Scaptotrigona nigrohirta, Nogueira & Santos-Silva & Carvalho & Carvalho-Zilse & Alves & Oliveira, 2022

Nogueira, David Silva, Santos-Silva, José Augusto dos, Carvalho, Matheus Mourão, Carvalho-Zilse, Gislene Almeida, Alves, Rogério Marcos de Oliveira & Oliveira, Marcio Luiz, 2022, Two new species of Scaptotrigona Moure, 1942 from the Amazon forest (Hymenoptera: Apidae: Meliponini), EntomoBrasilis (e 985) 15, pp. 1-10 : 2-6

Scaptotrigona nigrohirta sp. nov. Nogueira & Santos-Silva

urn:lsid:zoobank.org:pub:66BBD0DA-E731-4379-8B9B-D57F66CB6A63

Nannotrigona (Scaptotrigona) nigrohirta: POSEY 1982: 457 [nomen nudum, list]; POSEY 1983: 71 [nomen nudum, list]; CAMARGO & PEDRO 2013 [nomen nudum, catalog, incertae sedis, citation].

Nannotrigona (Scaptotrigona) xanthotricha: COIMBRA JR. 1985 : 66 [geographic record, folk name: yoidíter-éy].

Scaptotrigona nigrohirta : POSEY & CAMARGO 1985: 250, 252, 254, 261, 266-268 [nomen nudum, nest entrance, behavior, habitat, figures 2D and 4A, folk names: imrê ñy kamrek, imrênhy-kamrek, “red imrê”]; POSEY 1986: 256, 261 [nomen nudum, habitat, folk name: imrê-ñy-kamrek]; CAMARGO & POSEY 1990: 18, 19, 21-23, 25, 31, 35, 36 [nomen nudum, nest entrance, behavior, habitat, folk name: imerê-ñy-kamrek, figures 2D and 4A]; CAMARGO & ROUBIK 1991: 27 [nomen nudum, nest]; CAMARGO 1994: 53 [nomen nudum, behavior, nest]; KERR et al. 1996: 35 [nomen nudum, nest entrance, figure 3D]; VENTURIERI 1999: 94 [nomen nudum, geographic record, floral resource]; VENTURIERI & IMPERATRIZ-FONSECA 2000: 356 [nomen nudum, geographic record, management, nest, behavior]; OLIVEIRA 2002: 616, 628 [nomen nudum, nest, folk name: cu-de-vaca, bui txashkaati]; CRUZ-LANDIM et al. 2006: 169 [nomen nudum, class III glands size]; CORTOPASSI-LAURINO et al. 2006: 280 [nomen nudum, geographic records]; FRANÇOSO & ARIAS 2013: 847 [nomen nudum, COI sequence KC853354 ]; REZENDE et al. 2019: 255, 256, 257, 258, 260, 262, 264, 265, 266 [nomen nudum, geographic record, floral resource]; FERREIRA 2020: 46, 47, 57, 61, 62, 67 [nomen nudum, geographic record, floral resource].

Trigona (Scaptotrigona) xanthotricha: BAUMGARTNER & ROUBIK 1989: 16-19 [geographic record, behavior].

Scaptotrigona xanthotricha : BAUMGARTNER & ROUBIK 1989: 14 [geographic record]; NATES-PARRA 2001: 240 [geographic record]; COSTA 2010: i-iii, vii-xxi, 1, 4-6, 13, 16, 18-20, 23, 26-28, 30, 32-34, 36, 38, 40-44, 47-50, 59-61, 65, 70-72, 74, 76, 79, 81, 83, 84, 86-88, 100-102, 109-111, 114-126, 128, 134-136, 139, 140, 142-144, 146, 148-151, 153, 159 [behavior, nest entrance, honey, pollen and queen production, chapter 1 figures: 1, 3-10; chapter 4 figures: 1-3]; BARBOSA-COSTA & CARVALHO-ZILSE 2013: 8, 91-103 [biology, geographic record, figures 1, 2, 4]; CARVALHO-ZILSE 2013: 21, 22 [nest entrance, figure 1D]; COSTA-NETO et al. 2016: 143 [geographic record, folk name: trombeta de macaco]; MAIA et al. 2020: 1378 [geographic record];

Scaptotrigona sp.n nigrohirta : DELGADO & VELA 2012: 81,

82, 85 [nomen nudum, floral resource, geografic record, characteristics, figure 1C].

Scaptotrigona aff. xanthotricha : FERREIRA et al. 2020: 1-7 [as Scaptotrigona xanthotrica , sic (page 2), poisoning test, folk name: canudo]; PIRES et al. 2020: 59252, 59254, 59258, 59260, 59262 [geographic record, honey characters, folk name: canudo]; VIANA et al. 2021: 4 [as Scaptotrigona aff. xanthotricha , sic, geographic record, folk name: canudo];

Diagnosis: Bees in general yellowish brown with black terga with yellowish brown apical band and with or without yellowish brown maculations on T1 and T2. Head black with lower half (up to half of the antennal socket) depigmented and vitreous, as well as the clypeus and labrum. Very similar to S. xanthotricha Moure, 1950 , except for the rounded shape on the posterior border of the metabasitarsus ( Figure 2E ). In general,it has dark brown to black hairs on the metabasitarsus and vertex. Area of the gena strongly carinated in the shape of two spines ( Figure 2C ).

Description: Worker ( Figures 1 , 2A, 2C, 2E , 3B and 4A ).

Coloration: Integument predominantly yellowish-brown. Labrum, clypeus, radicle, malar area and supraclypeal area yellowish-brown. Gena yellowish-brown with superior area black. Mandible brown with apical half reddish-brown and with brown condyles. Scape yellowish-brown. Pedicel dark brown. Flagellum dark brown with brown in ventral view of F1 to F10. Pronotal lobe, propodeum, scutellum and axilla yellowish-brown. Mesepisternum yellowish-brown with a brown maculation on inferior area. Mesoscutum black. Legs yellowish-brown with metabasitarsus and other tarsomeres brown. Tegula yellowish vitreous. Pterostigma and wing venation brown. Wing membranes slightly smoky with microtrichia brown more abundant in the apical area of the Rs vein. T1 with vertical area yellowish-brown vitreous and discal area black with apical band yellowish-brown. T2-T6 black with apical band yellowish-brown. Sterna yellow.

Punctation: Integument predominantly punctate. Head and mesosoma shiny and punctate. Legs and sterna smooth and shiny with pilligera puntaction with rugose aspect. Terga matte.

Pubescence: Frons with yellow simple setae (0.2 mm). Malar area practically glabrous with very short setae (0.03 mm). Basal area of scape with long, simple and yellowish setae (0.10 mm). Vertex with black long simple setae (0.26 mm). Clypeus and supraclypeal area in lateral view with decumbent yellow simple setae (0.01 mm). Labrum with yellow simple setae (0.21 mm). Coxae with whitish simple hairs: procoxa (0.25 mm), mesocoxa (0.29 mm) and metacoxa (0.42 mm). Mesocoxa and metacoxa with hairs with delicate crimped apex. Trochanters with whitish simple hairs with delicate crimped apex: protrochanter (0.25 mm), mesotrochanter (0.24 mm) and metatrochanter (0.37 mm). Basal area of profemur with whitish simple hairs with delicate crimped apex (0.21 mm), and short branched hairs (0.07 mm) and apical area with erect setae (0.11 mm) and very short branched hairs on ventral face of profemur (0.04 mm). Basal area of mesofemur with whitish simple hairs with delicate crimped apex (0.26 mm), and short branched hairs (0.05 mm), and apical area with short erect brown setae (0.07 mm) and very short branched hairs on ventral face of mesofemur (0.05 mm). Metafemur with whitish hairs with delicate crimped apex on basal area (0.09 mm) and with long erect branched brown to black setae on apical area (0.26 mm). Protibia (0.24 mm), mesotibia (0.35 mm) and metatibia (0.36 mm) with simple and branched black setae on posterior border. Corbicula area with long black setae with delicate crimped apex (0.61 mm). Probasitarsus with brown to black simple setae (0.33 mm). Mesobasitarsus with brown to black simple and branched setae and branched hairs (0.26 mm). Metabasitarsus with simple brown and black setae (0.27 mm). Mesoscutum with decumbent simple (0.05 mm) and branched (0.08 mm) yellowish hairs and anterior area with erect simple and branched black setae (0.20 mm). Mesoscutellum with simple (or with branch) yellow and black setae (0.37 mm). Sides of propodeum with long simple yellowish hairs (0.22 mm) and short branched hairs (0.10 mm). Mesepisternum with simple erect yellowish setae (0.14 mm) and branched decumbent yellowish hairs (0.05 mm). Inferior area of mesepisternum with simple yellowish hairs with delicate crimped apex (0.35 mm). Metepisternum with very abundant decumbent yellowish branched hairs (0.10 mm). T1 with decumbent yellowish branched hairs (0.05 mm). T2 with decumbent yellowish branched hairs (0.06 mm) and long simple and branched black setae (0.15 mm). T3 with branched black setae (0.19 mm) and simple decumbent branched yellowish hairs (0.11 mm). T4 with branched black setae (0.24 mm) and branched yellowish hairs (0.14 mm). T5 with branched and simple brown to black setae (0.20 mm) and branched yellowish hairs (0.08 mm). T6 with branched and simple brown to black setae (0.21 mm) and decumbent branched yellowish hairs (0.08 mm). Sterna with abundant long yellowish hairs with delicate crimped apex (0.42 mm).

Structures: Body length 6.06 mm. Head 1.24x wider than long (2.24 mm: 2.79 mm, length and width, respectively). Forewing length 5.77 mm. Length of compound eye 3.82x breadth (1.57 mm: 0.41 mm, length and width of the compound eye, respectively). Inner orbits concave, converging downwards (1.76 mm: 1.64 mm, superior and inferior interorbital distance, respectively). Maximum interorbital distance greater than the length of the eyes (1.89 mm: 1.57 mm). Malar area 2.12x of the flagellum diameter (0.34 mm: 0.16 mm). Clypeus with width 1.85x its length (0.7 mm: 1.3 mm = length and width of clypeus respectively). Clypeus-ocellar distance 1.31 mm. Alveolorbital distance equal to interalveolar distance (0.50 mm). Vertex slightly convex. Ocellorbital distance slightly greater than interocellar distance (0.52 mm: 0.50 mm). Scape length 6.73x its width (1.01 mm: 0.15 mm). Flagellum width slightly greater than the pedicel width (0.16 mm: 0.13 mm). Mandible with the second tooth wider than the first. Mesotibial spur 0.26 mm long. Metatibia with triangular corbicula; length of metatibia 2.1x its width (2 mm: 0.95 mm). Metabasitarsus 1.78x longer than broad (1.14 mm: 0.64 mm) and with posterior bord angled and projected. Intertegular distance 1.82 mm. With 7 hamuli on posterior wing. Mesoscutellum 1.27x wider than long (0.79 mm: 1.27 mm, length and width, respectively).

Etymology and remarks: This species has been widely cited in the literature, however, it has never been officially published to this moment. It was being described by Jesus Santiago Moure , however, he did not have the opportunity to publish it before his death. The specific epithet given by him was “nigrohirta ”, which we decided to keep in his honor. The etymology comes from Latin which means “black erect setae”, and probably alludes to the simple setae present in the vertex, basitarsi and terga of most specimens. When we are studying nest material, we noticed that this character varies, as there are individuals with yellowish-brown setae in these areas. Also, this name can be an allusion to the setae present in the anterior portion of the mesoscutum, as they are black and elongated in relation to the setae of the same region in S. xanthotricha , which are shorter and brownish.

Several studies that used other species names such as S. xanthotricha (or S. aff. xanthotricha ) to refer to S. nigrohirta sp. nov. (e.g., COIMBRA JR. 1985; BAUMGARTNER & ROUBIK 1989; NATES-PARRA 2001; COSTA 2010; CARVALHO-ZILSE 2013; BARBOSA-COSTA &

CARVALHO-ZILSE 2013; FERREIRA et al. 2020; MAIA et al. 2020; PIRES et al. 2020; VIANA et al. 2021), however, it is evident that they are distinct species, since the T1 of S. nigrohirta sp. nov. it has abundant branched hairs while S. xanthotricha are sparse ( Figure 2A and B ), the shape of the metabasitarsus is different, with the middle posterior border of S. xanthotricha is angled and S. nigrohirta sp. nov. is rounded ( Figure 2E, F and G ), in addition to the M and Cu1 veins of the forewing being strong in S. nigrohirta sp. nov. ( Figure 3A, B and C ). In relation to S. pectoralis (Dalla Torre, 1896) , the yellow depigmentation on the forehead is lower, until the midline of the antennal socket in S. nigrohirta sp. nov. ( Figure 1B and 2D ) and S. xanthotricha ( Figure 3D ).

Based on the distribution of species presented by CAMARGO & PEDRO (2013) and PEDRO (2014), on the genetic support presented by DUARTE et al. (2014) and in the data presented in this present study, we define here that the distribution of occurrence of S. xanthotricha is restricted to areas of Atlantic forest and S. nigrohirta sp. nov. to areas of the Amazon rainforest.

Furthermore, S. nigrohirta sp. nov. can be distinguished from S. fulvicutis (Moure, 1964) , which is the other yellow Amazonian species, because S. fulvicutis has a yellow mesoscutum, the face in frontal view with strong punctuation and clypeus with abundant setae, while S. nigrohirta sp. nov. has dark brown to black mesoscutum, face with punctuation sparse and the clypeus practically glabrous, with sparse and very short setae.

Variations: Some individuals from Campo Novo (Rondônia, Brazil), Barcelos and Jaú National Park (Amazonas, Brazil) have a yellow (depigmented) spot on the forehead above the upper socket line, as well as in S. pectoralis ( Figure 2D ). An individual from Terra Indígena São Marcos-Guaribas (Roraima, Brazil) has a black spot at the base of the clypeus. Some individuals from Belterra (Pará, Brazil) have two yellow triangles joined by a vertex at T2 ( Figure 2A ).

A label of one of the bees from Campo Novo (Rondônia, Brazil) was handwritten “ Scaptotrigona nigrohirta var. xanthina in litt.”, however, without identifying who wrote. And another from the Toototobi indigenous area (Yanomami indigenous people, Barcelos, Amazonas, Brazil) written “ N. (Scaptotrigona) chorerraensis Schwarz MS, Camargo 1982Det. ”, but this name was never formally published for the species.

Biological notes: Bees are highly defensive and build the nest entrance in a cylindrical shape ( Figure 4A ) ( POSEY & CAMARGO

1985; COSTA 2010; BARBOSA-COSTA & CARVALHO-ZILSE 2013).

Geographical distribution ( Figure 6 ): Brazil (Acre, Amazonas, Maranhão, Pará, Rondônia, Roraima), Peru (Nauta).

Examined Material: S. nigrohirta sp nov.: Holotype: worker. Label data: BRA, AM, Manaus, 26.xi.2021, INPA 1, ninho portaria . Paratypes (273), workers. BRAZIL: Amazonas: 2 ( INPA), Brasil - AM, PARNA do Jaú, 14-20 Out 2003, M.L. Oliveira & J.A. Cunha leg., Em ninho / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2009; 1 ( INPA), BRASIL. Amazonas. Serra do Aracá, Acampamento base. 108 m. a. 00°52’26.4”N / 63°27’18.0”W, 09/viii/2007. ninho, M.L. Oliveira & A. S. Filho leg. / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2009; 6 ( INPA), RPSP 814c / Samaúma, rio Demini, AM , Brasil, 24.VII.1999, 62°46’W. 00°22’S, Camargo, Pedro, Mazucato / Scaptotrigona sp.n., Camargo & Moure, MS, Det. Camargo, 2000 / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2007; 5 ( INPA), BRA, AM, Iranduba, Sítio Hélio, 08.xi.2021, Carvalho-Zilse e eq. (12); 1 ( INPA), BRASIL - AMAZONAS, LÁBREA - RIO ITUXI, 1 - X - 76, EDUARDO PRETAS / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2009; 3 ( INPA), BRA, AM, Manaus, 19.vii.2019, INPA 1, D. Nogueira leg.; 63 ( INPA), BRA, AM, Manaus, 26.xi.2021, INPA 1, ninho biblioteca; 35 ( INPA), BRA, AM, Manaus, 26.xi.2021, INPA 1, ninho portaria; 3 ( RPSP) idem; 3 ( MZSP) idem; 3 ( MPEG) idem; 3 ( ZSM) idem; 1 ( INPA), BRA, AM, Maués, Sítio Dedeco, 27.x.2021, Carvalho-Zilse e eq. (15); 3 ( INPA), BRA, AM, Maués, Sítio Dedeco, 27.x.2021, Carvalho-Zilse e eq. (24); 5 ( INPA), BRA, AM, Maués, Sítio Dedeco, 27.x.2021, Carvalho-Zilse e eq. (28); 3 ( INPA), RPSP 803c / Samaúma, rio Daarã, AM, BRASIL 17-20.VII.1999, 64°45’W, 00°26’S, Camargo, Pedro, Mazucato / Scaptotrigona sp.n., Camargo & Moure, MS, Det. Camargo, 2000 / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2007; 6 ( INPA), RPSP 766c / Plano, rio Negro, AM, BRASIL 07- 09.VII.1999, 65°59’W. 00°18’S, Camargo, Pedro, Mazucato / Scaptotrigona sp.n., Camargo & Moure, MS, Det. Camargo, 2000 / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2007; Pará: 2 ( INPA), BRASIL Pará. Santarém.V, Alter do Chão. 18/VIII /2002, Glória Assis. leg. / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2007; 3 ( INPA), BRASIL, Pará, Alter do Chão. Ilha Campo. 24.ix.1989. U.C. Barbosa / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2007; 6 ( INPA), BRASIL, PA, Belterra, FLONA Tapajós, 100 m, 02°36’15”S - 54°56’35”W / 15-16.iv.2008, manual, J.A.Rafael, F.F.Xavier F° / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2009; Rondônia: 107 ( INPA), Brasil Rondonia, Ig. [Igarapé] Tracoazinho, Reg. CpoNovo [Campo Novo] / 5 viii 1985, V. Py.Daniel, L Aquino / Scaptotrigona nigrohirta sp.n Moure, MS, Det.Camargo,2007; 2 ( INPA), BRASIL: Rondônia, Vilhena, 22 vii - 5 viii 1983, F.J.A. Peralta / 124-c / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2007; Roraima: 6 ( INPA), Brasil, Roraima, 23-à 28-II-2003 / M.L.Oliveira & S.J.R. Silva leg. A.I. São Marcos Guaribas, Em Ninho / Scaptotrigona nigrohirta sp.n Moure, MS, Det. Camargo, 2009.

S. pectoralis (36): COSTA RICA: San José: 34 ( ZSM), Costa Rica Pr San José, Villa Lapas b. Tarcoles , Trockenwald 50 m, N9°45’326” W64°36’511”, 18.04.2003 1. M. Kraus / Scaptotrigona pectoralis Det. D. Nogueira, 2021 ; 1 ( ZSM), idem except: 19.04.2003 ; MEXICO: Chiapas: 1 ( INPA), Mex, Chis, Jaltenango Paz , 886 m, 15,78 N; 92,62 W, 3/11/2017, Philippe Sagot / [verse] Scapec ♀ / [verse] Scaptotrigona pectoralis ♀ det. Sagot Ph. 2017 .

S. xanthotricha (24): BRAZIL: Bahia: 1 ( INPA), BRA, Bahia, Salvador , UFBA, 23.ix.2019 ; 1 ( INPA), BRA, Bahia, Salvador, UFBA ( Ondina ), 11.iv.2013 / Scaptotrigona xanthotricha Moure, 1950 , Det. D. Nogueira, 2019; Minas Gerais: 1 ( INPA), Reinventário EPDA de Peti, 9779 - 28895 / São GonÇalo do Rio Abaixo MG, BRASIL 23/09/2003, N. G. F o n s e c a / Scaptotrigona xanthotricha (Moure, 1950) , L. M. Santos, det. 2006 ; 17 ( INPA), BRA, Minas Gerais, ViÇosa, UFV Apiário , 17.ix.2019 / Scaptotrigona xanthotricha Moure, 1950 , Det. D. Nogueira, 2019; São Paulo: 4 ( ZSM), Brasilien São Paulo , v. Ihering / [verse] Trigona dorsalis , 1913 Friese det., Ym. [?] / Scaptotrigona xanthotricha, Det. D. Nogueira, 2021 .

S. fulvicutis (16): BRAZIL: Amazonas: 1 ( INPA), BRASIl. Amazonas. Itacoatirara. AM-010. km 215. Faz Aruana. 0302S / 5850W. 21.viii.1991 F.J. Peralta / Scaptotrigona fulvicutis (Moure, 1964) Det. Camargo, 2001 ; 1 ( INPA), BRASIL: Amazonas 60 Km. N. Manaus Fazenda Esteio ZF-3 Km-23 7-5- 90 M. L. Oliveira / 2150 / Scaptotrigona fulvicutis (Moure, 1964) Det. Camargo, 2007 ; 1 ( INPA), idem except: 23-8-90 / 2692; 1 ( INPA), idem except: 4-6-90 / 2287; 1 ( INPA), idem except: 27-11-90 / 0893; 1 ( INPA), idem except: 12-3-90 / 1893; 1 ( INPA), idem except: 7-4-90 / 2168; 1 ( INPA), idem except: 03-4-90 / 2046; 1 ( INPA), idem except: 23-2-90 / 1730; 1 ( INPA), LO-1 500m / Brasil. Amazonas REBio Uatumã 29-III-2008 Presidente Figueiredo A. C. O. Araújo & H.F. Guarlento / Scaptotrigona fulvicutis (Moure, 1964) Det. Camargo, 2009 ; Pará: 6 ( INPA), BRASIL. Pará Oriximiná. Porto Trombetas Platô Avião 12/ II/2008 0554685/9806548-UTM Y. Antonin, M.L. Oliveira leg / Scaptotrigona fulvicutis (Moure, 1964) Det. Camargo, 2009 .

Figures

Figure 1. Holotype of Scaptotrigona nigrohirta sp. nov.. Metatibia and metabasitarsus (A), head in frontal view (B), metasoma in dorsalposterior view (C), body in lateral view (D) and body in dorsal view (E). Scales 0.5 mm (A and B) and 1 mm (C, D and E). Source: authors.

Figure 2. Structures of Scaptotrigona Moure, 1942. T1 and T2 of S. nigrohirta sp. nov. (A) and S. xanthotricha Moure, 1950 (B). Head in lateral view of S. nigrohirta sp. nov. (C) and S. pectoralis (Dalla Torre, 1896) (D). Metabasitarsus of S. nigrohirta sp. nov. (E), S. xanthotricha (F) and S. pectoralis (G). Scales: 0.5 mm (A, B, C and D) and 0.25 mm (E, F and G). Source: authors.

Figure 3. Forewing and head of Scaptotrigona Moure, 1942 species. Forewing of S. hylaeana sp. nov. (A), S. nigrohirta sp. nov. (B), and S. xanthotricha Moure, 1950 (D). Head of S. xanthotricha (D) and S. pectoralis (Dalla Torre, 1896). Scales 1 mm (A, B, and C) and 0.5 mm (D and E). Source: authors.

Figure 4. Nest structures of Scaptotrigona Moure, 1942 species. Nest entrance of Scaptotrigona nigrohirta sp. nov. in a trunk in Instituto Nacional de Pesquisas da Amazônia (INPA, Manaus, Amazonas, Brazil) (A). Nest entrance of Scaptotrigona hylaeana sp. nov. in a trap-nest in Scientific Meliponary of Grupo de Pesquisas em Abelhas from INPA (Manaus, Amazonas, Brazil) (B and C). Brood comb in superior view (D), food pots (E) and queen (F) of S. hylaeana sp. nov.. Scale bar 1 centimeter. Source: authors.

Figure 6. Distribution records of some species of Scaptotrigona Moure, 1942: S. hylaeana sp. nov., S. nigrohirta sp. nov., S. fulvicutis (Moure, 1964), S. pectoralis (Dalla Torre, 1896) and S. xanthotricha Moure, 1950.