Research Article |
Corresponding author: Leonela Olivera ( lolivera@fcnym.unlp.edu.ar ) Academic editor: Martin Husemann
© 2023 Leonela Olivera, María Cecilia Melo, Pablo M. Dellapé.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Olivera L, Melo MC, Dellapé PM (2023) Revisiting the South American Acanthocephalini (Hemiptera, Coreidae): Spilopleura Stål (status novum). Evolutionary Systematics 7(1): 35-50. https://doi.org/10.3897/evolsyst.7.94403
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The family Coreidae is composed of some of the larger terrestrial Heteroptera, with showy colors and expansions on the legs, the antenna or the pronotum. Among the Neotropical fauna, the tribe Acanthocephalini Stål, including 18 genera, is recognized by the strongly deflexed juga, and the conspicuously projecting tylus. In the same work in which Stål established the tribe, he described two new subgenera to the genus Acanthocephala Laporte (currently synonymized under it): A. (Metapodius) (replaced for Metapodiessa Kirkaldy) and A. (Spilopleura). In the present work, the subgeneric name Spilopleura is removed from synonymy under Acanthocephala and elevated to generic rank, and the species A. parensis (Dallas) and A. ochracea Montandon are transferred therein. The genus Spilopleura status nov. and both included species are redescribed and illustrated, including characters from male and female genitalia; and a distributional map is given with the first records of the species from Argentina, Colombia, Paraguay, and Peru.
Acanthocephala, Heteroptera, new genus, neotropical fauna
The Coreidae, or leaf-footed bugs, have a cosmopolitan distribution showing greater diversity in tropical and subtropical areas, and includes some of the larger terrestrial Heteroptera, often with striking colors and expansions on the hind femora or tibiae, pedicellus and/or basiflagellomere, and the pronotum (
Stål (1870) established the Neotropical tribe Acanthocephalini (originally as the subfamily “Acanthocephalina”) including eight genera: Acanthocephala Laporte, Empedocles Stål, Laminiceps Costa, Lucullia Stål, Thymetus Stål, Petalops Amyot and Serville, Plaxiscelis Spinola, and Stenocelidea Westwood. Subsequently, two of these genera (Plaxiscelis and Stenocelidea) were removed (
In the same work, Stål described two new subgenera in Acanthocephala: A. (Metapodius) including six species (later replaced for Metapodiessa by
At present, the genus Acanthocephala includes 26 species distributed from Canada to Argentina (
In the present work, we remove Spilopleura from synonymy under Acanthocephala and elevate it to generic rank including two species: A. parensis (Dallas), the only species originally included by Stål in the subgenus, and A. ochracea Montandon. The genus Spilopleura status nov. and both included species are diagnosed, redescribed, and illustrated, including characters from male and female genitalia. A distributional map of the species is also included.
Specimens were examined under an Olympus SZ1000 and a Nikon SMZ1000 stereomicroscope. Digital images were taken with a Micrometrics 391CU, 3.2m, Accu-Scope digital camera attached to the Nikon SMZ1000 stereomicroscope. Images were stacked using HeliconFocus 6.7.1 software. The genital capsule of males and genital segments of females were dissected and cleared with a saturated potassium hydroxide solution (10%), and in the case of the male genitalia, the aedeagus was also immersed in lactic acid for final clearing and better handling. Finally, the dissected structures were washed in distilled water, place in glycerin for observation and preserved in microvials attached to the same pin of mounted specimens. For the description of the genitalia the terminology used is that of
The following abbreviations are used for the institutions cited in the material examined:
We selected 23 measurements to described species, as follows:
total length : in lateral view, from apex of tylus to tip of abdomen;
head length : in dorsal view, from apex of tylus to posterior margin of ocellar tubercles;
max head width : in dorsal view, from outer margin of right eye to outer margin of left eye;
tylus length : in dorsal view, from apex of tylus to the beginning of post-tylus depression;
post-tylus depression length : in dorsal view, from tip to tip of depression;
eye width : in dorsal view, from inner margin to outer margin of an eye, maximum width;
interocular space : in dorsal view, distance between inner margin of eyes;
ocellar distance : in dorsal view, from outer margin of right ocellus to outer margin of left ocellus, maximum width;
interocelar space : in dorsal view, distance between inner margin of ocelli;
antennal articles length : from base to apex each;
antennal articles width : maximum width each;
length of labial segments : in lateral view, from base to apex each;
humeral angles width : in dorsal view, maximum width;
scutellum length : in dorsal view, from base to apex;
scutellum width : in dorsal view, maximum width at base of scutellum;
hemelytra length : in dorsal view, from base to apex;
profemur width : maximum width;
metafemur width : maximum width;
metafemur length : from base to apex;
metatibia width : maximum width;
metatibia length : from base to apex;
abdomen length : in lateral view, from base to apex;
abdomen width : in dorsal view, maximum width.
All measurements are in millimeters and are expressed as: minimum -(average)- maximum.
Family Coreidae
Subfamily Coreinae
Tribe Acanthocephalini
1870 Acanthocephala (Spilopleura)
Stål, 150 [n. subgen.; type species: Metapodius parensis
1938 Acanthocephala (Spilopleura): Blöte, 275.
2010 Acanthocephala (Spilopleura): Packauskas, 13 [synonymized under Acanthocephala].
2021 Acanthocephala (Spilopleura): CoreoideaSF Team [as synonym of Acanthocephala].
Spilopleura parensis (Dallas).
Pronotum without tubercles, posterior pronotal lobe always punctate; apex of scutellum flat, never incrassated or with an apparent callosity; with a conspicuously rounded or conical, but never spinose, projection on the metathoracic acetabulum; and metatibia conspicuously expanded.
Male. Head. Dorsally flat, lateral margins parallel (Fig.
Female. Structure and color similar to male. Thorax. Supracoxal area of metapleura not incrassate posteriorly. Metathoracic acetabulum without a conspicuous projection. Metatrochanter unarmed or with only one small ventral setiferous tubercle. Ventral margin of profemur with only one row of spines. Metafemur flattened and less developed than in males; spines of the dorsal margin without a conspicuous larger spine; ventral margin with two rows of flat spines that become larger to the apex and ends in a flat bidentate projection; posterior margin tuberculate or not. Expansions of the metatibia unarmed and more developed than in males. Abdomen. Expansion of abdominal sternite III absent. Female genitalia: Sternum VII with plica and fissure present. First gonocoxae large and triangular, distal angle rounded, proximal region of the outer lateral margin with a finger-like process for insertion of the muscles of the vagina (dorsal apodeme). First gonapophysis elongate, digitiform and hollow, basal and middle thirds punctate and with oblique striations and long setae, inner region of the distal third weakly sclerotized and with short and thick peg-like setae; first ramus straight and well sclerotized. Second gonocoxa elongate. Second gonapophysis elongate and well sclerotized, thickened distal region with a wide lateral tooth; second ramus well sclerotized, curved in basal region. Spermatheca: Type III, subtype D (
1852 Metapodius parensis Dallas, 432 [n. sp., Brazil].
1859 Metapodius parensis: Dohrn, 26.
1864 Metapodius parensis: Costa, 76.
1870 Acanthocephala (Spilopleura) parensis: Stål, 9: 150 [n. comb.].
1871 Metapodius parensis: Walker, 50.
1873 Metapodius parensis: Walker, 33.
1894 Acanthocephala parensis: Lethierry and Severin, 31.
1938 Acanthocephala (Spilopleura) parensis: Blöte, 275.
1938 Acanthocephala
(Spilopleura) parensis var. tristis Blöte, 20: 275 [n. var., Peru; sin. by
2010 Acanthocephala parensis: Packauskas, 19.
2021 Acanthocephala parensis: CoreoideaSF Team.
Lectotype
(here designated) ♀ (photograph), Metapodius parensis Dallas, type, syntype, BMNH(E) #651541 (
(Table
Distribution and summary of material examined of the species of Spilopleura stat. nov. Abbreviations: ARG – Argentina, BOL – Bolivia, BR – Brazil, COL – Colombia, PAR – Paraguay, PE – Peru.
Species | Type locality | Distribution | Comments | Material examined |
---|---|---|---|---|
S. parensis (Dallas, 1852) comb. nov. | Pará (BR) | ARG, BOL, BR, COL, PAR, PE | Type species of Spilopleura stat. nov. | 41 males, 37 females |
S. ochracea (Montandon, 1895) comb. nov. | Cumbase (BR) | BR, PAR, PE | 3 males, 2 females |
Species dark brown to black, with the apex of scutellum, apex of metatibia and metatarsus yellowish. Distiflagellomere bicolorous, dark brown with a yellow ring on apical third near the apex. Veins of clavus and embolium contrasting reddish. Abdominal tergites black, with a central yellowish line extending along all segments. Eyes with its major axis perpendicular to the dorsal margin of head in lateral view. Thoracic pleurae with three conspicuous patches of decumbent yellowish setae. Metathoracic acetabulum in males with a rounded projection. Metatibia with fusiform expansions. Posteroventral margin of pygophore with a small median depression. Expansion of the spermathecal duct small and oval.
Male. General color dark brown to black. Head. With sparse erect yellowish setae; dorsal region black with two paler areas next to ocelli (Fig.
Spilopleura parensis (Dallas). Male genitalia. A–C Pygophore: A Ventral view: B Dorsal view; C Lateral view; D Right paramere in both views; E, F Aedeagus: E Ventral view; F Dorsal view. Abbreviations: antO: pygophore anterior opening; Ar: paramere arm; bSh: paramere basal shank; Cj: Aedeagus conjunctiva; dApp1: dorsal conjunctival appendages I; Ds: dorsal sac of conjunctiva; ejD: ejaculatory duct; ejR: ejaculatory reservoir; End: endosoma; gp: gonopore; iMbSh: inner margin of paramere basal shank; laM: pygophore lateral margin; oMbSh: outer margin of paramere basal shank; Pha: phallosoma; postO: pygophore posterior opening; pvM: pygophore posteroventral margin; vApp: ventral conjunctival appendages; Vs: vesica.
Variation (observed in some specimens examined): Apex of tylus slightly projected upwards.
Female. Thorax. Humeral distance as wide as the width of abdomen. Posterior row of spines on ventral margin of pro- and mesofemora absent. Metafemur at least two times wider than profemur (Fig.
Spilopleura parensis (Dallas). Female genitalia. A Ventral view of the pregenital region and genital plates; B First gonocoxa and first gonapophysis; C First gonapophysis; D Second gonocoxa and second gonapophysis; E Spermatheca. Abbreviations: dA: dorsal apodem; dd: distal duct; df: distal flange; di: dilatation of the sphermatecal duct; fi: fisura; fz: flexible zone; Gp8: first gonapophysis; Gp9: Second gonapophysis; Gx8: first gonocoxa; Gx9: second gonocoxa; pd: proximal duct; pf: proximal flange; pgS: peg-like setae; pl: plica; Pt8: paratergite VIII; Pt9: paratergite IX; Ra1: first ramus; Ra2: second ramus; sr: seminal receptacle; VII: Sternite VII.
(n= 5♂ | 5♀): Total length: ♂ 18 -(20.727)- 22.625 / ♀ 18 -(20.37530)- 23.74; head length: ♂ 1.992 -(2.054)- 2.112 / ♀ 1.944 -(2.097)- 2.136; max head width: ♂ 2.432 -(2.645)- 2.816 / ♀ 2.432 -(2.609)- 2.72 tylus length: ♂ 0.736 -(0.837)- 0.992 / ♀ 0.64 -(0.789)- 0.96; post-tylus depression length: ♂ 0.288 -(0.313)- 0.336 / ♀ 0.24 -(0.256)- 0.272; eye width: ♂ 0.688 -(0.726)- 0.752 / ♀ 0.72 -(0.771)- 0.832; interocular space: ♂ 1.088 -(1.22)- 1.376 / ♀ 1.056 -(1.194)- 1.312; ocellar distance: ♂ 0.896 -(1.06)- 1.216 / ♀ 0.928 -(1.036)- 1.12; interocellar space: ♂ 0.576 -(0.72)- 0.8 / ♀ 0.608 -(0.707)- 0.832; antennal articles length: scape, ♂ 4 -(4.582)- 5 / ♀ 3.635 -(4.185)- 5; pedicel, ♂ 3.5 -(3.816)- 4.25 / ♀ 2.875 -(3.511)- 4.25; basiflagellomere, ♂ 2.875 -(3.419)- 3.75 / ♀ 2.875 -(3.221)- 3.75; distiflagellomere, ♂ 7.25 -(8.1)- 9.125 / ♀ 6.75 -(7.432)- 8.125; antennal articles width: scape, ♂ 0.32 -(0.355)- 0.384 / ♀ 0.256 -(3.18)- 0.352; pedicel, ♂ 0.192 -(0.358)- 0.256 / ♀ 0.192 -(0.217)- 0.256; basiflagellomere, ♂ 0.192 -(0.226)- 0.256 / ♀ 0.192 -(0.211)- 0.224; distiflagellomere, ♂ 0.228 -(0.25)- 0.256 / ♀ 0.224 -(0.288)- 0.352; labial segments length: I, ♂ 1.692 -(1.875)- 2.021 / ♀ 1.115 -(1.854)- 1.739; II, ♂ 1.551 -(1.725)- 1.88 / 1.504-(1.684)- 1.88; III, ♂ 1.363 -(1.493)- 1.696 / ♀ 1.363 -(1.485)- 1.786; IV, ♂ 1.363 -(1.433)- 1.568 / ♀ 1.269 -(1.365)- 1.457; humeral angles width: ♂ 6.65 -(8.03)- 8.55 / ♀ 7.125 -(7.983)- 8.75; scutellum length: ♂ 2.538 -(2.877)- 3.29 / ♀ 2.585 -(2.873)- 3.243; scutellum width: ♂ 2.538 -(2.925)- 3.29 / ♀ 2.632 -(2.978)- 3.29; hemelytra length: ♂ 15.625 -(17.375)- 18.75 / ♀ 15.125 -(17.139)- 18.75; profemur width: ♂ 0.768 -(0.828)- 0.912 / ♀ 0.72 -(0.777)- 0.832 ; metafemur width: ♂ 1.76 -(2.558)- 3.2 / ♀ 1.6 -(1.75)- 1.952; metafemur length: ♂ 10.75 -(12.276)- 13.75 (12.276)/ ♀ 10.625 -(10.833)- 11.5; metatibia width: ♂ 1.52 -(1.945)- 2.24 / ♀ 2.304 -(2.733)- 3.104; metatibia length: ♂ 10.875 -(12.152)- 13.75 / ♀ 9.626 -(11.152)- 11.75; abdomen length: ♂ 9.75 -(10.983)- 12.5 / ♀ 9.625 -(11.041)- 13.5; abdomen width: ♂ 5.5 -(6.5)- 7.125 / ♀ 6.125 -(7.272)- 8.15.
Bolivia, Brazil, Peru, and first records from Argentina, Colombia, and Paraguay (Fig.
A single female specimen with a label “syntype” deposited in the Natural History Museum of London (detailed in material examined and examined through photographs) is the only type material of Metapodius parensis Dallas we found.
To ensure nomenclatural stability, we select this specimen (Fig.
The analysis of the characters exhibited in the photograph examined of the male paratype of A. (S.) parensis var. tristis Blöte allows us to confirm the synonymy under Spilopleura parensis as proposed by Packauskas.
1895 Acanthocephala ochracea Montandon, 8, pl. 1, fig. 3 [n. sp., Brazil, illustration].
1913 Acanthocephala ochracea: Bergroth, 129.
2010 Acanthocephala ochracea: Packauskas, 18.
2021 Acanthocephala ochracea: CoreoideaSF Team.
Holotype
♀ (photograph), Acanthocephala (Metapodius) ochracea Montandon, Bresíl du Nord, Cumbase, type, holotype, Romania (
(Table
Species entirely yellowish ochraceous, with the anterior region of the metaepisternum darker, and with two dark brown central lines extending along all abdominal tergites. Eyes with its major axis oblique to the dorsal margin of head in lateral view. Thoracic pleurae with sparse setae, never forming a patch. Metatibia with lanceolate expansions. Posteroventral margin of pygophore entire, without a median depression. Expansion of the spermathecal duct tubular, long and curved.
Male. General color yellowish ochraceous. Head. With sparse erect and semierect yellowish setae (Fig.
Spilopleura ochracea (Montandon). Male genitalia. A–C Pygophore: A Ventral view; B Dorsal view; C Lateral view; D Right paramere in both views; E–F Aedeagus: E Ventral view; F Dorsal view. Abbreviations: antO: pygophore anterior opening; Ar: paramere arm; bSh: paramere basal shank; Cj: Aedeagus conjunctiva; dApp1: dorsal conjunctival appendages I; Ds: dorsal sac of conjunctiva; ejD: ejaculatory duct; ejR: ejaculatory reservoir; End: endosoma; gp: gonopore; iMbSh: inner margin of paramere basal shank; laM: pygophore lateral margin; oMbSh: outer margin of paramere basal shank; Pha: phallosoma; postO: pygophore posterior opening; pvM: pygophore posteroventral margin; vApp: ventral conjunctival appendages; Vs: vesica.
Variation (observed in one specimen examined): Anterior margin of metaepisternum entirely ochraceous, without a darker area.
Female. Thorax. Humeral distance 1.2 times wider than the abdomen. Posterior row of spines on ventral margin of profemur absent. Metafemur ochraceous, at least 2.4 times wider than profemur (Fig.
Spilopleura ochracea (Montandon). Female genitalia. A Ventral view of the pregenital region and genital plates; B First gonocoxa and first gonapophysis; C First gonapophysis; D Second gonocoxa and second gonapophysis; E Spermatheca. Abbreviations: dA: dorsal apodem; dd: distal duct; df: distal flange; di: dilatation of the sphermatecal duct; fi: fisura; fz: flexible zone; Gp8: first gonapophysis; Gp9: Second gonapophysis; Gx8: first gonocoxa; Gx9: second gonocoxa; pd: proximal duct; pf: proximal flange; pgS: peg-like setae; pl: plica; Pt8: paratergite VIII; Pt9: paratergite IX; Ra1: first ramus; Ra2: second ramus; sr: seminal receptacle; VII: Sternite VII.
(n= 2♂ | 1♀): Total length: ♂ 24.375–24.625 / ♀ 21; head length: ♂ 2.72–2.88 / ♀ 2.496; max head width: ♂ 2.688–2.848 / ♀ 2.528 ; tylus length: ♂ 1.12–1.224 / ♀ 1.12; post-tylus depression length: ♂ 0.24 / ♀ 0.264; eye width: ♂ 0.672–0.72 / ♀ 0.624; interocular space: ♂ 1.32–1.44 / ♀ 1.248; ocellar distance: ♂ 1.024 / ♀ 0.896; interocellar space: ♂ 0.704 / ♀ 0.444; antennal articles length: scape, ♂ 4.625–5.32 / ♀ 3.807; pedicel, ♂ 4.5 / ♀ 3.76; basiflagellomere, ♂ 4.25 / ♀ 3.525; distiflagellomere, ♂ 9.875 / ♀ 5.605; antennal articles width: scape, ♂ 0.416–0.512 / ♀ 0.384; pedicel, ♂ 0.288 / ♀ 0.224; basiflagellomere, ♂ 0.288 / ♀ 0.224; distiflagellomere, ♂ 0.288 / ♀ 0.288; length of labial segments: I, ♂ 2.115–2.176 / ♀ 1.88; II, ♂ 1.88–2.08 / ♀ 1.88; III, ♂ 2.528–2.585 / ♀ 2.538; IV, ♂ 1.739–1.92 / ♀ 1.692; humeral angles width: ♂ 7.747–8.93 / ♀ 7.5; scutellum length: ♂ 3.232–3.296 / ♀ 2.82; scutellum width: ♂ 2.88 / ♀ 2.632; hemelytra length: ♂ 18–18.125 / ♀ 16.625; profemur width: ♂ 1.056 / ♀ 0.658; metafemur width: ♂ 2.656–2.848 / ♀ 1.598; metafemur length: ♂ 12.625–13.125 / ♀ 10.625; metatibia width: ♂ 3.04–3.2 / ♀ 2.867; metatibia length: ♂ 12.125–13 / ♀ 11.25; abdomen length: ♂ 11.875–12.5 / ♀ 11.25; abdomen width: ♂ 6.625–6.935 / ♀ 6.125.
Stål (1870) established the subgenera mainly based on the shape of the metatibial expansions, the coloration of the abdominal tergites, and the presence of abdominal expansions in segment three. In Spilopleura the metatibial expansions are fusiform or lanceolate, and are developed along all the tibial length with the apex narrow in both sexes, contrary to Acanthocephala where the metatibial expansions are fusiform only in males, or are well developed along all the tibia in both sexes but ends with the apex rounded or straight. In the case of the abdominal characters, the third abdominal sternite is not expanded laterally like in some Acanthocephala which present sub-triangular expansions; and the abdominal terga present contrasting longitudinal central lines (one or two) that extend along all abdominal segments, character absent in Acanthocephala.
In addition to the characters observed by Stål (1870), we were able to recognize a set of generic character states shared with other Acanthocephalini genera, but that allows to separate Spilopleura from Acanthocephala, such as: the apex of scutellum flat, the pronotal disc never tuberculate, and the metathoracic acetabulum in males with a conspicuous rounded or conical projection. On the other hand, in Acanthocephala, the apex of scutellum is swollen; the pronotal disc is always tuberculate; and the metathoracic acetabulum in males lacks a conspicuous projection.
Both genera share, but with different degrees of development, a conspicuously expanded metatibia, a character state also exhibited by Cervantistellus, Empedocles, Ichilocoris, Lucullia, Meluchopetalops, Stenometapodus, Thymetus and Zygometapodus. Among these, Ichilocoris, Cervantistellus, Lucullia, Meluchopetalops, Stenametapodus, Zygometapodus, and Spilopleura have the pronotal disc not tuberculate, unlike Empedocles, Thymetus, and Acanthocephala, which present the posterior pronotal lobe with tubercles of different degrees of development. On the other hand, Lucullia, Zygometapodus and Spilopleura present a conspicuous conical or rounded, never spine-like, projection in the metathoracic acetabulum in males. Lucullia and Spilopleura also have the apex of scutellum flat, never swollen as in the other genera. However, Lucullia species can be easy recognized and separated from Spilopleura by the metallic green iridescence coloration, the presence of a wide yellow longitudinal stripe extending across pronotal disk to apex of scutellum; and by the fusiform and slightly expanded metatibial expansions in both sexes.
This work was funded by the Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Argentina; and the Universidad Nacional de La Plata. We would like to thank John M. Leavengood for the critical reading of the manuscript.
Spilopleura Stål (status novum) distribution
Data type: occurences
Explanation note: Spilopleura Stål (status novum) distribution.