Paratrigona lineata ( Lepeletier, 1836 ), 1994

Oliveira, Favízia Freitas de, Madella-Auricchio, Cláudia Renata & Freitas, Breno Magalhães, 2020, A new species of Paratrigona Schwarz, 1938 from northeastern Brazil, with notes on the type material of Melipona lineata Lepeletier, 1836 (Hymenoptera: Anthophila: Apidae), Journal of Natural History (J. Nat. Hist.) 54 (25 - 26), pp. 1637-1659 : 1650-1653

Paratrigona lineata ( Lepeletier, 1836) ( Figures 3–5 )

Melipona lineata Lepeletier, 1836 , Histoire Naturelle des Insectes- Hyménoptères. Paris:

Roret Vol. 1, 1–547 pp. [430; original description]. Trigona (Paratrigona) lineata ; Moure,

1943 [147; notes, geographic record]; Schwarz, 1938 [489; 492–494; 497; partim;

taxonomic note, key, figures: worker, male, wing (plate 59, A, A’, B, B’; plate 60, C)];

Trigona lineata ; Cockerell, 1917 [127; partim; taxonomic note]; Schwarz 1932 [239, 259;

citation]; Paratrigona (Paratrigona) lineata ; Moure, 1951 [58; taxonomic notes]; Trigona

(Paratrigona) lineata lineata ; Schwarz, 1943 [4–6; key, taxonomic note], Schwarz, 1948

[16, 171, 347, 348, 350–352, 354, 356–366, 370, 372, 374], 376, 377, 380, 383, 384,

390–392, 397, 400, 402, 403, 408, 410; partim; nesting site citation, taxonomy, key,

synonymy, diagnosis, geographic records, geographic range, description: male, figures:

male, worker]; Paratrigona lineata ; Camargo and Wittmann, 1989 [223; citation: nesting

site as obligate subterranean]; Moure, 1990 [117; taxonomic note]; Camargo, 1996 [108,

109, 110, 115, 119; partim; geographic range, biogeography, cladogram]; Camargo and

Roubik, 2005 [33, 35–37, 40, 42; taxonomic characters, comparative notes, phylogeny];

Camargo and Pedro, 2007 [424, partim; geographic record]; Camargo and Pedro, 2013

[424, partim; online catalogue; geographic record]; Paratrigona lineata lineata ;

Camargo, 1988 [357; taxonomic notes; geographic records]; Camargo and Moure,

1994 [33–40, 42, 44, 46, 58, 69–77, 79–82, 86–89, 91–93, 99, 100, 106; partim; biogeo-

graphy, key, taxonomy, diagnosis, geographic records, geographic range, phylogeny,

figures: worker, male); Camargo and Moure, 1996 [103, 110; partim; geographic range,

biogeography].

In the original description of Melipona lineata, Lepeletier (1836) did not mention how many specimens he included in the type material, mentioning only that the material was from Goiás State, Brazil [‘ Du Brézil : Capitainerie de Goyan’] and was deposited in the MNHP Collection [‘ Musée de France’] . We found two specimens in this collection ( Figures 3 , 4 ), both specimens bear identical labels (a small white rectangular label, with ‘Type’ handwritten in black ink, a white circular label with provenance data handwritten in black ink, and a cream-coloured square label printed with the phrase ‘ Sud de La Capit e de Goyaz’) ( Figures 3 (f), 4(f)) . One of the specimens has two additional labels: a small rectangular label with the name lineata handwritten in black ink and a larger label with the word ‘Lectotype’ printed on red paper (probably this red-printed label was added later by Dr. Jesus Santiago Moure, but there is no evidence to confirm this) ( Figure 3 (f)).

Camargo and Pedro (2007: 424) mentioned that the lectotype would have been designated by Moure (in litt., in 17.IV.1958), but both specimens deposited in the MNHP collection have no identification label added by Moure . They also stated that in his personal notes, Moure did not mention other type specimens besides the lectotype. In order to avoid future confusion about the identity of the type material, the specimen bearing the rectangular label with lineata handwritten in black ink and the larger lectotype red label had received an extra lectotype label printed on blue paper added by the first author of this article ( Figure 3 (f)). The second type specimen also had added a paralectotype label printed in green paper ( Figure 4 (f)).

The lectotype characteristically has a broad anchor-shaped stain occupying the central area of the clypeus ( Figure 3 (a)), with the labrum and mandibles completely yellowish (the mandibles brown contoured, especially at the apex); the supraclypeal area completely yellow; the parocular area with a wide yellow band, wider below until just before the tentorial pit and gradually narrowing upward; the scape yellow, with a brown band on the dorsal surface ( Figure 3 (a)); the pronotum with a wide band, interrupted in the middle; narrow band on the laterals of the mesoscutum; the pronotal lobe yellow; the axillae yellow on the dorsal surface ( Figure 3 (b–d)); the apical edge of the mesoscutellum with a broad band ( Figure 3 (b–d)); and a yellow spot at the base of the anterior, middle and posterior tibiae ( Figure 3 (b,e)). The specimen is very dirty, but it is possible to see the whitish pilosity on the mesosoma and head, and slightly more yellowish pilosity on the mesoscutum, especially the thicker setae.Relatively long setae are present on the entire mesoscutum ( Figure 3 (c,d)); the entire dorsal surface of the mesoscutellum ( Figure 3 (c,d)), including the apical border and ventral surface; as well as the entire surface of the mesepisternum.The metasoma is slightly more reddish than the head and the mesosoma, with the lower sides of T1 rather yellowish.The wing membrane is hyaline with a strongly iridescent lustre and brown veins ( Figure 3 (b)); the tegules are yellowish with a translucent spot on the disc. The microreticulation is quite strong in the basal area of the propodeum; on the mesepisternum the microreticulation is visible but duller (matt) and covered with very short plumose pilosity interspersed with much longer yellowish setae.

Lectotype measurements

Approximate total length: 4.1 mm; head width: 1.8 mm; thorax width: 2 mm; Anterior wing length: 3.8 mm. The specimen is extremely dirty and very fragile, making other measurements impossible.

Camargo and Moure (1994) mentioned that, as also in Moure’s personal notes, the type specimen of Melipona lineata Lepeletier, 1836 has erect setae on the entire surface of the mesoscutum, mesoscutellum and mesepisternum sides, a pattern also present in specimens from São Paulo and Minas Gerais ( Figure 5 (b,c)), what was confirmed in our reanalysis of the lectotype.

The paralectotype deposited at MNHP is doubtless a different species ( Figure 4 ), which means that the type material is composed of two species. It has smaller facial markings (a longitudinal stripe in the middle of the clypeus and two small stripes on each side of the clypeus, as well as narrower parocular bands) ( Figure 4 (a,b)). The mesoscutellum bears long erect setae only on the sides of the ventral surface ( Figure 4 (d)), but these are much shorter than in the lectotype. This specimen also lacks long erect setae on the mesoscutum and on the upper 2/3 of the mesepisternum ( Figure 4 (d)), differing from P. incerta only the reduction of stains on clypeus (possibly a variation).

Also according to Camargo and Moure (1994), the specimens from northeastern Brazil, including Piauí (UruÇui-Una), Maranhão (Balsas, Fortaleza dos Nogueiras and S. Raimundo das Mangabeiras), Ceará (Tabuleiro do Norte), and Paraíba (Santa Luzia, Serra do Brandao), have a glabrous mesoscutum and the dorsal surface of the mesoscutellum (entire disc) with short but very evident erect setae only on the distal margin of the mesoscutellum and the sides of the mesepisternum. Therefore, based on these characters, the specimens that they considered a variation of Paratrigona lineata lineata are treated here as Paratrigona intermedia Oliveira, Madella-Auricchio & Freitas sp. nov ( Figure 8 ).

In view of the geographic distribution and patterns of variation in specimens from northern, northeastern and southeastern Brazil, including those mentioned by Camargo and Moure (1994), we conclude that Paratrigona lineata ( Lepeletier, 1836) sensu Camargo and Moure (1994) is a species repository, from which we separate Paratrigona intermedia Oliveira, Madella-Auricchio & Freitas as a new species. As a consequence the new concept and distribution for P. lineata ( P. lineata lineata ) here are different than that assumed by Camargo and Moure (1994) and Camargo and Pedro (2013), being restricted, according to their geographical records, to Bolivia (Santa Cruz) and Brazil (Pará, Goiás, Minas Gerais, Bahia, Mato Grosso, São Paulo, Paraná).

Lectotype conservation state: very dirty specimen. The left middle tibia and following parts are glued to the pin. The specimen is missing the left posterior distitarsus from the second tarsus and the right posterior distitarsus from the first tarsus, and the entire flagellum of the left antenna. The right posterior wing is badly damaged at the lower margin.

Key to species of Paratrigona (workers) from the lineata species group [Modified from Camargo and Moure (1994) and Gonzalez and Griswold (2011)]

1 Anterior margin of mesoscutum a gently rounded slope and its surface uniformly matt and reticulated; absense of maculations on frons, genae (except for some small spots in some specimens of Paratrigona subnuda ), mesepisternum and metepisternum (including epimerum); antennal scape not enlarged-flattened, cylindrical, slightly wider, equal to or narrower than width of F3; submarginal angle acute (less than 90°); malar space narrower than half the diameter of F3; head elongated..... 2 ( lineata group)

2(1) Mesoscutellum short-paraboloid, length about 0.55 times width; parocular yellow band usually sinuous at level of antennal alveolus.. Paratrigona nuda ( Schwarz, 1943) Mesoscutellum short-semicircular, length 0.50 to 0.51 times width; parocular yellow band not sinuous at level of antennal alveolus........................................................................... 3 3(2) Scape narrower than width of F3 ............................................................................................. 4 Scape equal to or slightly wider than width of F3...................................................................... 8 4(3) Long erect setae relatively dense on upper front to vertex ( Figure 5 (a)), disc of mesoscutum and upper 2/3 of mesepisternum ( Figure 5 (c)) ( Bolivia, northern Brazil south to Paraná)................................................................ Paratrigona lineata ( Lepeletier, 1836) Long erect setae only surrounding ocelli (very few in some species) and vertex ( Figures 1 (a), 6(a), 7(a)); disc of mesoscutum without long erect setae ( Figures 1 (d), 6(e), 7(d)); upper 2/3 of mesepisternum with or without long erect setae............................................ 5 5(4) Upper 2/3 of mesepisternum with long erect setae, longer than decumbent plumose pubescence [ Figures 1 (c), 2(a); and figure 9 of Gonzalez and Griswold (2011)] ........................................................................................................................................................... 6 Upper 2/3 of mesepisternum without long erect setae ( Figures 6 (c), 7(c))...................... 7 6(5) Erect setae on upper 2/3 of mesepisternum dense and long (at least half of median ocellar diameter) (figure 9 of Gonzalez and Griswold 2011); mesoscutellum dorsally with long erect setae (about as long as median ocellar diameter), including disc and distal half (figure 10 of Gonzalez and Griswold 2011); longest setae on mesoscutellum on yellow stripe ( Paraguay)................ Paratrigona wasbaueri Gonzalez & Griswold, 2011 Erect setae on upper 2/3 of mesepisternum fine, sparse and short (about 1/4 of median ocellar diameter) ( Figure 1 (c)); setae on mesoscutellum short (longest setae about half of median ocellar diameter), restricted to distal margin and some on dorsal yellow stripe ( Figure 1 (d)); longest setae on mesoscutellum on ventral surface (northeastern Brazil: Maranhão, Piauí, Paraíba and Ceará)....................................................................................... ................................... Paratrigona intermedia Oliveira, Madella-Auricchio & Freitas , sp. nov 7(5) Mesoscutellum with erect setae on distal margin, including also some on dorsal surface adjacent bordering the distal margin ( Figure 6 (d,e)); face with pale yellow maculations ( Figure 6 (a)) (northern Argentina to Bolivia)............................................................ ................................................................................. Paratrigona glabella Camargo & Moure, 1994 Mesoscutellum with erect setae on ventral surface only, absent on distal margin ( Figure 7 (d)); face with bright-yellow maculations ( Figure 7 (a)) (northern Minas Gerais, Brazil). ................................................................................... Paratrigona incerta Camargo & Moure, 1994 8(3) Scape distinctly broader (1.2–1.3 times) than width of F3 (Pará and Amazonas, Brazil).............................................................. Paratrigona crassicornis Camargo & Moure, 1994 Scape about as broad as or slightly broader than width of F3 (southeastern Brazil)....... ....................................................................................................... Paratrigona subnuda Moure, 1947 Malar area length about 2/3 width of F3; frontal maculation not distinctly broad above bifurcation; clypeus entirely yellow, at most darkened on disc (Amazonas, Brazil)........... ................................................................................. Paratrigona compsa Camargo & Moure, 1994

Figures

Figure 1. Holotype female (worker) of Paratrigona intermedia Oliveira, Madella-Auricchio & Freitas sp. nov., deposited at the MHNBA, Salvador, Bahia State, Brazil: (a). Frontal view of head; (b). Lateral view of body; (c). Mesepisternum in profile, and iridescence on wing; (d). Dorsolateral view of mesoscutum and mesoscutellum. Arrows indicate presence or absence of setae.

Figure 3. Lectotype of Melipona lineata Lepeletier, 1836 deposited at the MNHP, Paris, France: (a). Frontal view of head; (b). Lateral view of body; (c, d). Dorsolateral view of mesoscutum and mesoscutellum; (e). Metatibia; (f). Labels of lectotype. Arrows indicate presence of setae.

Figure 4. Paralectotype of Melipona lineata Lepeletier, 1836 deposited at the MNHP, Paris, France: (a). Frontal view of head; (b). Lateral view of head; (c). Lateral view of body; (d). Dorsolateral view of mesosoma; (e). Metatibia; (f). Labels of paralectotype. Arrows indicate absence of setae.

Figure 5. Specimen of Paratrigona lineata (Lepeletier, 1836) from Januária (Minas Gerais State, Brazil), deposited at the BIOSIS, Institute of Biology, Federal University of Bahia, Brazil: (a). Frontal view of head; (b). Lateral view of body; (c). Dorsolateral view of mesosoma; (d). Labels. Arrows indicates presence of setae.

Figure 6. Paratype of Paratrigona lineata glabella Camargo & Moure, 1994, deposited at the Prof. J.M.F. Camargo Collection from the RPSP, University of São Paulo, Ribeirão Preto, Brazil: (a). Frontal view of head; (b). Lateral view of body; (c). Lateral view of mesepisternum; (d). Dorsal view of mesoscutellum; (e). Dorsal view of mesosoma; (f). Labels. Arrows indicate absence or presence of setae. Ownership rights to these images and their copyright belong to RPSP (Coleção Entomológica ‘Prof. J.M.F. Camargo’). Photo by E.A.B. Almeida.

Figure 7. Holotype of Paratrigona incerta Camargo & Moure, 1994, deposited at the Prof. J.M.F. Camargo Collection from the RPSP, University of São Paulo, Ribeirão Preto, Brazil: (a). Frontal view of head; (b). Lateral view of body; (c). Lateral view of mesepisternum; (d). Dorsal view of mesosoma; (e). Labels. Arrows indicate presence or absence of setae. Ownership rights to these images and their copyright belong to RPSP (Coleção Entomológica ‘Prof. J.M.F. Camargo’). Photo by E.A.B. Almeida.

Figure 8. Geographic distribution for species of Paratrigona Schwarz, 1938 from the lineata group in South America, based on the present data and the material cited by; Camargo and Moure (1994), and by; Camargo and Pedro (2013).