Research Article |
Corresponding author: Daercio A.A. Lucena ( daerciobio@gmail.com ) Academic editor: Michael Ohl
© 2021 Daercio A.A. Lucena, Eduardo A.B. Almeida, Fernando C.V. Zanella.
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:
Lucena DAA, Almeida EAB, Zanella FCV (2021) Amiseginae and Cleptinae from northeastern Brazil, with the description of four new species (Hymenoptera, Chrysididae). Journal of Hymenoptera Research 81: 57-85. https://doi.org/10.3897/jhr.81.60048
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The diversity of two subfamilies of cuckoo wasps in northeastern Brazil is reviewed. Four new species are described and illustrated: Amisega boyi Lucena, sp. nov., A. sertaneja Lucena, sp. nov., and Duckeia dudui Lucena, sp. nov. (Amiseginae), and Cleptidea nordestina Lucena, sp. nov. (Cleptinae). These new species of Amisega and Duckeia represent the first records of both genera in the core zone of the Caatinga dry region, and they seem to be endemic to this portion of the region. The other two Amisega species previously recorded in northeastern Brazil are restricted to ecotone habitats between Caatinga and Atlantic forest. Cleptidea nordestina Lucena, sp. nov. (Cleptinae) is the most septentrional record for the fasciata species group in South America. With the present contribution, the total recorded diversity of Amiseginae and Cleptinae in northeastern Brazil is represented by the following species: A. boyi Lucena, sp. nov., A. flavipes Kimsey, 1987, A. sertaneja Lucena, sp. nov., A. similis Kimsey, 1987, and D. dudui Lucena, sp. nov. (Amiseginae), and C. nordestina Lucena, sp. nov. (Cleptinae).
Chrysidoidea, deciduous forest, diversity, rain forest, semiarid, taxonomy
Cuckoo wasps are a species-rich cosmopolitan group of aculeate hymenopterans, currently with more than 2,500 valid species distributed into 94 extant genera and four subfamilies (e.g., taxonomic data modified from
Northeastern Brazil corresponds to an area roughly equivalent to Colombia and Ecuador together, totaling 1,552,157 Km2 and extending from 07°12' to 48°20'S, and the easternmost point in South America, nearly 34°47' to 48°45'W (
The driest biome, Caatinga, has traditionally been considered to harbor the poorest biodiversity in comparison with more humid neighboring biomes, which also reflected in a relative overlook of its biology. Despite that, this biome has received growing attention from researchers (e.g.,
Caatinga is mostly characterized by high temperatures throughout the year, with a marked seasonal climate marked by a strong hydric deficit (Fig.
Easternmost biogeographic regions of South America (modified from
Herein, we expand the knowledge of Chrysididae fauna in northeastern Brazil by describing and illustrating four new species in the Amiseginae and Cleptinae. We comment on the new species’ habitats and discuss some geographic records of previously known taxa from the northeastern region. Additionally, we discuss and update the list of taxa with occurrence in the core region of Caatinga.
For the analyses of geographic occurrence of species into the semiarid limits, we used delimitation presented by
Most specimens were collected using malaise traps standing in the field for at least two consecutive months (Patu and Mossoró-RN), a whole year (Jequié-BA), and four consecutive years (Santa Teresinha-PB) (detailed descriptions of localities and sampling methods were provided in
Specimens were photographed at Laboratório de Biologia Comparada e Abelhas, Departamento de Biologia, FFCLRP/USP, using a Leica DFC425 camera attached to a Leica M205C stereomicroscope, equipped with a Leica FlexiDome lighting diffuser, and light system model Leica LED5000 HDI. Images were improved using Leica application suite software-LAS to combine multiple photos, then enhanced with Adobe Photoshop.
The general morphological terminology follows primarily
The Amiseginae are specialized parasitoids of walking stick eggs (e.g.,
Amisega Cameron, 1888: 457. Type species: Amisega cuprifrons Cameron, 1888: 457 (by monotypy).
This genus is one of the largest in the Amiseginae. Currently, Amisega includes 24 valid species (
Vertex and anterolateral border of pronotum with rugose-striate marks; anterior declivity of pronotum without medial pit; dorsum of propodeum polished; scapal basin punctate-rugulose, vaguely cross-ridged; metasoma with distinct purple highlights; legs brown, lighter on pro- and mesotibiae, and tarsomeres; malar space 1.5× MOD. Amisega boyi Lucena, sp. nov. most resembles A. flavipes Kimsey, 1987 and A. similis Kimsey, 1987. This new species can be distinguished from both species based on the following combination of characters: rugose marks on vertex and pronotum (lacking rugose marks on vertex in A. similis, and only striatiform punctures on anterior border of pronotum in A. flavipes); scapal basin punctate-rugulose, vaguely cross-ridged medially (entirely cross-ridged in A. flavipes and A. similis); dorsum of propodeum polished and impunctate (dorsum of propodeum smooth and scarcely punctated in A. flavipes and A. similis); sparse punctation on disc of T2, with more than 1PD apart and medial impunctate polished strip clearly defined (disc of T2 densely punctated in A. similis, and medial impunctate polished strip not defined in A. flavipes); and purple highlights on metasoma (blue highlights in A. similis). The light brown antenna, mandible, and clypeus, and the anterior declivity of pronotum lacking medial pit also help to distinguish this new species from A. flavipes (dark brown antenna, mandible, and clypeus, and with well-marked medial pit on anterior border of pronotum); and the greenish blue body coloration contrasts with the light blue pattern of A. similis.
Holotype, male. Body length: 2.8 mm (Fig.
Head: 0.95× wider than high; toruli barely separated, inner margins slightly produced, forming a short lamellar projection; scape long, cylindrical, slightly wider basally, 3.8× longer than its maximum width; F1 length 2.2× breadth, 1.25× longer than F2, F2 1.15× longer than F3; F4 0.9× F3, F5–F10 slightly shorter than F4, F11 acute; lower margin of clypeus evenly round; subantennal distance 0.2× MOD; malar space 1.5× MOD; POL 1.6× OL, 4.3× OOL; inner ocular margin convergent submedially, LID 0.95× scape length; eye height 1.75× breadth.
Mesosoma: anterior declivity of pronotum lacking medial pit, lobe slightly separated from tegula; scutum notaulus barely distinguishable, impressed only basally (Fig.
Metasoma: lateral margins of T1–T3 sharp, but not carinate; first metasomal segment with ventral keel; posterior margin of S1 concave, posterior margins of S2–S4 nearly straight.
Coloration: head greenish blue, with greenish highlights on frons, scapal basin, and gena; bluish purple tint on vertex (Fig.
Sculpturing: head regularly punctate, punctures becoming clumped among ocelli and near face; vertex with rugose marks marginally (Fig.
Vestiture: short erect light brown setation on vertex, gena, and frons, with relatively longer and denser setation on occiput; mandible, clypeus and labrum with distinct long gold setation; antenna with short, decumbent, pale setation; eye with sparse microtrichia; dorsum of mesosoma with dense, relatively long, light brown setation; lateral pronotum, posterior border of mesopleuron, dorsum of propodeum, and metapleuron-propodeum, glabrous; wing membrane entirely setose; outer surface of metatibia, and pro- and metafemora, with long dark setae; venter of tarsomeres with abundant irregularly-sized spines; T1–T2 mostly glabrous, except for sparse short setae placed marginally; T3–T5 with marginal stripe of dense long dark setae; S2–S5 abundantly setose.
Female. Unknown.
Holotype: Brazil • ♂; Rio Grande do Norte, Mossoró, “Faz. Sta. Júlia” [Fazenda Santa Júlia], Malaise 1, caatinga; 05°01'10"S, 37°22'56"W; 14.iv.2008; Fernandes, DRR & cols; RPSP.
• 3♂ paratypes: same data as holotype (1♂
Brazil, RN: Mossoró.
The holotype is missing F6–F11 of the left antenna. The metasoma is damaged, with distal segments partly detached and upside down. Two paratypes lack heads, and another paratype has a detached metasoma. This species is only known from Mossoró, Rio Grande do Norte state (sympatric with A. sertaneja Lucena, sp. nov., see below), within the semiarid region. All specimens were collected in a Caatinga native vegetation fragment adjacent to a melon crop Cucumis melo L. (Cucurbitaceae) (
The new species is named after Daniell Fernandes (Boy), collector of many specimens used in this study.
Unknown.
Amisega flavipes Kimsey, 1987: 67. Holotype ♀ (USU: not examined). BRAZIL: Bahia, Encruzilhada.
Brazil• 1♂ paratype; Bahia, Encruzilhada; xi.1975; M Alvarenga;
Brazil, BA: Encruzilhada, Jequié.
This species can be readily distinguished from other species of the northeastern Brazil by the following combination of characters: scapal basin cross-ridged; dark brown antenna, clypeus, mandible, and metaleg; and lack of distinct striation on vertex and dorsum of mesosoma (Fig.
Unknown.
Dorsum of head and mesosoma densely striate, with sparse punctures inserted among striae; propodeum with rugose marks, becoming irregularly striate posteriorly, posterior declivity with some sparse punctures; lateral pronotal pit sulci-form; scapal basin transversely cross-ridged; metasoma brown, without blue nor purple highlights; legs light brown; malar space 1.7× MOD. This new species can be readily distinguished from other species occurring in northeastern Brazil, Amisega boyi Lucena, sp. nov., Amisega flavipes Kimsey, and A. similis Kimsey, based on densely striate vertex and mesosoma; rugose marks on propodeum; and lack of blue or purple highlights on the metasoma in both genders. Furthermore, A. sertaneja sp. nov. has the lateral pronotal pit shallow and sulci-form (pit well-marked in A. flavipes), and concolorous light brown legs in both sexes (bicolored in A. similis, and dark brown in males of A. flavipes). The extensive transverse striation on dorsum of head and mesosoma also resembles A. striata Kimsey (Costa Rica). However, A. sertaneja sp. nov. is readily distinguished from A. striata by its relatively shorter body (3.1 mm vs. 4 mm), lack of reddish highlights on the legs, brassy tints dorsally on the body, and the entirely dark brown scape (paler ventrally in A. striata).
Amisega sertaneja Lucena, sp. nov., holotype, male A lateral habitus B metasoma, dorsal view C head and mesosoma, lateral view D dorsal habitus E head, frontal view F head and mesosoma, dorsal view G head, dorsal view H metasoma, lateral view I propodeum, dorsal view. Scale bars: 1 mm (A, D), 0.5 mm (F, G), 0.2 mm (B, C, E, H, I).
Holotype, male. Body length: 3.1 mm (Fig.
Head: 1.2× higher than broad; toruli barely separated, inner margins slightly produced, forming a short lamellar projection (Fig.
Mesosoma: anterior declivity of pronotum with shallow medial pit, lateral pit near pronotal lobe sulci-form, lobe slightly separated from tegula (Fig.
Metasoma: lateral margins of T1–T3 sharp, but not carinate (Fig.
Coloration: head dark blue, with greenish highlights on frons, scapal basin, and gena; eye, antenna, and mouthparts light brown; distal flagellomeres light brown; mesosoma mainly dark blue, with greenish highlights dorsally; mesopleuron with ventral light brown tint, posterior margin with dark brown stripe; metapleuron-propodeum dark brown with marginal bluish highlights; tegula light brown; wing membrane light fuscous, veins brown; legs brown, darker on metacoxa; metasoma entirely brown (Fig.
Sculpturing: frons and vertex heavily striate (Fig.
Vestiture: short sparse pale setation on vertex, gena, and frons, with relatively longer and denser setation on lateral border of face and occiput; mandible, clypeus and labrum with distinct long gold setation; antenna with very short, decumbent, pale setation; eye with sparse microtrichia; dorsum of mesosoma with short sparse light brown setation; lateral pronotum, posterior border of mesopleuron, dorsum of propodeum, and metapleuron-propodeum, glabrous; wing membrane entirely setose; outer surface of metatibia, and pro- and metafemora, with long dark setae; venter of tarsomeres with abundant irregularly-sized spines; T1–T2 mostly glabrous, except for sparse short setae placed marginally; T3–T5 with marginal stripe of long dense dark setae; S2–S5 abundantly setose.
Female (Fig.
Holotype: Brazil • ♂; Rio Grande do Norte, Mossoró, “Faz. Sta. Júlia” [Fazenda Santa Júlia], Malaise 1, caatinga; 05°1'10"S, 37°22'56"W; 14.iv.2008; Fernandes, DRR & cols; RPSP • 9 paratypes same data as holotype (2♂1♀ RPSP; 1♂1♀ UNILA; 2♂
Brazil, PB: Santa Teresinha; RN: Mossoró.
The holotype is missing F10–F11. This species is only known from localities in the core zone of the Caatinga dry region. It is sympatric with A. boyi Lucena, sp. nov. in Mossoró-RN, which suggests both species are endemic to Caatinga. All specimens of both new species were collected between January to May, which is the period of most intense rainfall in the region.
Body size: 2.8–3.0 mm; F1 length 2.6–3.3× breadth.
The name is a Brazilian gentilic adjective for person living in the semiarid region.
Unknown.
Amisega similis
Brazil • 1♂ paratype; Pernambuco, Caruaru; iv.1972; M Alvarenga;
Brazil, BA: Encruzilhada; MG: Pedra Azul; PE: Caruaru; RJ: Rio de Janeiro and Mangaratiba.
This species can be distinguished from other species of the northeastern Brazil by its bicolored legs, blue highlights on dorsum of metasoma, and lack of striation on vertex and dorsum of mesosoma.
Unknown.
1 | Vertex and dorsum of mesosoma densely striate (e.g., Fig. |
Amisega sertaneja Lucena, sp. nov. |
– | Vertex and dorsum of mesosoma lacking striae or with only stratiform punctures or rugose marks on vertex and anterolaterally on pronotum (e.g., Fig. |
2 |
2 | Pronotum without any striae or rugose marks; metasomal terga with metallic blue highlights; malar space 1.3× MOD long; male F1 length 3.5× breadth | Amisega similis Kimsey |
– | Pronotum with discrete anterolateral stratiform punctation or rugose marks; metasomal terga with metallic purple highlights; malar space 1.5× MOD long; male F1 length shorter than 3× breadth | 3 |
3 | Anterior declivity of pronotum lacking distinct medial pit; scapal basin punctate-rugulose, vaguely cross-ridged medially (Fig. |
Amisega boyi Lucena, sp. nov. |
– | Anterior declivity of pronotum with deep medial pit (Fig. |
Amisega flavipes Kimsey |
Duckeia
Duckeia is a rarely collected taxon. Currently, the genus comprises three species, two of them were recorded in Rio de Janeiro state, southeastern Brazil, in an area of Atlantic rainforest, and one species is known from Costa Rica (
Body short, stocky build, not compressed laterally; posterior margin of eye lacking carina; upper gena slightly produced; malar space about half eye height; antennae brown, lighter on distal flagellomeres; legs light brown, becoming yellow in tarsomeres; metasoma with dense appressed punctation on marginal borders; dorsum of metasoma with distinct blue highlights. Duckeia dudui sp. nov. closely resembles D. gracile Kimsey, 1987. The new species differs from D. gracile by the following combination of characters: elongated head, 1.3× higher than broad (0.9× higher than broad in D. gracile); long malar space, about half of eye height (one third eye height in D. gracile); light brown legs (darker in D. gracile); short, sparse pale setation on dorsum of meso- and metasoma (relatively longer and denser setation on dorsum of meso- and metasoma in D. gracile); marginal borders of T1–T2 with contiguous punctures (punctation on marginal borders of T1–T2 more than 0.5–1.0 PD in D. gracile); dorsum of metasoma with distinct bluish highlights marginally on terga (only T2 with faint bluish tint in D. gracile). Furthermore, D. dudui sp. nov. has a stocky built and relatively short body, contrasting with the laterally compressed and relatively elongated body of D. gracile.
Duckeia dudui Lucena, sp. nov., holotype, female A lateral habitus B head, frontal view C head and mesosoma, dorsal view D occiput (female paratype) E head and mesosoma, lateral view F head, dorsal view G metasoma, dorsal view H amplified view of T1–T2, dorsal view I metasoma, lateral view J wings (female paratype). Scale bars: 0.5 mm (A, C, E, G, I), 0.2 mm (B, D, F, H, J).
Holotype, female. Body length: 3.4 mm (Fig.
Head: pyriform, 1.3× higher than broad (Fig.
Mesosoma: pronotum with discrete medial longitudinal sulcus, posterior margin not elevated upon scutum, lateral and posterior pronotal pits well-marked, lobe separated from tegula by about 0.7× tegular diameter (Fig.
Metasoma: lateral margins of T1–T2 carinate (Fig.
Coloration: head dark brown green, with green highlights on frons, vertex and gena, darker basally at malar area; mouthparts light brown; antenna brown, lighter on pedicel and distal flagellomeres; mesosoma dark green brownish, with bluish purple tints dorsally on propodeum; mesopleuron with ventral and posterior light brown stripe; tegula light brown; wing membrane slightly pale to subhyaline, veins brown; legs light brown, becoming yellowish on tibiae and tarsomeres; metasoma dark brown, metallic blue highlights on dorsum, becoming greenish marginally on T3–T4; disc of T1 and T2 brownish; sterna mostly dark brown, except for marginal bluish highlights on S2.
Sculpturing: head contiguously punctate (Fig.
Vestiture: sparse long golden pale setation on vertex, gena and clypeus, frons and face with relatively shorter and sparser setation; flagellomeres with decumbent dense pale setation, sparser and longer on scape and pedicel; eye with sparse microtrichia on upper border; mesosoma setose, except glabrous disc of metapleuron-propodeum; mesosomal dorsum with sparse, pale setation, longer on metanotum; mesopleuron with long dense pale setation, becoming longer ventrally; wing membrane entirely setose (e.g., Fig.
Male. Unknown.
Holotype: Brazil • ♀; Rio Grande do Norte, Patu, “caatinga-arm. Malaise” [Malaise trap in caatinga]; 06°06'S, 37°37'W; ix.2008; DRR Fernandes & colls; RPSP • 4 paratypes, same data as holotype (1♀
Brazil, RN: Patu.
The holotype is missing part of left metaleg. Two paratypes are poorly conserved, lacking antennae, legs and metasoma, thus, preventing gender determination. This species is only known from Patu county. Specimens were collected in the base of “Serra do Lima” (inselberg), 248 m above sea level (
The species is named after Sebastião Antônio de Araújo, Dudu (in memoriam), grandfather of the first author.
Unknown.
Cleptinae are rarely collected and there are only few specimens housed in Brazilian collections. These wasps are parasitoids of prepupal larvae of Tenthredinoidea sawflies (e.g.,
Cleptidea
This genus is neotropical, occurring mostly in forested warm habitats from southwestern Mexico, through Central America, into South America as far south as northern Argentina (
Head bicolored, mostly dark green extending from vertex until upper half of face, mid face to malar area orange; clypeus raised and square-shaped, with lower margin straight; scutum, scutellum and metanotum testaceous; propodeum entirely black, except for the whitish tip of lateral angle. Cleptidea nordestina sp. nov. most resembles C. fasciata (Dalman). The new species can be readily distinguished from C. fasciata by the following combination of unique characteristics: bicolored head, hyaline marks at body (especially trochanters, pronotum, and propodeal angles), clypeus produced, orange metanotum, and entirely black propodeum.
Cleptidea nordestina Lucena, sp. nov., holotype, female A lateral habitus B head, frontal view C dorsal habitus D head and mesosoma, dorsal view E metasoma, lateral view F metasoma, dorsal view G propodeum, dorsal view H head and pronotum, dorsal view. Scale bars: 1 mm (A–F), 0.5 mm (G, H).
Holotype, female. Body length: 6.7 mm (Fig.
Head: 1.2× wider than high; toruli 0.9× MOD separated; scape more or less cylindrical, broader apically, 3.4× longer than its maximum width; F1 length 3.1× breadth, 3.1× longer than F2; F2–F10 sub-equal in size, slightly flattened ventrally, F8–F11 excavated ventrally, F11 acute apically, slightly longer than F10; clypeus protruded medially, squared lobe, with lower margin slightly convex; malar space about 0.6× MOD; ocelli set in compact triangle on vertex, with transverse sulcus linking laterals; posterodorsal border of eye surpassing lateral ocellus dorsal limit; inner ocular margin sub-parallel, slightly convergent above; LID 1.1× eye height; eye height 1.5× breadth; OOL 2.3× OL, 2.6× POL; frons with short longitudinal sulcus below median ocellus briefly interrupted.
Mesosoma: pronotum crossed antero- and submedially by transverse crenulate grooves, longitudinal sulcus culminating in a deep fovea posteriorly (Fig.
Metasoma: posterior margin of S1 distinctly concave medially; posterior margin of S2–S4 slightly convex.
Coloration: head basally orange, extending from malar area to mid face, paler basolaterally, metallic dark green hue extending from upper half of face to vertex and gena (Fig.
Sculpturing: head heavily sculptured, densely punctate, rugulose-lacunose, particularly on vertex and frons, becoming sparser on gena, and impunctate basally near torulus and clypeus; pronotum, scutum and scutellum mostly rugulose-lacunose, lobe of pronotum impunctate; mesopleuron faintly costate above, punctate to rugulose-lacunose below; metanotum punctate on disk; femora and tibiae sparsely punctate; dorsum of propodeum heavily carinate forming areolate enclosures, posterior declivity longitudinally crossed by three main carinae, somewhat rugulose in-between; T1 nearly impunctate and polished posteromedially, with sparse tiny lateral punctures; T2 sparsely punctate on disk, marginal areas impunctate; T3–T4 densely punctate on disc, with broad impunctate stripes marginally; S1 and S4 nearly impunctate, with sparse marginal punctures; S2–S3 finely punctate on disk becoming sparser marginally.
Vestiture: head with sparse, long, pale setae on vertex; frons, face, gena and scape with relatively shorter and denser setation; flagellomeres with decumbent dense pale setation; eye with tiny, sparse, sub-erect microtrichia; dorsum of mesosoma with long, sparse, pale setation, longer on venter of mesopleuron; marginal depression of metanotum with patch of pale setae posteriorly; wing membrane entirely setose, with some distinctive, long, erect dark setae inserted proximally; legs with abundant long pale setation, comparatively longer on metaleg; inner surface of tarsi with short, thick setation and short spines irregularly distributed; disc of T1 and T2 glabrous, sparse setation only marginally, T3 and T4 with long dense setae posteriorly, S1 and S4 glabrous, S2 and S3 with long sparse setation; gonapophysis with some erect setae apically.
Male. Unknown.
Host. Unknown.
Holotype: Brazil • ♀; Bahia, Jequié, campus II UESB, Malaise III; 11.v.2007; Silva-Jr. JC & cols; RPSP.
Brazil, BA: Jequié.
The holotype is missing distal flagellomeres of the left antenna. The new species is only known from Jequié, located in a transition zone between Atlantic rainforest and Caatinga (Fig.
Biogeographic regions of easternmost South America, shown in Fig.
The name refers to the region where the new species was collected, and it is a Brazilian gentile adjective for a person native from the northeastern region.
Since
The present records of Duckeia species in quite distant sites in Central America and eastern Brazil without references to intermediate areas suggest a rudimentary knowledge of its diversity. We examined the type material of D. cyanea and D. vagabunda (
The diversity of Amiseginae and Cleptinae is probably the most underestimated in comparison with other groups of cuckoo wasps in South America. Their relatively small size and peculiar parasitoid lifestyles contribute to these wasps being rarely collected and frequently misidentified as other hymenopterans. The new species herein described in Amiseginae and Cleptinae enhance our understanding of chrysidid diversity in a poorly known Brazilian region. These new findings also reinforce the importance of long-term collecting studies and strategic samplings, which provide precious audited data potentially useful for selecting priority areas for conservation.
We are grateful to the aforementioned curators who kindly loaned specimens or allowed us to study specimens in their collections, and to Lynn Kimsey (
DAAL was supported by scholarships granted by the São Paulo Research Foundation (FAPESP): 2014/12407-0, 2015/12326-3, and 2017/25081-4; and FAPESP grant 2018/09666-5 to EABA. Additional funding for this research came from the CNPq (Brazilian National Council for Scientific and Technological Development) grants 422019/2018-6 and 310111/2019-6 and REFAUNA Program to EABA, and 501850/2009-0 to FCVZ. We are grateful for the support by Coordenação de Aperfeiçoamento de Pessoal de Nível Superior – Brazil (CAPES) – Finance Code 001.