A new species and first record of Vates Burmeister, 1838 from the Atlantic Rainforest (Mantodea: Vatinae)
Author
Rivera, Julio
45A27931-0384-4E29-8D22-66B6FE37E657
Unidad de Investigación en Entomología y Medio Ambiente, Universidad San Ignacio de Loyola, Avenida La Fontana 550, La Molina, Lima, Peru.
jrivera@usil.edu.pe
Author
Herculano, João Felipe
4F583CB4-7B12-4C4D-9316-FCEC24BF7D8C
Instituto de Pesquisas, Jardim Botânico do Rio de Janeiro, Laboratório de Fitossanidade, Brazil.
jfherculano@gmail.com
Author
Lanna, Leonardo Moutinho
BF12F0C8-1EB4-4857-B9E5-72E16D48ED7B
Instituto de Pesquisas, Jardim Botânico do Rio de Janeiro, Laboratório de Fitossanidade, Brazil.
leonardomlanna@gmail.com
Author
Cavalcante, Sávio
23A50810-AAC2-4A3C-9C90-2D115BD1D9CF
Instituto de Pesquisas, Jardim Botânico do Rio de Janeiro, Laboratório de Fitossanidade, Brazil.
savcavalcante@gmail.com
Author
Teixeira, Maria Lúcia França
4C66B903-2AB3-45E9-BD6F-F015A55EC242
Instituto de Pesquisas, Jardim Botânico do Rio de Janeiro, Laboratório de Fitossanidade, Brazil.
malu@jbrj.gov.br
text
European Journal of Taxonomy
2020
2020-01-28
598
1
25
journal article
10.5852/ejt.2020.598
7cce35d8-9550-4d28-acc6-259a5733e207
2118-9773
3659604
8BE6D600-5575-415B-8D90-6041B74408C0
Vates phoenix
sp. nov.
urn:lsid:zoobank.org:act:
EB7BB1D8-786F-401C-B408-2D41260C7797
Figs 2–5
,
6
A–H, 8–9;
Table 1
Diagnosis
The new species can be easily recognized by the following combination of characters: i) cuticular projections above lateral ocelli strongly reduced (almost entirely missing in some specimens); ii) antennomeres of males asymmetrical (s-shaped); iii) hindwing of females with large, yellowish white and partially opaque area that spreads over most or part of the membrane; iv) anterodorsal lobe of hind tibiae at least 50% the length of tibial length (i.e., not narrowly restricted to its middle section).
Etymology
The specific epithet refers to the Phoenix, a mythical, immortal creature that is born again from its own ashes after being consumed by fire. The new species is a homage to the Museu Nacional of
Rio de Janeiro
, which was destroyed during a massive fire on
September 2, 2018
. The entire entomological collection, representing more than 5 million specimens, was destroyed, including all praying mantis specimens. Only a few specimens of
Vates
borrowed in the context of this study, including our new species, survived the event.
Vates phoenix
sp. nov.
thus symbolically attempts to link the past and the future of the Museu Nacional, as it represents the rebirth of the
Mantodea
collection and our hopes for the revival of an even stronger institution in the not too distant future.
Material examined
Holotype
BRAZIL
•
♂
;
Rio de Janeiro
,
Valença
,
Fazenda Recanto
;
22°07′15.3″ S
,
43°51′01.2″ W
; alt.
560 m
;
15 Nov. 2015
;
Projeto Mantis
leg.;
white cloth light trap
;
MNRJ-ENT6-28441
.
Allotype
BRAZIL
•
1 ♀
;
Rio de Janeiro
,
Rio de Janeiro
City
,
Jardim Botânico
do
Rio de Janeiro
; 22°96′73.717″ S,
43°22′50.381″ W
; alt.
7 m
;
29 May 2018
;
M.L.F. Teixeira
leg.; manual collection;
MNRJ-ENT6-28442
.
Paratypes
BRAZIL
–
Rio de Janeiro
•
1 ♂
; same collection data as for holotype;
MNRJ-ENT6-28445
•
1 ♂
; same collection data as for holotype;
31 Dec. 2016
;
white cloth light trap
;
William Moura
leg.;
MNRJ-
ENT6-28443
•
2 ♂♂
;
Reserva Ecológica de Guapiaçu
,
Cachoeiras de Macacu
;
22°27′10.309″ S
,
42°46′13.011″ W
; alt.
37 m
;
18 Dec. 2017
;
Projeto Mantis
leg.;
white cloth light trap
;
MNRJ-
ENT6-28446
,
MNRJ-ENT6-28447
•
1 ♂
;
Rio de Janeiro
City,
Corcovado
;
22°57′06″ S
;
43°12′37″ W
; alt.
600 m
;
Jan. 1936
;
D. Mendes
leg.;
MNRJ-ENT6-28448
•
3 ♂♂
;
Angra dos Reis
,
Jussaral Train Station
(note: now in ruins, the station closed down in 1996);
22°56′26″ S
,
44°16′26″ W
; alt.
351 m
;
Sep. 1934
;
D. Mendes
leg.; MNRJENT6-28450,
MNRJ-ENT6-28453
,
MNRJ-ENT6-28454
•
3 ♂♂
; same collection data as for preceding;
Sep. 1935
;
D. Mendes
leg;
MNRJ-ENT6-28449
,
MNRJ-ENT6-28451
,
MNRJ-ENT6-28452
•
1 ♀
; same collection data as for allotype;
May 1935
;
MNRJ-ENT6-28455
. –
São Paulo
•
1 ♂
;
Angatuba
;
22°56′26″ S
,
44°16′26″ W
; alt.
7 m
;
Nov. 1917
;
A. Marques
leg.;
MNRJ-
ENT6-28456
.
Description
Male
(
holotype
; MNRJ-ENT6-28441)
HABITUS. Live specimen (
paratype
) in
Fig. 2A
; pinned specimen (
holotype
) in
Fig. 2C.
MEASUREMENTS. See
Table 1
(specimen FR01).
HEAD. Eyes rounded (
Fig. 3A
). Vertex flat, higher than imaginary line connecting dorsal margin of compound eyes. Juxtaocular bulges flat, aligned to vertex. Ocellar tubercles (
Fig. 3A
) bearing two poorly developed conical projections, one over each lateral ocellus, only scarcely spaced (variation of this character across examined specimens is shown in
Fig. 3
B–E). Central ocellus elliptical, lateral ocelli rounded. Antennae (
Fig. 3
F–H) with scape and pedicel light brown, rest of antennae dark brown, proximal-most antennomeres cylindrical, eventually turning strongly asymmetrical, conferring antennae with a pectiniform appearance, distal-most antennomeres more or less filiform. Lower frons (
Fig. 3A
) sub-pentagonal, wider than high and with upper margin arcuate, surface smooth, concave and medially darkened. Maxillary palps light brown. Inner margin of labial palpi dark, basal-most segment with dark spots.
THORAX. Pronotum (
Fig. 3I
) elongated, metazona triangular in cross-section. Supracoxal dilation moderately pronounced and broadly rounded; ratio metazona / prozona = 4.12 (variation across specimens 4.5–3.75). Distal margin of prozona uniformly curved, margins with small, spine-like, blunt tubercles, denser along prozona than along metazona and mostly absent proximally. Dorsal surface of metazona keeled along its midline (keel more pronounced proximally). Pronotum predominantly green, except for darkened prozone and lateral margins of metazona. Variation in pronotal size and shape across examined specimens shown in
Fig. 3
J–L.
PROTHORACIC LEGS. Forecoxae triangular in cross-section; ventral margin pale green, except for a small and distally positioned dark spot on its anterior aspect, and a larger preapical spot posteriorly; dorsal margin bearing five spine-like, darkened tubercles interleaved with smaller, paler ones; anterior aspect of forecoxae light colored, darked apically, rest of structure dark brown. Spination formula: F =4DS/14AvS/4PvS; T =14AvS/8(R)–10(L)PvS. Forefemora light brown, three-banded, with a small dark spot near trochanter; dorsal margin of forefemora slightly sinuous; discoidal spines I, II and III mostly pale with darkened apex, spine IV entirely dark; anteroventral spines II, IV, VI, X and XII slightly reclined and entirely dark, spine XV larger than the others, entirely dark and not curved; remaining spines smaller, pale and with darkened apex; genicular spines developed and present on both sides of femora; tibial spur groove located in proximal ¼ of femora. Foretibiae light brown, dorsally three-banded.
Table 1.
Standard body measurements of
Vates phoenix
sp. nov.
(in mm). Foretibial measurement D1 does not include the tibial spur, whereas D2 does. Measurements under FR01 correspond to the holotype and JB01 to the allotype. Abbreviations: FR = Fazenda Recanto; JB = Jardim Botânico do Rio de Janeiro; RG = Reserva Ecológica do Guapiaçu; CO = Corcovado; JU = Jussaral; AN =Angatuba. Standard measurements
sensu
Brannoch
et al
. (2017)
.
Character |
FR01 |
JB01 |
FR02 |
FR03 |
RG01 |
RG02 |
CO |
JU01 |
JU02 |
JU03 |
JU04 |
JU05 |
JU06 |
AN |
JB02
|
Sex |
♂
|
♀
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♂
|
♀
|
Body length |
64.0 |
67.5 |
66.0 |
66.0 |
64.0 |
64.5 |
64.0 |
67.0 |
68.0 |
64.0 |
68.5 |
67.5 |
68.5 |
66.0 |
65.0 |
Head width |
5.0 |
6.9 |
6.0 |
6.0 |
6.0 |
6.0 |
5.5 |
6.0 |
6.0 |
6.0 |
6.0 |
6.0 |
6.5 |
6.0 |
7.0 |
Head length |
2.5 |
3.0 |
2.5 |
2.5 |
2.5 |
2.5 |
2.0 |
2.0 |
2.5 |
2.5 |
2.5 |
2.5 |
2.5 |
2.3 |
3.0 |
Lower frons width |
1.5 |
2.5 |
1.5 |
1.5 |
1.5 |
1.5 |
damaged |
1.5 |
1.0 |
1.0 |
2.0 |
1.5 |
1.0 |
1.0 |
2.5 |
Lower frons length |
1.0 |
1.4 |
1.0 |
1.0 |
1.0 |
1.0 |
1.5 |
1.0 |
1.0 |
0.6 |
1.0 |
1.0 |
1.0 |
1.0 |
2.5 |
Pronotal length |
20.5 |
23.2 |
22.5 |
21.3 |
20 |
19 |
19.5 |
21.5 |
20.5 |
19.5 |
21.5 |
21.5 |
21.5 |
21 |
23 |
Prozone |
4.0 |
4.2 |
4.5 |
4.0 |
4.0 |
4.0 |
3.5 |
3.5 |
3.5 |
3.5 |
4.0 |
4.0 |
4.0 |
4.0 |
4.1 |
Metazone |
16.5 |
19 |
18.0 |
17.3 |
16.0 |
15.0 |
16.0 |
18.0 |
17.0 |
16.0 |
17.5 |
17.5 |
17.5 |
17.0 |
18.9 |
Pronotal width |
4.0 |
5.2 |
4.0 |
4.5 |
4.0 |
4.0 |
4.0 |
4.5 |
4.0 |
4.0 |
4.3 |
4.0 |
4.0 |
4.0 |
5.0 |
Ratio metazone/ prozone |
4.12 |
3.65 |
4.50 |
3.85 |
4.00 |
3.75 |
4.00 |
4.00 |
4,25 |
4.00 |
04.05 |
4.37 |
4.37 |
4.37 |
3.78 |
Forewing length |
42.0 |
40.3 |
44.0 |
42.3 |
41.0 |
43.5 |
40.0 |
43.0 |
43.5 |
43.0 |
44.0 |
43.0 |
43.5 |
40.5 |
40.0 |
Forewing width |
9.0 |
10.0 |
9.0 |
8.5 |
8.0 |
8.5 |
- |
- |
- |
- |
- |
- |
- |
- |
- |
Hindwing length |
37.5 |
35.2 |
38.0 |
37.6 |
39.5 |
39.0 |
- |
- |
- |
- |
- |
- |
- |
- |
- |
Hindwing width |
17.0 |
18.6 |
16.0 |
17.5 |
16.0 |
16.0 |
- |
- |
- |
- |
- |
- |
- |
- |
- |
Forecoxal length |
11.0 |
14.4 |
11.0 |
11.0 |
11.0 |
11.0 |
11.0 |
11.0 |
11.0 |
10.6 |
11.0 |
11.5 |
11.5 |
11.5 |
13.5 |
Forecoxal width |
1.5 |
2.5 |
1.5 |
1.6 |
1.5 |
1.5 |
2.0 |
2.0 |
1.5 |
1.5 |
2.0 |
2.0 |
2.0 |
1.6 |
2.0 |
Forefemoral length |
14.0 |
15.3 |
14.5 |
12.5 |
12.5 |
11.5 |
12.0 |
13.0 |
13.0 |
12.3 |
12.6 |
13.0 |
12.6 |
12.5 |
15.0 |
Forefemoral width |
1.6 |
2,3 |
2.0 |
1.6 |
1.6 |
1.6 |
1.5 |
2.0 |
2.0 |
2.0 |
2.0 |
2.0 |
2.0 |
1.6 |
3.0 |
Foretibial length (D1) |
5.0 |
6.0 |
5.5 |
5.0 |
4.6 |
4.5 |
4.5 |
5.0 |
5.0 |
5.0 |
5.0 |
5.0 |
5.0 |
4.5 |
5.5 |
Foretibial length (D2) |
7.0 |
9.0 |
7.5 |
7.33 |
7.0 |
7.0 |
7.0 |
7.5 |
7.0 |
7.0 |
7.5 |
7.0 |
7.33 |
7.33 |
8.0 |
Mesocoxa |
4.0 |
6.5 |
5.5 |
4.0 |
4.0 |
4.0 |
4.0 |
4.0 |
4.5 |
4.0 |
4.5 |
4.5 |
5.0 |
4.5 |
6.0 |
Mesofemur |
11.0 |
12 |
11.0 |
11.0 |
10.0 |
10.0 |
10.0 |
10.0 |
10.0 |
10.0 |
11.0 |
11.0 |
11.0 |
11.0 |
12.3 |
Mesotibia |
8.3 |
9,6 |
9.0 |
8.6 |
9.0 |
8.0 |
8.0 |
8.0 |
8.5 |
9.0 |
9.0 |
8.5 |
8.6 |
8.6 |
10.0 |
Mesotarsus |
6.0 |
7,7 |
6.0 |
6.0 |
6.0 |
6.0 |
6.0 |
5.0 |
6.0 |
6.5 |
6.0 |
5.5 |
6.5 |
6.0 |
7.0 |
Metacoxa length |
4.0 |
5.7 |
4.0 |
4.5 |
4.5 |
4.0 |
4.0 |
4.0 |
4.0 |
4.0 |
4.5 |
4.5 |
5.0 |
4.5 |
5.5 |
Metafemur length |
13.5 |
14,3 |
15.0 |
13.5 |
13.5 |
13.0 |
12.0 |
13.0 |
12.0 |
12.0 |
14.0 |
13.5 |
13.66 |
13.6 |
15.3 |
Metatibia length |
11.5 |
14,3 |
11.5 |
12.0 |
12.0 |
11.5 |
11.0 |
12.0 |
11.5 |
12.0 |
12.5 |
12.0 |
12.0 |
11.6 |
13.0 |
Metatarsus length |
6.0 |
8.5 |
7.0 |
7.5 |
7.0 |
7.5 |
6.5 |
7.0 |
7.0 |
7.5 |
8.5 |
8.0 |
8.5 |
8.5 |
8.0 |
WINGS. Forewings (
Fig. 2C
) surpass abdomen by ¼ of its length in resting position. Costal area distally tapering, membrane opaque and predominantly green, with a yellowish longitudinal strip along margin of costal vein; discoidal area entirely hyaline with yellowish to brownish longitudinal veins, apex slightly darkened, with more densely reticulate venation. Hindwings (
Fig. 2C
) hyaline and colorless, yellowish to brownish longitudinal veins, apex of discoidal area densely reticulated and dark brown.
Fig. 2.
Adults of
Vates phoenix
sp. nov.
A–B
. Live specimens photographed in a studio.
A
. Paratype, ♂, from Reserva Ecológica de Guapiaçu (MNRJ-ENT6-28446).
B
. Allotype, ♀, from Jardim Botânico do Rio de Janeiro (MNRJ-ENT6-28442). –
C–D
. Pinned adults.
C
. Holotype, ♂, from Fazenda Recanto (MNRJ-ENT6-28441).
D
. Allotype, ♀ (MNRJ-ENT6-28442). Scale bars: C–D = 10 mm.
Fig. 3.
Vates phoenix
sp. nov.
, male morphology.
A
. Head, frontal view, holotype (MNRJ-ENT6-28441). –
B–E
. Variation in ocellar tubercles across localities (not to scale).
B
. Fazenda Recanto, holotype (MNRJ- ENT6-28441).
C.
Reserva Ecológica de Guapiaçu, paratype (MNRJ-ENT6-28447).
D
. Jussaral, paratype (MNRJ-ENT6-28449).
E
. Corcovado, paratype (MNRJ-ENT6-28448). –
F–H
. Antenna, highlighting sectional variation of antennomeres.
F
. Basal section.
G
. Mid section.
H
. Apex. –
I–L
. Pronota, highlighting size variation.
I
. Fazenda Recanto, holotype (MNRJ-ENT6-28441).
J
. Reserva Ecológica de Guapiaçu, paratype (MNRJ-ENT6-28447).
K
. Jussaral, paratype (MNRJ-ENT6-28449).
L
. Angatuba, paratype (MNRJ-ENT6-28456). Scale bars: A = 1 mm; I–L = 5 mm.
MESO- AND METATHORACIC LEGS. Mesothoracic femora (
Fig. 4A
) with a marked posteroventral keel that forms two evident lobes: a proximal, elongated but scarcely produced lobe, and a pre-apical, short and produced lobe whose shape resembles that of a shark’s dorsal fin (variation of this character across specimens can be seen in
Fig. 4
B–D). Metathoracic femora (
Fig. 4E
) with a marked posteroventral keel that forms a single, preapical lobe, also shaped like a shark’s dorsal fin (variation of this character across specimens can be seen in
Fig. 4F
). Mesothoracic tibiae (
Fig. 4G
) with two medial lobes: a produced, anterodorsal lobe almost as long as tibia itself, and an anteroventral one, almost equally produced but clearly shorter (variation of this character across specimens can be seen in
Fig. 4
H–J). Metathoracic tibiae (
Fig. 4K
) with two elongated lobes, anterodorsal one longer and wider than anteroventral one, and also more prominent than its homologue on mesothoracic tibiae (variation of this character across specimens can be seen in
Fig. 4
L–M).
ABDOMEN. Slightly compressed dorsoventrally, widest between segments III–V. Tergites I–III with a whitish coloration, remaining tergites darkened. Supraanal plate triangular in shape, wider than longer, apex medially notched, forming two small, lateral lobes. Cerci cylindrical and elongated, not surpassing apex of subgenital plate, last cercomere conical.
GENITALIA (note:
holotype
was not dissected; the following description corresponds to genital structures of
paratypes
). Left phallomere (
Fig. 5
A–E): sclerite L4B longer than wide, its left margin projects anteriorly; anterior process (afa) glabrous, sinuous, basal half broad but tapering distally, apex more strongly sclerotized, with sharp, pointy apex; posteromesal lobe (loa) elongated and sinuous, glabrous, lacking projections; posterior process (Paa) elongated, slightly curved left, apex curved anteriorly. Ventral phallomere (
Fig. 5
F–I): sclerite L4A roughly oval, proximal left margin slightly sinuous and strongly sclerotized, forming a small, medial projection followed by a membranous notch; posterior process (Pda) elongated, apex strongly scletorized and tapering distally, uniformly curved right, although its distal third curves anteriorly. Right phallomere (
Fig. 5
J–N): roughly triangular, distal margin folded anteriorly; anterior apodeme (an) of sclerite R3 elongated, distally broadened, bearing a small, basal process near its articulation to main posterior lobe (fda), the latter with a relatively short and broad dextral extension (bm); process anteromesal to Pia (Pva) elongated and finger-like, slightly curved ventrally, strongly sclerotized, with a moderately truncated apex; process posterolateral to Pva (Pia) elongated and well sclerotized.
Female
(
allotype
; MNRJ-ENT6-28442)
HABITUS. Live specimen (
allotype
) in
Fig. 2B
; pinned specimen (
allotype
) in
Fig. 2D.
MEASUREMENTS. See
Table 1
(specimen JB01).
HEAD. Eyes rounded (
Fig. 6A
). Vertex flat, higher than imaginary line connecting dorsal margin of compound eyes. Parietal suture darkened, juxtaocular bulges flat and aligned to vertex. Ocellar tubercles (
Fig. 6B
) bearing two poorly developed conical projections, one over each lateral ocellus and barely spaced (variation of this character across specimens can be seen in
Fig. 6C
). Ocelli rounded, lateral ocelli twice as large as central ocellus. Antenna filiform, scape and pedicel light brown, flagellomeres dark brown. Lower frons sub-pentagonal, wider than high, smooth, medially darkened, upper margin arcuate, smooth and concave. Maxillary palps light brown; inner margin of labial palpi darkened, basalmost segment with dark spots.
THORAX. Pronotum elongated (
Fig. 6D
), metazona triangular in cross-section. Supracoxal dilatation moderately pronounced and broadly rounded; ratio metazona / prozona = 3.65 (additional
paratype
female = 3.78). Distal margins of prozona uniformly curved, margins with small, spine-like, blunt tubercles, denser along prozona than along metazona and mostly absent proximally. Dorsal surface of metazona keeled along its midline (keel more pronounced proximally). Pronotum predominantly green, except for darkened prozone and lateral margins of metazona.
Fig. 4.
Mid- and hind legs of
Vates phoenix
sp. nov.
, highlighting variation across male specimens.
A–D
. Mid femora.
A
. Holotype from Fazenda Recanto (MNRJ-ENT6-28441).
B
. Paratype from Reserva Ecológica de Guapiaçu (MNRJ-ENT6-28447).
C
. Paratype from Jussaral (MNRJ-ENT6-28449).
D
. Paratype from Angatuba (MNRJ-ENT6-28456). –
E–F
. Hind femora.
E
. Holotype from Fazenda Recanto (MNRJ-ENT6-28441).
F
. Paratype from Angatuba (MNRJ-ENT6-28456). –
G–J
. Mid tibiae.
G
. Holotype from Fazenda Recanto (MNRJ-ENT6-28441).
H
. Paratype from Reserva Ecológica de Guapiaçu (MNRJ-ENT6-28447).
I
. Paratype from Jussaral (MNRJ-ENT6-28449).
J
. Paratype from Angatuba (MNRJ-ENT6-28456). –
K–M
. Hind tibiae.
K
. Holotype from Fazenda Recanto (MNRJ- ENT6-28441).
L
. Paratype from Jussaral (MNRJ-ENT6-28449).
M
. Paratype from Angatuba (MNRJ- ENT6-28456). Scale bar: A–M = 5 mm.
Fig. 5.
Male genital structures of
Vates phoenix
sp. nov.
, highlighting variation across specimens.
A–B
. Left phallomere, paratype (MNRJ-ENT6-28445).
A
. Dorsal view.
B
. Ventral view. –
C–E
. Left phallomere, partial ventral view.
C
. Paratype (MNRJ-ENT6-28445).
D
. Paratype (MNRJ-ENT6-28443).
E
. Paratype (MNRJ-ENT6-28446). –
F–I
. Ventral phallomere.
F
. Dorsal view, paratype (MNRJ- ENT6-28445).
G
. Partial ventral view, same as in F.
H
. Partial dorsal view, same as in F.
I
. Partial ventral view, paratype (MNRJ-ENT6-28446). –
J–L
. Right phallomere, dorsal view.
J
. Fazenda Recanto, paratype (MNRJ-ENT6-28445).
K
. Reserva Ecológica de Guapiaçu, paratype (MNRJ-ENT6-28446).
L
. Fazenda Recanto, paratype (MNRJ-ENT6-28443). –
M–N
. Right phallomere, details of ventral structures.
M
. Fazenda Recanto, paratype (MNRJ-ENT6-28443).
N
. Reserva Ecológica de Guapiaçu, paratype (MNRJ-ENT6-28446). Scale bars: A–B, F, J–L = 2 mm; C–E, G–I = 1 mm; M–N = 500 μm.
PROTHORACIC LEGS. Forecoxae triangular in cross-section, ventral margin pale yellowish brown, except for small dark spot on its anterior end, and a larger, preapical one on its posterior aspect; dorsal margin with nine spine-like, darkened tubercles interleaved with smaller, paler ones; color patterning in general similar to that of males. Forefemora light brown, three-banded, and with a small spot in anterior region of each femur near trochanter. Dorsal margin of forefemora slightly sinuous. Spination formula of forelegs: F =4DS/15AvS/4PvS; T=15(R)–16(L)AvS/13(R)–11(L)PvS. Discoidal spines I, II and III pale with dark spots at their base and apex, spine IV entirely dark. Anteroventral spines II, IV, VI, VIII and X slightly reclined, spines I–XII entirely black, XIII and XIV black laterally and at apex, XV with black base and apex, spine XV largest, straight and darkened at its base and its tip; posteroventral spines black at their base and tips; a well developed genicular spine on each sides of femora; tibial spine groove located in proximal ¼ of femora. Foretibiae light brown, dorsally three-banded.
WINGS. Forewings (
Fig. 2D
) surpass abdomen by ¼ of their length in resting position. Forewings with membrane of costal area opaque and predominantñy green, with a yellowish longitudinal strip along margin of costal vein, distal-most portion of costal vein and membrane around radial vein darkened; discoidal area with green and opaque membrane, longitudinal veins mostly yellow, veins densely reticulated distally, stigma with a darkly pigmented spot; anal area small, mostly opaque, veins and membrane withish. Hindwing with narrow costal area, tapering distally and partially opaque, proximal half with membrane and veins whitish, although cells become dark brown distally; discoidal area largely hyaline with yellowish veins, proximal region of membrane whitish, distal portion smoky brown, opaque and heavily reticulated; anal area largely hyaline, veins and proximal area of membrane whitish, distal portion below discoidal area faint brown.
MESO- AND METATHORACIC LEGS. Mesothoracic femora (
Fig. 6E
) with a marked posteroventral keel that forms two evident lobes: a proximal one that is elongated but scarcely produced, and a pre-apical one that is short and produced, shaped like a shark’s dorsal fin. Metathoracic femora (
Fig. 6F
) with a marked posteroventral keel that forms a single preapical lobe, also shaped like a shark’s dorsal fin. Mesothoracic tibiae (
Fig. 6G
) with two medial lobes: a produced, anterodorsal one, almost as long as tibia itself, and an anteroventral one, almost equally produced but clearly shorter. Metathoracic tibiae (
Fig. 6H
) with two elongated lobes, anterodorsal one longer and wider than anteroventral one, and also more prominent than its homologue on mesothoracic tibiae.
ABDOMEN. Fusiform, slightly flattened dorsoventrally, widest between segments III–V, brownish with some contrasting spots in middle of each tergite. Cerci elongated, surpassing apex of subgenital plate, cercomeres cylindrical.
Differential diagnosis
Svenson
et al
. (2015)
listed all species of
Vates
they considered valid, totaling 13 spp. All these species are herein discussed in relation to our new species. The males of
V. phoenix
sp. nov.
share with those of
V. biplagiata
Sjöstedt, 1930
,
V. luxuriosa
Beier, 1958
,
V. amazonica
(Westwood, 1889)
,
V. pectinicornis
(Stål, 1877)
,
V. foliata
(Lichtenstein, 1802)
and
V. lobata
(
Fabricius, 1798
)
the asymmetrical, s-shaped antennomeres, whereas the females share with those of
V. serraticornis
Stål, 1877
,
V. festae
Giglio-Tos, 1914
,
V. weyrauchi
Beier, 1958
, and likely also
V. boliviana
Giglio-Tos, 1914
(which remains known from males only) the yellowish white tinge of the hindwing membrane. None of the preceding species has both males with s-shaped antennomeres and females with yellowish white hindwings. Therefore,
V. phoenix
sp. nov.
possesses a unique combination of character states unknown in other species of
Vates
, making its identification straightforward. In the absence of either one of the sexes for effective comparisons, the reduced ocellar tubercles of
V. phoenix
sp. nov.
represent a distinct character state for the species, common to both sexes, and unique among members of the genus (compare, for instance, with
Fig. 6
I–J). Our new species can also be easily distinguished from
V. pectinata
Saussure, 1871
and
V
.
chopardi
(Deeleman-Reinhold, 1957)
for lacking the dorsal, preapical lobe of the forefemora, a distinct character shared by the latter two species (
Fig. 7
) — which
Roy (2012)
suggested as likely synonyms. Additionally, we were unable to compare
V. phoenix
sp. nov.
with the
type
specimen of
Vates lobata
(
Fabricius, 1798
)
. This species was originally described from “Cajennae” (= Cayenne,
French Guiana
) as
Mantis
lobata
(
Fabricius 1798
)
in the ‘Supplementum’ to ‘Entomologica Systematica’ — not to be confused with
Mantis
lobata
Fabricius, 1781
, a synonym of
Harpagomantis tricolor
(Linnaeus, 1758) (Galinthiadidae)
sensu
Beier (1934)
. Examination of the Banks collection (JR) housed by the Natural History Museum, London (containing a sizable portion of Fabricius
types
) did reveal the presence of
Mantis
lobata
Fabricius, 1781
, though no specimen attributable to
Mantis
lobata
Fabricius, 1798
(i.e.,
Vates
) was found there. The taxa that
Fabricius (1798)
described from “Cajennae” in his ‘Supplementum’ were owned by Louis Augustin Guillaume Bosc d’Antic (N. Kristensen, pers. com. 2011), a French naturalist whose insect collection, in part studied by Fabricius, dispersed across European natural history institutions after his death in 1828 (
Notton 2007
). A good portion of this material eventually made it to Paris and Geneva; however, examination of these and other European collections also containing some of Fabricius’
types
, such as the Zoological Museum in Copenhagen (N. Kristensen, pers. com. 2011) and the Hunterian Museum in Glasgow (online catalogue: http://collections.gla.ac.uk/) did not turn up any specimen attributable to the
type
of
Mantis
lobata
Fabricius, 1798
. This specimen is most likely lost. Regardless of the fate of the
type
specimen, the new species can be distinguished from
Vates lobata
on the basis of its distribution, as
French Guiana
and Rio de Janeiro are biogeographically distant and unrelated regions that do not share any praying mantis species. Finally,
Vates obscura
Toledo Piza, 1983
, listed as valid in
Svenson
et al
. (2015)
, had already been synonymized with
V. biplagiata
in an earlier publication (
Agudelo & Rivera 2015
), and thus we conclusively remove this species from the
Vates
checklist.
Fig. 6.
Vates phoenix
sp. nov.
, female morphology (both specimens are from Jardim Botânico do Rio de Janeiro), and head morphology of
Vates
spp.
A
. Head, frontal view, allotype (MNRJ-ENT6-28442). –
B–C
. Variation in ocellar tubercles (not to scale).
B
. Allotype (collected in 2018; MNRJ-ENT6-28442).
C
. Paratype (collected in 1935; MNRJ-ENT6-28455). –
D
. Pronotum, dorsal view, allotype (MNRJ- ENT6-28442). –
E–H
. Mid- and hind legs, allotype (MNRJ-ENT6-28442).
E
. Mid-femur, detail of distal section.
F
. Hind femur, detail of distal section.
G
. Mid-tibia.
H
. Hind tibia. –
I
.
Vates biplagiata
Sjöstedt, 1930
, head, frontal view (male specimen from Peru).
J
.
Vates weyrauchi
Beier, 1958
, head, frontal view (male specimen from Peru). Scale bars: A = 1 mm; D–H = 5 mm.
Systematic remarks
Morphological comparison of male genitalia provided insights on the affinities of the new species. The only species whose male genital structures are known are
Vates chopardi
(
Lombardo 2000
: figs 25–27), and
V. biplagiata
and
V. festae
(
Medellín & Salazar 2011
: fig. 14). Morphological comparisons between
V. phoenix
sp. nov.
and
V. biplagiata
genitalia showed that both species share a strong, proximally bent afa, whereas the accentuated sigmoidal shape of the same can also be observed in
V. chopardi
, though in the latter the afa is not bent to the same degree as in
V. biplagiata
. Differences in genital structures are more accentuated between
V. phoenix
sp. nov.
and
V. festae
, the latter with a much straighter afa and, and a shortened and robust Pda on the ventral phallomere. Coincidently, the phylogeny of
Vatini
proposed in
Svenson
et al
(2015)
recovered three main clades within
Vates
: one containing
V. chopardi
, sister to another clade comprised of
V. festae
and
V. weyrauchi
, altogether sister to a more diverse clade containing
V. biplagiata
. Our analysis of external morphology and existing reports of male genital structures thus suggests a closer affinity to members of this latter clade, which in
Svenson
et al
. (2015)
also included
V. luxuriosa
,
V. amazonica
,
V. pectinicornis
and two additional, unidentified species. Further phylogenetic studies are necessary to test this hypothesis and resolve evolutionary affinities with all members of
Vates
.
With the description of our new species and the clarification of previous records, the following species are considered valid: 1)
V. amazonica
(Westwood, 1889)
; 2)
V. biplagiata
Sjöstedt, 1930
; 3)
V. boliviana
Giglio-Tos, 1914
; 4)
V. chopardi
(Deeleman-Reinhold, 1957)
; 5)
V. festae
Giglio-Tos, 1914
; 6)
V. foliata
(Lichtenstein, 1802)
; 7)
V. lobata
(
Fabricius, 1798
)
; 8)
V. luxuriosa
Beier, 1958
; 9)
V. pectinata
Saussure, 1871
; 10)
V. pectinicornis
(Stål, 1877)
; 11)
V. serraticornis
Stål, 1877
; 12)
V. weyrauchi
Beier, 1958
; 13)
V. phoenix
Rivera
et al
.
sp. nov.
The possible synonymy between
V. pectinata
and
V
.
chopardi
, as suggested by
Roy (2012)
, is pending confirmation.
Fig. 7. A
.
Vates pectinata
Saussure, 1871
, holotype, ♂, and labels (Muséum d’histoire naturelle de la Ville de Genève, Geneva, Switzerland); note: the male and female specimens portrayed in fig. 8d–c of Svenson
et al
. (2016) and reported as
Vates pectinata
cannot be attributed to this species.
B
.
Lobovates chopardi
Deeleman-Reinhold, 1957
, holotype, ♂, and labels (National Museum of Natural History, Leiden, the Netherlands); note: wings are colorless and hyaline in the actual specimen, whitish tinge is due to a photo originally taken against a white background.
Natural history and behavior
Rehn (1935)
noted that species of
Vates
were difficult to come across in nature. Things have not changed much since Rehn’s times, as there still is very little information in the literature regarding the biology of
Vates
.
Below we present natural history information of
Vates
spp resulting from our own observations and literature accounts, and summarize this information to place it in the context of our new species.
Neither nymphs nor adults of
V. phoenix
sp. nov.
were ever found in their natural habitat. Unlike members of other sympatric genera, all broadly sampled across
Rio de Janeiro
by Projeto Mantis’ research team, we were unable to locate specimens of
Vates
in any manually sampled plant formations in the three years that this project spanned. This suggest that
Vates
spp. likely prefer higher layers within the vegetation as perching and hunting grounds, thus making their collection at ground level difficult. In fact,
Dantas
et al.
(2008)
reported
Vates
spp. flying to canopy light traps
45 m
above the ground near Manaus,
Brazil
. Interestingly, the latter study reported collecting a few females, thus evidencing enhanced flying capabilities of this sex in
Vates
. Behavioral observations conducted on the only wild-caught female indeed showed that this sex is able to sustain controlled gliding for short distances while moving across perching sites. Our anatomical examinations of
V. phoenix
sp. nov.
revealed that females have relatively larger ocelli compared to other non-Vatini females, a pattern consistent with flying capabilities in female praying mantises and other insects (
Battiston
et al
. 2018
). Further, reduced sexual dimorphism in wing size and shape is evident in the examined specimens (e.g.,
Fig. 2
C–D). This likely explains why females of
Vates
are sometimes lured to light traps, as shown by
Dantas
et al
. (2008)
and corroborated by some of us through field work (JR, pers. obs.). Collecting efforts using this sampling method could eventually result in collecting additional female specimens of
V. phoenix
sp. nov.
Angatuba (São Paulo) and Reserva Ecológica de Guapiaçu (Rio de Janeiro) represent the westerneasternmost limits of
Vates phoenix
sp. nov.
based on the available data. Unfortunately, it is difficult to determine the extent of the original distribution of
V. phoenix
sp. nov.
and other sympatric species, as the original Atlantic Rainforest circumscribed by these localities is now heavily altered by urban development and farming, thus affecting this and likely other praying mantis species (
Rodrigues & Cancello 2013
). Sampled localities showed that
V. phoenix
sp. nov.
ranges from
7 to
600 m
. Distributional records compiled from the literature revealed that several species of
Vates
are found at mid- to higher elevations. For instance, the dominant species along the oriental slopes of the Peruvian / Ecuadorian Andes and neighbouring lowlands are
V. weyrauchi
(
100– 1000 m
),
V. luxuriosa
(
400– 1000 m
),
V. biplagiata
(
180 – 2900 m
) and
V. festae
(
1000 – 1700 m
) (
Lombardo & Agabiti 2001
;
Rivera & Vergara-Cobián 2017
), with the latter also found on the eastern slopes of the central and oriental cordilleras of central
Colombia
(
450– 2400 m
) (
Medellín & Salazar 2011
;
Svenson
et al
. 2015
) and in the vicinity of Manaus,
Brazil
(ca
90 m
) (
Dantas
et al
. 2008
). The relatively narrow vertical distribution of
V. phoenix
sp. nov.
is comparable to that of
V. amazonica
and
V. chopardi
, both predominantly lowland species mostly found below
500 m
, the former across the Amazon basin and the latter along the Mexican Pacific and Atlantic lowlands (
Svenson
et al
. 2015
). All this information suggests that several species of
Vates
have a broad altitudinal range (e.g.,
V. biplagiata
), and that the genus has a tendency to be more diverse at higher elevations. However, distributional records across the Amazonian lowlands are still scarce for most species and more surveys are necessary to infer patterns of vertical distribution across the genus.
Reared specimens preferred perching clinging upside-down, often keeping their raptorial forelegs stretched forward to form a 90º angle relative to the body (
Fig. 8
). This posture was assumed either
de novo
or enhanced in response to a nearby observer the insect possibly perceived as a potential threat.
Robinson (1969)
reported a similar behavior in
Pseudovates chlorophaea
(Blanchard, 1836)
(cited as
Phyllovates chlorophaea
therein) in
Panama
, regarding it as a combination of leaf and stick mimicry. More efforts are necessary to unveil further aspects of the natural history of
Vates
spp, and of
Vatini
in general.