Research Article |
Corresponding author: Matthias Seidel ( matth.seidel@gmail.com ) Academic editor: Andrey Frolov
© 2022 Matheus Bento, Mary Liz Jameson, Matthias Seidel.
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:
Bento M, Jameson ML, Seidel M (2022) New species and illustrated key of Macraspis (Scarabaeidae, Rutelinae, Rutelini) from the Amazon biome of Brazil. ZooKeys 1124: 161-189. https://doi.org/10.3897/zookeys.1124.91156
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The phytophagous scarab genus Macraspis MacLeay (Scarabaeidae, Rutelinae, Rutelini) is reviewed from the Brazilian Amazon region. Three new species are described and illustrated from the states of Amazonas, Pará, and Rondônia: M. buehrnheimi sp. nov., M. opala sp. nov., and M. phallocardia sp. nov. Two species, Macraspis fernandezi Neita-Moreno and M. oblonga Burmeister, are recorded for the first time in Brazil (new country records). Macraspis maculata crosarai Soula is a new synonym of Macraspis maculata Burmeister; hence this species no longer includes subspecies. Furthermore, Macraspis cincta parensis Soula, 2005 is deemed unavailable under the provisions of ICZN Articles 16.4.1 and 16.4.2. An illustrated key to 15 species and subspecies of Macraspis from the Brazilian Amazon enables identification of this speciose leaf chafer genus.
Chafers, identification, morphology, Neotropical Region, taxonomy
The Neotropical genus Macraspis MacLeay, 1819 (Scarabaeidae, Rutelinae, Rutelini) is a distinctive and widely distributed leaf chafer group that occurs from Mexico to Argentina (
Macraspis spp. (A–O). Lateral and ventral views of pro- and mesothorax of A, B M. morio and D, E M. p. pseudochrysis. Dorsal view of head and anterior portion of pronotum of C M. festiva and F M. c. chloraspis. Anterior view of mesothoracic legs of G M. festiva H M. oblonga I M. p. pseudochrysis and J M. c. chloraspis. Female metatibial spurs of K M. c. chloraspis and L M. phallocardia sp. nov. Anterior view of mesotarsus of M M. p. pseudochrysis N M. c. chloraspis and O M. fernandezi. Scale bars: 1 mm.
Macraspis spp. (A–O). Dorsal view of head of A M. oblonga B M. olivieri and C M. xanthosticta. Lateral view of mesothorax of D M. festiva E M. maculata and F M. c. chloraspis. Female pygidium of G M. oblonga H M. olivieri and I M. morio. Dorsal view of female abdomen of J M. p. pseudochrysis and K M. lepiouffi paratype. Metatarsus of L M. morio and O M. olivieri. Female last abdominal ventrites of M M. morio and N M. p. pseudochrysis. Scale bars: 0.5 mm (A–C), 1 mm (D–O).
Macraspis buehrnheimi sp. nov. (A–G) and Macraspis fernandezi Neita-Moreno, 2014 (H–J) Macraspis buehrnheimi sp. nov. A dorsal view of holotype male and B ventral view of holotype male C dorsal view paratype female and D ventral view view paratype female E–F aedeagus of holotype (E dorsal F caudal G lateral views). Macraspis fernandezi Neita-Moreno, 2014 from Caracaraí, Roraima state, Brazil (new country record) H aedeagus dorsal view I aedeagus caudal view and J aedeagus lateral view. Scale bars: 2 mm (A–D); 0.5 mm (E, G, H, J); 0.2 mm (F, I).
We examined the Macraspis species and subspecies with distributions in the Brazilian Amazon biome (defined as northern Brazil including the states of Acre, Amapá, Amazonas, Pará, Rondônia, Roraima, as well as small portions of Mato Grosso, Tocantins, and Maranhão states) (IBGE 2004) (Fig.
Macraspis currently includes 68 species and 21 subspecies (
The type material and additional specimens used for descriptions, comparisons, and key are deposited in the following institutions (acronym and curators parenthesized): Seção de Entomologia da Coleção Zoológica da Universidade Federal do Mato Grosso, Cuiabá, Brazil (CEMT; Fernando Zagury Vaz-de-Mello); Coleção Entomológica do Instituto Oswaldo Cruz, Rio de Janeiro, Brazil (CEIOC; Márcio Felix, Claudia Leal Rodrigues); Musée des Confluences, Lyon, France (CCECL; Cédric Audibert); Coleção Zoológica Prof. Paulo Bührnheim, Universidade Federal do Amazonas, Manaus, Brazil (CZPB; Fábio Siqueira Pitaluga de Godoi); Coleção Entomológica Pe. Jesus Santiago Moure, Universidade Federal do Paraná, Curitiba, Brazil (
Body length was measured from the apex of the clypeus to the apex of the elytra; body width was measured across the humeri. Regarding punctures, surfaces were considered punctostriate when punctures were confluent and elongated, densely punctate if punctures were nearly confluent to less than two puncture diameters apart, moderately punctate if 2–6 punctures diameters apart, and sparsely punctate if punctures were separated by more than six puncture diameters. The punctures were considered small when less than 0.019 mm in diameter, moderate when 0.02–0.049 mm, and large if larger than 0.05 mm. Setae were defined short if between 0.1–0.19 mm in length, moderately long if between 0.2–0.39 mm, and long if longer than 0.4 mm. Regarding density of setae, surfaces were considered densely setose when many setae completely covered the surface, moderately setose when the surface visible and with many setae, and sparsely setose when the surface was visible and with only a few setae.
Morphological terms follow
Like many other Macraspis species, identification of the two new species described here requires examination of male genitalia. Female external genitalia of these new species showed conspicuous differences in the proximal and distal gonocoxites. However, caution must be taken concerning the identification of females based only on external genital characters because the whole range of intraspecific variation is not known for these structures. Accordingly, females of these species can be more reliably identified when collected with associated males. Therefore, females were not included in the diagnoses and key.
For purposes of the identification key, we include species known to occur only within the Brazilian Amazon biome (
Images were taken using a Leica DFC295 camera attached to a Leica M165C stereomicroscope and were processed using the Leica Application Suite (LAS) v. 4.1 and Helicon Focus (HeliconSoft) software. The photographic illumination follows
The verbatim label data from type specimens are transcribed in quotation marks, with “/” used to separate lines on the same label, and “//” to separate labels. Label data from non-type specimens are provided as follows: country, state or province, locality, date, collector or old collection (quantity, sex symbol, collection acronym).
The geographic coordinates were obtained with Google Maps and the georeferenced points were plotted on the distribution map generated by the web software Simplemappr (
Based on examination of specimens in 12 collections, we recorded seven species as occurring in the Brazilian Amazon: M. buehrnheimi sp. nov., M. fernandezi Neita-Moreno, M. lateralis (Olivier), M. martinezi auzerali Soula, 1998, M. oblonga Burmeister, M. opala sp. nov., and M. phallocardia sp. nov. We provide species treatments and a distribution map for all species including M. maculata Burmeister, 1844 (Fig.
Macraspis cincta parensis Soula, 2005 was described based on two specimens with the doubtful locality of Obidos, Pará state, Brazil. In the original description
Lacking females of M. buehrnheimi sp. nov., M. fernandezi, M. lateralis, M. opala sp. nov., M. phallocardia sp. nov.
1 | Protarsomere V enlarged; anterior protarsal claw thickened and unequally cleft; pygidium strongly convex, with posterior margin ventrally positioned, male | 2 |
– | Protarsomere V not enlarged; anterior protarsal claw not thickened and equally cleft; pygidium weakly convex to plano-convex, with posterior margin dorsoapically positioned, female | 16 |
2 | Mesometaventral process thickened and oblique in relation to the longitudinal axis of the body (Fig. |
M. morio Burmeister |
– | Mesometaventral process weakly flattened and parallel in relation to the longitudinal axis of the body (Fig. |
3 |
3 | Large body size (length 14–23 mm); head without supraorbital strigae (Fig. |
4 |
– | Small body size (length 7–14 mm); head with supraorbital strigae (Fig. |
9 |
4 | Mesotibia medially wider than half the mesofemur width (Fig. |
5 |
– | Mesotibia medially narrower than half the mesofemur width (Fig. |
6 |
5 | Body non-metallic green; posterior border of mesocoxal cavity narrow (Fig. |
M. pseudochrysis pseudochrysis Landin |
– | Body metallic green; posterior border of mesocoxal cavity wide (Fig. |
M. lepiouffi Soula |
6 | Dorsum unicolored, with elytra uniformly green and without metallic maculation | M. peruviana Ohaus |
– | Dorsum bicolored, with elytra yellow with or without metallic green maculation | 7 |
7 | Pronotum entirely green (lateral margins same color as disc) | M. festiva Burmeister |
– | Pronotum with disc green and lateral margins yellow | 8 |
8 | Clypeus rugopunctate at apex, with anterior margin strongly raised and subtruncated (Fig. |
M. oblonga Burmeister |
– | Clypeus densely punctate at apex, with anterior margin weakly raised and rounded (Fig. |
M. olivieri (Waterhouse) |
9 | Mesotibia medially narrower than mesofemur at middle (Fig. |
M. chloraspis chloraspis Laporte |
– | Mesotibia medially as wide as mesofemur at middle (Fig. |
10 |
10 | Paramera apically dilated (Fig. |
11 |
– | Paramera apically narrowed (Figs |
12 |
11 | Lateral articular areas of tectum truncated and weakly projected distally (Fig. |
M. fernandezi Neita-Moreno |
– | Lateral articular areas of tectum pointed and strongly projected distally (Fig. |
M. buehrnheimi Bento, Jameson & Seidel, sp. nov. |
12 | Apex of pygidium smooth (Fig. |
M. opala Bento, Jameson & Seidel, sp. nov. |
– | Apex of pygidium with strong, concentric sculpturing (Fig. |
13 |
13 | Smaller specimens (length 7–9 mm); anterior margin of clypeus weakly notched medially (more evident in females) (Fig. |
M. xanthosticta Burmeister |
– | Larger specimens (length 10–11.7 mm); anterior margin of clypeus not notched; scutellar shield strongly constricted basolaterally; paramera with apex narrowly rounded or parabolic | 14 |
14 | Elytron metallic green with two median, yellow maculae or a single transverse yellow band | M. martinezi auzerali Soula |
– | Elytron usually metallic green and without yellow maculae or band | 15 |
15 | Lateral articular areas of tectum thickened and deflected outward (Fig. |
M. phallocardia Bento, Jameson, Seidel, sp. nov |
– | Lateral articular areas of tectum compressed and straight (Fig. |
M. lateralis (Olivier) |
16 | Mesometaventral process thickened and oblique to the body’s longitudinal axis (Fig. |
M. morio Burmeister |
– | Mesometaventral process weakly flattened and parallel to body longitudinal axis (Fig. |
17 |
17 | Large specimens (length 14–23 mm); head without supraorbital strigae (Fig. |
18 |
– | Small specimens (length 7–14 mm); head with supraorbital strigae (Fig. |
23 |
18 | Mesotibia medially wider than half the mesofemur width (Fig. |
19 |
– | Mesotibia medially narrower than half the mesofemur width (Fig. |
20 |
19 | Body non-metallic green; pygidium apically obtuse in dorsal view (Fig. |
M. pseudochrysis pseudochrysis Landin |
– | Body metallic green; pygidium apically acuminate in dorsal view (Fig. |
M. lepiouffi Soula |
20 | Dorsum unicolored, with elytra uniformly green and without metallic maculation | M. peruviana Ohaus |
– | Dorsum bicolored, with elytra yellow with or without metallic green maculation | 21 |
21 | Pronotum entirely green (lateral margins same color as disc) | M. festiva Burmeister |
– | Pronotum with disc green and lateral margins yellow | 22 |
22 | Clypeus rugopunctate at apex, with anterior margin strongly raised and subtruncated (Fig. |
M. oblonga Burmeister |
– | Clypeus densely punctate at apex, with anterior margin weakly raised and rounded (Fig. |
M. olivieri (Waterhouse) |
23 | Inner metatibial spur evenly and strongly curved (Fig. |
M. chloraspis chloraspis Laporte |
– | Inner metatibial spur straight or barely curved (Fig. |
24 |
24 | Smaller specimens (length 7–9 mm); anterior margin of clypeus weakly notched medially (Fig. |
M. xanthosticta Burmeister |
– | Larger specimens (length 10–11.7 mm); anterior margin of clypeus not notched; scutellar shield more strongly constricted basolaterally | M. martinezi auzerali Soula |
(1 male, 1 female). Holotype male deposited at CZPB, labeled: “BRASIL, Amazonas, Coari, / rio Urucu, LUC – 09, 4°51'56"S, 65°04'56"W, / 25/I–10/II/1995, P. F. / Bührnheim et al col.” (white, printed) // “à luz mista / de mercúrio” (white, printed) // “HOLOTYPE / Macraspis buehrn-/ heimi Bento, Jameson, / Seidel, 2022 / M. Bento, det. 2022. Paratype: same data as holotype (1 ♀, CZPB).
Male genitalia are required for identification: lateral articular areas of tectum pointed and strongly projected distally (Fig.
Holotype male (Fig.
Comparison of male endophalli of Macraspis buehrnheimi sp. nov. and Macraspis fernandezi Neita-Moreno, 2014. Macraspis buehrnheimi sp. nov. (A–D) in A dorsal B lateral and D lateral views, with C detail showing lateral sclerite. Macraspis fernandezi Neita-Moreno, 2014 (E–H) in E dorsal F lateral and H ventral views, with G detail showing lateral sclerite. Scale bars: 0.5 mm.
Paratype (1 female) (Fig.
Comparison of female external genitalia of Macraspis buehrnheimi sp. nov. (A–C) and Macraspis phallocardia sp. nov (D–G). Macraspis buehrnheimi sp. nov. in A ventral B lateral and C dorsal views. Macraspis phallocardia sp. nov. in D ventral E lateral and F dorsal views. Abbreviations: dgx = distal gonocoxite; pgx = proximal gonocoxite; pct = proctiger; ppt = paraproct. Scale bars: 0.3 mm.
This species is named after the Brazilian zoologist Paulo Friederich Bührnheim (1937–2001), who greatly contributed to education and research in Amazonas state, Brazil. In addition, he founded the insect collection at the Universidade Federal do Amazonas (UFAM) and collected the type series of this species.
(Fig.
Macraspis buehrnheimi sp. nov. has the same color pattern as M. lateralis (Olivier, 1789), M. fernandezi Neita-Moreno, 2014, and M. phallocardia sp. nov. These species are only separated by careful comparison of male genitalia. The male aedeagus of M. buehrnheimi is most similar to that of M. fernandezi (unknown female) in that both have paramera apically dilated. Other characters that serve to separate M. buehrnheimi and M. fernandezi are (characters of M. fernandezi given in parenthesis): tectum abruptly narrowed towards the apical edge, with lateral articular areas pointed and strongly projected distally (tectum evenly narrowed towards the apical edge, with lateral articular areas truncated and weakly projected distally (Fig.
There are no reliable means to distinguish the female of M. buehrnheimi sp. nov. from that of M. phallocardia sp. nov. based on external morphology. Analysis of the external genitalia showed conspicuous differences in the proximal and distal portions of the gonocoxites of these species (compare Fig.
Holotype
male deposited at
(2 males). Brazil, Roraima, Caracaraí, Parque Nacional da Serra da Mocidade, 15–26.I.2016, F. F. Xavier, R. Boldrini, & P. Barroso (legs.) (♂,
(Fig.
This species was described based on a single male specimen from Colombia (
Cetonia lateralis Olivier, 1789: 80.
Cetonia virens
Fabricius, 1801: 141. Junior subjective synonym by
Macraspis lateralis var. cincticollis
Ohaus, 1898: 52. Junior subjective synonym by
Macraspis lateralis var. immaculata
Ohaus, 1898: 52. Junior subjective synonym by
Neotype
male (Fig.
Neotype male of Cetonia lateralis Olivier, 1789 (=Macraspis lateralis (Olivier)): A dorsal view B labels C aedeagus in frontal view D aedeagus in lateral view. A, B from
(2 males, 2 females). Brazil, Amazonas, Manaus, Reserva Ducke, 21.II.1978, Jorge Arias (leg.) (♀,
(Fig.
Macraspis maculata
Burmeister, 1844: 351 (original description);
Macraspis maculata crosarai
Soula, 1998: 38 (original description);
(2 males, 1 female). Lectotype male of Macraspis maculata (designated by Soula, 1998) deposited at
(18 males, 13 females). Brazil, Bahia, Prado, 05.III.1971, C. Elias (leg.) (♀,
(Fig.
Macraspis maculata has been recorded from the Brazilian Amazon (
Macraspis oblonga
Burmeister, 1844: 359 (original description);
(4 females). Brazil, Amazonas, Manaus, Fazenda Esteio, 30.VIII.1984, R. C. Klein (leg.) (♀,
Suriname (
This species is sympatric with M. olivieri, and both have a very similar color pattern. However, M. oblonga is easily distinguished from M. olivieri by (characters of M. olivieri given in parenthesis): clypeal apex rugopunctate, with anterior margin strongly raised and subtruncated (Fig.
(2 males). Holotype male deposited at
Apex of pygidium smooth (Fig.
Macraspis opala sp. nov. (A–H) and Macraspis maculata Burmeister (I–M): Macraspis opala sp. nov., holotype male A dorsal view B ventral view C pygidium in caudal view D spiculum gastrale E aedeagus, dorsal view F aedeagus, caudal view G lateral view, paramera (of paratype) and H caudal view. Macraspis maculata Burmeister, 1844 from Linhares, Espírito Santo state, Brazil, I spiculum gastrale J aedeagus in dorsal view K caudal view and L lateral view M pygidium in caudal view. Scale bars: 2 mm (A, B); 1 mm (C, M); 0.5 mm (E, G, J, L); 0.2 mm (F, H, K).
Holotype male (Fig.
Female. Unknown.
Paratype (1 male). The male paratype differs from holotype by the general coloration darker with stronger green reflections and elytra without yellow maculae; paramera slightly narrower, with lateral margins parallel and apex slightly more rounded (Fig.
The specific epithet is derived from the Latin word ‘opalus’ (= precious stone) in reference to opal gemstone, alluding to the metallic, multicolor cuticular surface.
(Fig.
This species is quite similar to Macraspis maculata Burmeister, 1844. However, M. opala sp. nov. has a slightly smaller body size and is distinguished from M. maculata by (characters of M. maculata given in parenthesis): apex of pygidium smooth, without sculpturing (apex of pygidium with strong and concentric sculpturing (Fig.
(4 males, 14 females). Holotype male deposited at
Male: lateral articular areas of tectum thickened and deflected outward (Fig.
Macraspis phallocardia sp. nov. (A–G) and Macraspis lateralis (Olivier, 1789) (H–J). Macraspis phallocardia sp. nov. A holotype male dorsal view B holotype male ventral view C paratype female dorsal view D paratype female ventral view E–G aedeagus of holotype in E dorsal view F caudal view G lateral view. Macraspis lateralis (Olivier, 1789) from Manaus, Amazonas state, Brazil H aedeagus in dorsal view I caudal view J lateral view. Scale bars: 2 mm (A–D); 0.5 mm (E, G, H, J); 0.2 mm (F, I).
Male holotype (Fig.
Comparison of male endophalli of Macraspis lateralis (Olivier, 1789) (A–D) and Macraspis phallocardia sp. nov. (E–H): Macraspis lateralis (Olivier, 1789) in A dorsal view B lateral view D lateral view and C showing detail of lateral sclerite. Macraspis phallocardia sp. nov. in E dorsal view F lateral view H ventral view and G showing detail of lateral sclerite. Scale bars: 0.5 mm (A, B, D, E, F, H); 0.2 mm (C, G).
Paratypes (3 males, 14 females): male paratypes differ from holotype in length (10.7–11.7 mm), width (5.8–6.7 mm), and form of the apex of paramera (more round to slightly narrowed). Female paratypes (Fig.
The specific epithet is Greek for ‘phallos’ (= penis) and ‘kardia’ (= heart), refers to the heart-shaped male paramera of this species.
(Fig.
Distribution of Brazilian species of Macraspis treated in this publication. The Brazilian Amazon biome is colored lime green. Brazilian states are abbreviated as follows: AC = Acre, AM = Amazonas, AP = Amapá, BA = Bahia, ES = Espírito Santo, GO = Goiás, MA = Maranhão, MG = Minas Gerais, MT = Mato Grosso = PA = Piauí, RO = Rondônia, RJ = Rio de Janiero, RR = Roraima, SP = São Paulo, TO = Tocantins.
This species has the same color pattern as M. buehrnheimi sp. nov., M. fernandezi, and M. lateralis. The male genitalia of M. phallocardia sp. nov. is more similar to that of M. lateralis, but these species are differentiated by (characters of M. lateralis given in parenthesis): tectum with lateral articular areas thickened and deflected outward (tectum with lateral articular areas compressed and straight; Fig.
We are grateful to all of the curators mentioned in Material and methods, especially Antoine Mantilleri (