Porto Seguro
[pasture], [
16°23’59’’S
;
39°07’34’’W
],
71 m
,
16.V.2012
, pitfall with human feces, L. Audino—
2♀
2♂
(
CEMT
; pattern 1,
one ♂
pattern A);
Porto Seguro
[pasture], [
16°23’49’’S
;
39°08’21’’W
],
16.V.2012
, L. Audino—
2♀
5♂
(
CEMT
; pattern 1,
one ♂
pattern A);
Porto Seguro
,
Reserva Particular do Patrimônio Natural Estação Veracel
, [
16°23’15.71’’S
;
39°10’11.04’’W
],
XI.2004
, pitfall with human feces, J. Louzada—
1♂
(
CEMT
; pattern 1);
MATO GROSSO
:
Alta Floresta
, [
09°54’27’’S
;
56°03’18’’W
],
III.2008
, pitfall with human feces, E. Berenguer—
20♀
6♂
(
CEMT
; pattern 1,
four ♂
pattern A);
Alta Floresta
, [
09°54’24’’S
;
56°02’39’’W
],
III.2008
, pitfall with human feces, E. Berenguer—
2♀
4♂
(
CEMT
; pattern 1,
two ♂
pattern A);
Alta Floresta
, [
09°51’06’’S
;
55°59’04’’W
],
IV.2008
, pitfall with human feces, E. Berenguer—
4♀
1♂
(
CEMT
, pattern 1);
Alta Floresta
, [
09°50’22’’S
;
56°00’21’’W
],
IV.2008
, pitfall with human feces, E. Berenguer—
2♀
1♂
(
CEMT
; pattern 1);
Alta Floresta
, [
09°46’55’’S
;
56°02’13’’W
],
IV.2008
, pitfall with human feces, E. Berenguer—
1♀
(
CEMT
; pattern 1);
Alta Floresta
, [
09°51’06’’S
;
55°59’04’’W
],
V.2008
, pitfall with human feces, E. Berenguer—
2♀
1♂
(
CEMT
; pattern 1, male pattern A);
Cotriguaçu
,
Fazenda São Nicolau
, [
09°49’43’’S
;
58°15’56’’W
],
10.XII.2009
, F. Vaz-de-Mello—
1♂
(
CEMT
; pattern 1);
Cotriguaçu
,
Fazenda São Nicolau
[fig plantation], [
09°49’32’’S
;
58°15’46’’W
],
220 m
,
10.XII.2009
, pitfall with human feces, F. Vaz-de-Mello—
1♀
(
CEMT
; pattern 1);
Cotriguaçu
,
Fazenda São Nicolau
, [
09°49’52’’S
;
58°15’17’’W
],
X.2009
, pitfall, F. Vaz-de-Mello—
5♀
(
CEMT
; pattern 1);
Cotriguaçu
,
Fazenda São Nicolau
, [
09°51’38’’S
;
58°12’53’’W
],
X.2009
, pitfall, F. Vaz-de-Mello—
1♂
(
CEMT
; pattern 1);
Cotriguaçu
,
Fazenda São Nicolau
, [
09°49’32’’S
;
58°15’46’’W
],
09.XI.2010
, pitfall with human feces, M. Gigliotti—
1♀
(
CEMT
; pattern 1);
Cuiabá
,
Fazenda Santhidi
, [
15°36’03.1’’S
;
56°05’48.56’’W
],
190 m
,
08.I.2010
, L. Silva—
1♀
2♂
(
CEMT
; pattern 2,
one ♂
pattern B);
Poconé
[Pantanal], [
17°50.73’S
;
57°24.17’W
],
15.XI.2011
, pitfall with human feces, J. Silva—
2♀
(
CEMT
; pattern 1);
Sinop
,
Embrapa
, [
11°51’S
;
55°35’W
],
370 m
,
III–XII.2016
, pitfall, L. Lopes & R. Pita—
1♀
(
CERPE
; pattern 2);
Tangará da Serra
,
Sítio Clemente
, [
14°39’46’’S
;
57°49’35’’W
],
368 m
,
12–14.XII.2012
, pitfall with human feces, R. Silva—
1♂
(
CEMT
; pattern 1);
RONDÔNIA
:
Porto Velho
,
Rio Madeira
, [
08°45’40.3’’S
;
63°54’01.58’’W
],
76 m
, no date and collector
—
1♀
(
INPA
; pattern 1);
MINAS GERAIS
:
Cordisburgo
,
Fazenda Pontinha
, [
19°07’31.09’’S
;
44°19’13.63’’W
],
XII.1993
, F. Vaz-de-Mello—
2♀
(
CEMT
; pattern 1);
Cordisburgo
,
Fazenda Pontinha
, [
19°07’31.09’’S
;
44°19’13.63’’W
],
I.1994
, F. Vaz-de-Mello—
1♀
1♂
(
CEMT
; pattern 1,
♂
pattern B);
Curvelo
,
Fazenda do Roberto Lubernau
[pasture], [
18°47’43’’S
;
44°38’51’’W
],
561 m
,
26.I.2012
, R. Macedo—
3♀
1♂
(
CEMT
; pattern 1,
♂
pattern A);
Lavras
, [
21°14’54.56’’S
;
45°00’04.95’’W
],
926 m
,
I.1999
, J. Louzada—
1♀
(
CEMT
; pattern 1);
Pompéu
,
Fazenda do Pedro
, [
19°09’17’’S
;
44°52’23’’W
],
458 m
,
22.I.2012
, pitfall with human feces, R. Macedo—
1♀
1♂
(
CEMT
; pattern 1);
Visconde do Rio Branco
, [
21°00’49.65’’S
;
42°50’15.4’’W
],
I.1998
, T. Cavalcanti—
2♀
(
CEMT
; pattern 1);
DISTRITO FEDERAL
:
Brasília
, [
15°49’36.09’’S
;
47°55’18.55’’W
],
1100 m
,
XI.1999
, at light, N. Degallier—
1♀
1♂
(
CMNC
; pattern 1,
♂
pattern A);
GOIÁS
:
Goiatuba
, [
18°00’46.03’’S
;
49°21’59.79’’W
],
803 m
,
II.1947
, no collector—
1♀
(
CMNC
; pattern 1); same data but,
XI.1943
, no collector—
1♀
(
MZSP
; pattern 1);
Mineiros
,
Parque Nacional
das
Emas
, [
17°56’29’’S
;
52°58’40’’W
],
15.III.2011
, pitfall with human feces, M. Souza—
1♀
(
CEMT
; pattern 1);
Mineiros
,
Parque Nacional
das
Emas
, [
18°04’06’’S
;
52°56’09’’W
],
15.III.2011
, pitfall with human feces, M. Souza—
1♂
(
CEMT
; pattern 2, B);
Mineiros
,
Parque Nacional
das
Emas
, [
18°00’29’’S
;
52°58’56’’W
],
15.III.2011
, pitfall with human feces, M. Souza—
1♀
(
CEMT
; pattern 1);
RIO DE JANEIRO
:
Guapimirim
,
II.1944
,
illegible collector—1 unsexed specimen (
MNRJ
);
Itaguaí
, [
22°51’47.49’’S
;
43°46’40.47’’W
],
26 m
,
I.1991
, C. Godinho—
1♀
(
CEMT
; pattern 1);
Rio de Janeiro
,
Deodoro
,
20.II.1946
,
W. Zikán
—1 unsexed specimen (
MNRJ
);
Rio de Janeiro
,
Ilha do Governador
,
16.XI.1955
,
M. Alvarenga
—1 unsexed specimen (
MNRJ
);
Rio de Janeiro
,
Jacarepaguá
,
02.III.1952
,
O. Rego
—1 unsexed specimen (
MNRJ
);
Seropédica
,
Estrada Rio-São Paulo
, km 47,
03.III.1948
,
W. Zikán
—1 unsexed specimen (
MNRJ
);
Seropédica
,
Estrada Rio-São Paulo
, km 47,
XII.1948
,
W. Zikán
—2 unsexed specimens (
MNRJ
);
SÃO PAULO
:
Bertioga
,
Itaguaré
, [
23°47’32.7’’S
;
45°49’32.23’’W
]
—
1♀
(
CMNC
; pattern 1);
Bragança Paulista
, [
22°57’11.41’’S
;
46°32’32.37’’W
],
849 m
,
D. Vaglio
—
1♀
(
MZSP
; pattern 1);
Intanhaém
, [
24°10’54.48’’S
;
46°47’06.12’’W
],
7 m
,
II.1966
, U. Martins—
1♀
(
MZSP
; pattern 1);
Itú
,
Fazenda Pau D’Alho
, [
23°15’50.4’’S
;
47°17’58.82’’W
],
601 m
,
05.XI.1960
, U. Martins—
1♂
(
MZSP
; pattern 1, A);
MATO GROSSO DO SUL
:
Selvíria
,
Fazenda Experimental da Universidade Estadual Paulista
[pasture], [
20°20’51.89’’S
;
51°24’51.88’’W
],
03.XI.1994
,light trap, C.Flechtmann—
1♀
(
CEMT
; pattern 1);
PARANÁ
:
Ponta Grossa
, [
25°05’57.93’’S
;
50°09’29.96’’W
],
953 m
,
III.1945
—
1♀
(
DZUP
; pattern 1);
Wenceslau Braz
, [
23°52’17.07’’S
;
49°48’08.85’’W
],
832 m
,
I.1941
, J. Leprevest—
1♀
(
MZSP
; pattern 1);
SANTA CATARINA
:
Anchieta
, [
26°34’S
;
53°22’W
],
08.XII.2014
, pitfall, V. Alves—
1♀
(
CEMT
; pattern 1);
Seara
,
Nova Teutonia
, [
27°11’S
;
52°23’W
],
300–500 m
,
06.I.1946
, F. Plaumann—
1♀
(
CMNC
; pattern 1);
Seara
,
Nova Teutonia
, [
27°11’S
;
52°23’W
],
300–500 m
,
IX.1957
, F. Plaumann—
1♂
(
MZSP
; pattern 1);
Seara
,
Nova Teutonia
, [
27°03’00’’S
;
52°24’00’’W
],
864 m
,
XII.1971
, F. Plaumann—
2♂
(
CMNC
; pattern 1, A);
Seara
,
Nova Teutonia
, [
27°03’00’’S
;
52°24’00’’W
],
864 m
,
I.1975
, F. Plaumann—
1♀
(
CMNC
; pattern 1).
BOLIVIA
:
SANTA CRUZ
:
Buena Vista
, 3,7 km southeast of
Buena Vista
,
Hotel Flora
and
Fauna
[transitional tropical forest], [
17°29.949’S
;
63°33.152’W
],
405 m
,
05–15.XI.2001
, M. Thomas & B. Dozier—
2♀
2♂
(
CMNC
; pattern 1,
♂
pattern A);
COCHABAMBA
:
Chapare
, S. F.
Chipiriri
, [
16°58’07’’S
;
65°23’41’’W
],
400 m
,
XI.1959
, no collector—
1♂
(
CMNC
; pattern 1, A).
PARAGUAY
:
CORDILLERA
:
Piribebuy
, [
25°27’53.37’’S
;
57°02’31.73’’W
],
255 m
,
XI.1949
, no collector—
1♂
(
CMNC
; pattern 1, A);
GUAIRÁ
:
Villarica
, [
25°46’33.82’’S
;
56°26’58.53’’W
],
183 m
,
IV.1942
, no collector—
1♂
(
MZSP
; pattern 1).
ARGENTINA
:
FORMOSA
: Parque Nacional
Río Pilcomayo
, [
25°08’43.11’’S
;
58°11’11.78’’W
],
76 m
,
17.XII.1990
, J. Peck & S. Peck—
1♀
(
CMNC
; pattern 1);
MISIONES
:
Puerto Iguazú
, [
25°50’53.71’’S
;
54°20’48.18’’W
],
294 m
,
X.1927
, no collector—
1♂
(
CMNC
; pattern 1, A)
.
Distribution
. Known from
Venezuela
(
Bolívar
,
Yaracuy
),
Suriname
(
Paramaribo
),
French Guiana
(Cayenne),
Colombia
(Amazonas,
Casanare
,
Vichada
),
Brazil
(
Acre
,
Amapá
, Amazonas,
Bahia
,
Ceará
, Distrito Federal,
Goiás
,
Maranhão
,
Mato Grosso
,
Mato Grosso do Sul
,
Minas Gerais
, Pará,
Paraíba
,
Paraná
,
Pernambuco
,
Piauí
,
Rio de Janeiro
,
Rio Grande do Norte
,
Rondônia
,
Roraima
,
Santa Catarina
,
São Paulo
),
Bolivia
(Chapare,
Santa Cruz
),
Paraguay
(
Cordillera
,
Guairá
) and
Argentina
(
Formosa
, Misiones) (
Fig. 7A
). According to literature data, this species was also cited from
Peru
by
Ratcliffe
et al
. (2015)
. However, since the specific locality was not mentioned we have included a “?” in distribution map (
Fig. 7A
).
Areas of endemism [South America]
:
Brazilian subregion
: South Brazilian dominion: Rondônia province. Boreal Brazilian dominion: Guianan Lowlands, Imerí, Napo, Pará, and Roraima provinces. South Brazilian dominion: Yungas province.
Chacoan subregion
: Chacoan dominion: Caatinga, Cerrado, and Chaco provinces. Parana dominion: Atlantic and Parana provinces (see
Morrone 2014
: fig. 12).
Notes about the
lectotype
of
Canthon xanthurus
Blanchard, 1847
. Analyzing the original description (
Blanchard 1847
), no
holotype
was originally fixed, and we could not find any explicit mention that
C
.
xanthurus
was described based on a single specimen. Following Recommendation 73F of the International Code of Zoological Nomenclature (
International Commission on Zoological Nomenclature 1999
), it is, therefore, assumed that the type series was based on more than one individual. Fernando Vaz-de-Mello found only a single
syntype
of
C
.
xanthurus
in the MNHN during his studies of
Scarabaeinae
type specimens housed in European museums (see
Vaz-de-Mello & Cupello [2018]
for more details). It is possible to recognize this specimen as part of the type series because, as indicated by its labels, it was collected by Alcide d’Orbigny in
Corrientes
,
Argentina
, which is in agreement with information on the collector and type locality from the original description (
Blanchard 1847
). Furthermore, the
lectotype
matches Blanchard’s succinct description.
Notes about the synonymy of
Canthon felix
Arrow, 1913
with
Canthon xanthurus
Blanchard, 1847
.
Canthon felix
was described by
Arrow (1913)
based on specimens from
Pernambuco
,
Ceará
(Baturité and Maranguape), Pará (Santarém), and
Rio de Janeiro
, in
Brazil
.
Bacchus (1978)
found
eight syntypes
(
4 males
and
4 females
) in the BMNH and designated the male from Santarém as the
lectotype
. The
paralectotypes
from Para, Santarém,
Pernambuco
, and Baturité were found; the others (
Rio de Janeiro
and Maranguape) were deemed lost. Decades later, Fernando Vaz-de-Mello found
two paralectotypes
of
C
.
felix
, the
lectotype
male from Santarém, Pará (BMNH), and a
paralectotype
female from
Pernambuco
(MNHN). We examined these
two specimens
from photographs, as well as the
lectotype
of
C. xanthurus
from
Argentina
and non-type material from different localities. The
lectotype
and
paralectotype
of
C
.
felix
have orange spots on almost the entire lateral edge of the body, mainly at elytra, larger density of pronotum and elytra punctures compared to
C
.
xanthurus
. The
lectotype
of
C
.
xanthurus
has orange spots only at base and apex of elytra, smaller density of pronotum and elytra punctures compared to
C. felix
.
Arrow (1913)
also explicitly stated that
C. felix
was described based on those differences above mentioned in comparison to the type of
C. xanthurus
(on dorsal orange spots as well as on the pronotal and elytral punctation). However, these morphological patterns co-occur in specimens from the same locality throughout the distribution range of
P
.
xanthurus
. Also, we verified these different patterns in specimens collected near the type localities of each species name, which are least
1,600 km
apart. Thus, we consider this variation as an intraspecific component, and we maintain here the synonym of
C
.
felix
with
C
.
xanthurus
proposed by
Pereira & Martínez (1960)
.
Morphological variation
.
Pseudocanthon xanthurus
exhibits considerable discrete intraspecific variation in body colour pattern and some structural components of male genitalia. Some specimens have orange spots on almost the entire lateral edge of the body, mainly in elytra (
Fig. 3C
). Other specimens have orange spots only at the base and apex of elytra (
Fig. 3B
). The aedeagus, in ventral view, has the apex of the parameres subparallel (
Fig. 5C
) or laterally projected (
Fig. 5B
). The frontolateral peripheral (FLP) endophallite can also present variation in width and degree of curvature. It can be narrow and straight, or wider and curved (
Fig. 6B–C
).
These patterns of variation are found throughout the range of the species, some populations being polymorphic (
Figs. 8–9
). Regarding the body colour patterns, specimens with pattern 1 have orange spots only at base and apex of elytra (
Fig. 3B
), and have been recorded in
Argentina
(Formosa, Misiones),
Bolivia
(Chapare, Santa Cruz),
Brazil
(Acre, Amapá, Amazonas, Bahia, Distrito Federal, Goiás, Maranhão, Mato Grosso, Mato Grosso do Sul, Minas Gerais, Pará, Paraná, Pernambuco, Piauí, Rio de Janeiro, Rondônia, Roraima, Santa Catarina, São Paulo),
Colombia
(Amazonas),
French Guiana
(Cayenne, Kourou),
Paraguay
(Cordillera, Guairá),
Suriname
(Paramaribo), and
Venezuela
(
Bolívar
,
Yaracuy
) (
Fig. 8A
). Pattern 2, in turn, includes specimens with orange spots on almost the entire lateral edge of body, mainly in elytra (
Fig. 3C
); these have been found in
Brazil
(Amazonas, Ceará, Goiás, Maranhão, Pará, Paraíba, Pernambuco, Piauí, Rio Grande do Norte),
Colombia
(Casanare, Vichada), and
Venezuela
(
Bolívar
) (
Fig. 8A
). The distribution of the two body colour patterns overlaps in several localities in
Brazil
(Amazonas, Goiás, Maranhão, Mato Grosso, Pará, Pernambuco, and Piauí) and
Venezuela
(
Bolívar
), being those populations polymorphic (
Fig. 8A
).
Regarding the parameres and the frontolateral peripheral endophallite, there are two general patterns.
Pattern A
consists of specimens having parameres apically with lateral projection (
Fig. 5B
) and narrow and straight frontolateral peripheral endophallite (
Fig. 6B
) and has been found in
Argentina
(Misiones),
Bolivia
(Chapare, Santa Cruz),
Brazil
(Acre, Amapá, Amazonas, Bahia, Distrito Federal, Maranhão, Mato Grosso, Minas Gerais, Pará, Pernambuco, Piauí, Roraima, Santa Catarina),
French Guiana
(Cayenne),
Paraguay
(Cordillera, Guairá), and
Venezuela
(
Bolívar
) (
Fig. 8B
).
Pattern B
, in turn, includes specimens having parameres apically subparallel (
Fig. 5C
) and with frontolateral peripheral endophallite wider and curved (
Fig. 6C
); these have been found in
Brazil
(Amazonas, Ceará, Goiás, Maranhão, Mato Grosso, Minas Gerais, Pará, Pernambuco, Piauí, Rio Grande do Norte),
Colombia
(Casanare), and
Venezuela
(
Bolívar
) (
Fig. 8B
). The distribution range of the two patterns also overlaps in several localities in
Brazil
(Amazonas, Maranhão, Mato Grosso, Minas Gerais, Pará, Pernambuco, Piauí) and
Venezuela
(
Bolívar
), being also those populations polymorphic (
Fig. 8B
)
.
FIGURE 2
. Details of the external morphology in species of
Pseudocanthon
Bates, 1887
.
A
,
P. vazdemelloi
new species
pronotal disc;
B
,
P
.
xanthurus
(
Blanchard, 1847
)
pronotal disc;
C
,
P
.
vazdemelloi
new species
detail of left lateral of elytron;
D
,
P
.
pantanensis
new species
detail of left lateral of elytron (arrow indicates eighth elytral stria present);
E
,
P
.
perplexus
(
LeConte, 1847
)
protibia in dorsal view (arrow indicates median emargination);
F
,
P
.
xanthurus
(
Blanchard, 1847
)
protibia in dorsal view;
G
.
P
.
pantanensis
new species
pygidium;
H
,
P
.
chaquensis
new species
pygidium. Scale bar: 0.5 mm (2A–D); 0.2 mm (2E–H).
FIGURE 3
. South American species of
Pseudocanthon
Bates, 1887
, dorsal view.
A
,
Pseudocanthon perplexus
(
LeConte, 1847
)
;
B–C
,
P
.
xanthurus
(
Blanchard, 1847
)
;
D
,
P
.
vazdemelloi
new species
;
E
,
P
.
pantanensis
new species
;
F
,
P
.
chaquensis
new species
. Scale bar: 1 mm (3A–E); 2 mm (3F).
We have not noticied any strict association between patterns of body colour and male genital morphology. In general, specimens with orange spots only at the base and apex of elytra (pattern 1;
Fig. 3B
) usually have aedeagus of pattern A (
Fig. 9
). However, in Maranguape (Ceará), Chapadinha (Maranhão), and Cordisburgo (Minas Gerais), some specimens have genitalia of pattern B (
Fig. 9
). Specimens with orange spots on almost the entire lateral edge of the body (pattern 2;
Fig. 3C
), on the other hand, usually present aedeagus of pattern B (
Fig. 9
). However, in Teresina (Piauí) and Caicara del Orinoco (
Venezuela
), specimens have aedeagus of pattern A (
Fig. 9
).
The
P. xanthurus
species concept herein recognized is a hypothesis based on the previously cited morphological and geographical evidence. Intraspecific variation in body colour patterns is common and was also recorded in many species of Deltochilini (
Silva
et al
. 2015
;
Valois
et al
. 2015
;
Silva & Valois 2019
). Furthermore, the morphological variations found by us between genital patterns A and B in
P. xanthurus
(
Figs. 5B–C
,
6B–C
) are weak if they were compared with the differences in aedeagus and endophallites between the recognized species of
Pseudocanthon
(
Figs. 5–6
).
FIGURE 4
. South American species of
Pseudocanthon
Bates, 1887
, ventral view.
A
,
Pseudocanthon perplexus
(
LeConte, 1847
)
;
B
,
P
.
xanthurus
(
Blanchard, 1847
)
;
C
,
P
.
vazdemelloi
new species
;
D
,
P
.
pantanensis
new species
(arrows indicate angulation at posterior edge of metafemur);
E
,
P
.
chaquensis
new species
(arrows indicate angulation at posterior edge of metafemur). Scale bar: 1 mm (3A–E); 2 mm (3F).
Alternatively, we are aware that each genital morph may represent a separate species lineage, both sympatric, and that each of these lineages can be polychromic. Some explanations for this pattern of shared polymorphism between sister species lineages can be the following: 1—incomplete lineage sorting, also called deep coalescence, with both descendant species inherited a polymorphism already present in the ancestral species; 2—parallelism, in which both descendant species inherited the same genetic “propensity” (in terms of genetic channelling, or biased variation) to express the character; 3—horizontal transfer (including hybridization), in which genes have somehow broken the confines of the close species lineages and moved horizontally through jumps between them (
Maddison 1997
;
Avise 2000
). All those genetic populational mechanisms mentioned above can generate those morphological patterns observed by us. However, as the present study was only based on morphological and geographic evidence, this is not part of the scope of our work, being practically impossible to test here these scenarios and confront them with our assumption. We state that a phylogeographical approach can be useful to test our species assumption. Future studies can corroborate if the morphological variation mentioned here reflects some phylogeographic structure or resulted from ecological responses to different environmental conditions along distributional range of this supposed species. Thus, we chose to describe at this time these patterns only as an intraspecific variation of
P. xanthurus
.