The taxonomic history of Trigona latitarsis Friese, 1900 and its clarification based on the observation of the types and literature data are treated in this study. The paper discusses the validity of the previously proposed lectotype, deposited in the ZMB (Berlin, Germany). Based on the type series deposited in HNHM (Budapest, Hungary) as well as the original description, a new lectotype and 15 paralectotypes from Amazon forest (São Paulo de Olivença, Amazonas) are designated. Data on the geographic distribution of Scaura latitarsis (Friese, 1900) are provided.

A história taxonômica de Trigona latitarsis Friese, 1900 é tratada nesse estudo, bem como esclarecimentos com base na observação dos tipos e em dados de literatura. É discutida a validade do lectótipo proposto anteriormente e depositado na ZMB (Berlim, Alemanha). Com base na série-tipo depositada no HNHM (Budapeste, Hungria), conforme descrição original, lectótipo e paralectótipos da floresta Amazônica (São Paulo de Olivença, Amazonas) são designados. Dados sobre a distribuição geográfica de Scaura latitarsis (Friese, 1900) são também fornecidos.

Identification, lectotype, Meliponini , morphology, stingless bee, taxonomy

The genus Scaura Schwarz, 1938 is composed of small and dark bees, with unusually enlarged hind basitarsus (wide as or wider than the hind tibia). The genal area is narrower than the compound eye in profile view and the malar space shorter than the diameter of the flagellum (Silveira et al. 2002). The distribution of the genus is broadly Neotropical, from Mexico to southern Brazil (Camargo and Pedro 2013).

Scaura was first described by Schwarz (1938) as a subgenus of Trigona Jurine, 1807, under the name Trigona (Scaura) latitarsis Friese, 1900, based on the enlarged hind basitarsus as the main diagnostic characteristic. Later, Schwarz (1948) included Melipona longula Lepeletier, 1836, in this subgenus, considering it to have two subspecies (originally treated as varieties): T. (S.) longula longula and T. (S.) longula tenuis (Ducke, 1916), based on width of the hind basitarsus, the edentate mandible and the rather triangular hind tibiae. Currently, these three forms are considered distinct species (Michener 1990, Camargo and Pedro 2013, Oliveira 2016), included in Scaura as an individualized genus apart from Trigona. This interpretation was initiated by Moure (1944) when he included three species of Scaura in a list of Peruvian species. Additional recognition characters of Scaura were provided by Moure (1951) who highlighted the absence of yellow spots on body, the malar space narrower than the flagellum diameter, the forewing with acute angle in submarginal cell and the abdomen narrower than the thorax.

In the original description of Trigona latitarsis, Friese (1900) described the female and the male, citing three workers and three males among the type material from Brazil (São Paulo) and Suriname. However, he does not identify the institution in which they were deposited, but he included several males from São Paulo among the syntypes that are deposited in the Hungarian Natural History Museum (HNHM - Budapest, Hungary). The specimen designated by Melo and Costa (2004) as lecotype of Trigona latitarsis is a worker from Jundiaí, São Paulo, which is deposited in the Museum für Naturkunde Berlin (ZMB, Berlin, Germany).

Based on some doubts arising from the study of the type material of Trigona latitarsis, the aim of the present study is to clarify some issues related to the taxonomic identity of Scaura latitarsis, based on the information available in the original description and of records species occurrence. The study also discusses the validity of the lectotype designated by Melo and Costa (2004), adds information on the syntypes, the type locality, morphological characters of the type series specimen, its taxonomic identify and, finally, the designation of a specimen of the type series as the new lectotype.

The type material designated by Friese was studied by the second author of this study, during her trip to the HNHM in 2014. All the specimens deposited in those collections which were identified and labeled by Friese as Trigona latitarsis were studied. Specimens found and identified as belonging to the type series of Trigona latitarsis were borrowed and brought to Laboratório de Bionomia, Biogeografia e Sistemática de Insetos (BIOSIS-UFBA, Salvador, Bahia, Brazil). There, they were studied, measured, and photographed with the use of a stereoscopic microscope Leica M165C, coupled with a digital camera Leica DFC295 and containing the Application Suite V4.1 Interactive Measurements, Montage program.

The material designated by Melo and Costa (2004) as lectotype of Trigona latitarsis (Figs 1–4) was loaned from Museum für Naturkunde Berlin (ZMB, Berlin, Germany) to the Laboratório de Hymenoptera, Instituto Nacional de Pesquisas da Amazônia (INPA, Manaus, Amazonas, Brazil), where it was studied with the use of the same stereomicroscope described above.

Figures 1–4. 

Lectotype of Trigona latitarsis Friese, 1900, designated by Melo and Costa (2004) deposited at the ZMB, invalidated in this study. Lateral view 1 Frontal view of head 2 Lateral view of hind leg and metasoma 3 Dorsal view 4 Scale bars in millimeters.

In the diagnosis below, the symbol “♀” refers to worker, “♂” refers to male, “Q” to queen, “S” to sterna and “T” to terga.

A male deposited in the HNHM is designated as the lectotype, which has a label with the following information: “Brasil, S. Paulo”, “631/156”.

A total of 15 paralectotypes (13 males and 2 worker) of Trigona latitarsis Friese, 1900 is also deposited in HNHM (drawer 78/47), which were studied, validated, and labeled by the second author (Fig. 25): five males, all with labels printed with the name “Brasilia” (corresponding to Brazil), and another label with the number “631/156” handwritten (one specimen also has a label written in black ink with Friese’s handwriting “Trigona latitarsis Friese, 1900”); six males with only a printed label with “Brasilia”; two males with a label handwritten in black ink, with the names "Brasil, São Paulo", also with another handwritten label with the number “631/156”; a worker with a handwritten label with the word “Brasil”, and another label handwritten and numbered “631/365”; a worker with a printed label in which is written “Brasil” and a handwritten label with the numbers “631/365”. Although the majority of syntypes of Trigona latitarsis deposited in the HNHM are males, and as Friese did not mention in the species description the females deposited at “Budapest collection”, there are no doubts that these two workers also belong to type series of the Trigona latitarsis, once the data confer with males labels (especially the number 631, which seems to be the number of the locality of collection – in this case "São Paulo" – or collector, as observed in Trigona latitarsis labels and also in the types labels of other species described by Friese in 1900 – same paper), all specimens are of the same species and had been kept separate together with males. It also important to mention that the putative type specimens are the only specimens of this species deposited at the HNHM. Probably, because they were among a large number of males, which was not usual for species of Meliponini described by him, Friese did not notice the two female specimens (therefore these two female were also marked as paralectotypes). Specimens from Suriname, mentioned by Friese (1900), were not found in the collections of Berlin, Vienna, Munich, Paris, or Budapest, which are collections where most of Friese’s types are deposited. All specimens in HNHM are labeled lectotype and paralectotypes by F. F. de Oliveira.

Figure 25. 

Syntypes of Trigona latitarsis Friese, 1900 deposited at the Entomological Collection of the HNHM.

Material deposited in the ZMB: The worker, designated by Melo and Costa (2004) as lectotype of Trigona latitarsis, does not belong to the types series of this species according to our study, as this specimen has the following label data: “Brasil, Jundiahy, 1899, Schrottky”, “Zool. Mus. Berlin”, “latitarsis” [writing with blue ink pen], “LECTOTYPE, Trigona latitarsis, Friese, 1900, desig. Melo & Costa, 2004”. In his publication, Friese (1900) refers to Jundiaí (a town in São Paulo State, Brazil) in the description of other species, but not in the original description of Trigona latitarsis, where he cites only “São Paulo”. If the type locality was Jundiaí, he would have written the full reference of the label, as he did for the other species described in the same paper, also including the collector’s name (e.g. Trigona schrottkyi – page 386–387: “Brasilia: Jundiaby, 20. Oktoh. 1898 (Schrottky) und Saõ Paulo”). Unfortunately, all type series of Melipona crassipes tenuis Ducke, 1916 that should be in MPEG (Museu Paraense Emílio Goeldi, Pará, Brazil) are lost. Ducke (1916), in the description of this species, refers to specimens from Pará (Itaituba), Amazon (Rivers Javary and Japurá) and Mato Grosso (Tributaries of Tapajóz, collected by G. Kuhlmann). This fact had already been mentioned by Ducke (1916) when he described the geographical distribution of Scaura latitarsis (as “Melipona latitarsis”), where he clearly mentions that, according to Friese, this species occurs (or would be present) in São Paulo de Olivença (p. 28 lines 1 and 2). The true Scaura latitarsis has an elongated abdomen with abundant branched setae on the fifth and sixth terga. The species which has been interpreted as S. latitarsis by several authors for many years including Melo and Costa (2004) has short subtriangular abdomen (Oliveira et al. 2013), with few whitish hairs on the fifth and sixth terga.

It is noteworthy that all Scaura found in ZMB were examined and none of them correspond morphologically to Scaura latitarsis, nor do the label data correspond to those cited in the original description by Friese (1900). Scaura latitarsis: Melo & Costa, 2004 (unjustified lectotype).

Scaura latitarsis has been interpreted over the years by a number of different authors based on specimens from different regions of Brazil, including specimens from São Paulo State in southeastern Brazil, like the lectotype designated by Melo and Costa (2004). However, analyzing the older literature on this species and the type material of Trigona latitarsis (the newly designated lectotype and paralectotypes) deposited in HNHM and cited by Friese in the original description (16 specimens, two workers, and 14 male), we conclude that Trigona latitarsis does not correspond morphologically to what has traditionally been interpreted as Scaura latitarsis, and that these specimens that are traditionally (mis)interpreted as S. latitarsis. The species that have been traditionally (mis)interpreted as S. latitarsis (from São Paulo State and other regions of Brazil, as well as from Bolivia, Colombia, Guyana, Peru, Suriname and Venezuela), including the lectotype designated by Melo and Costa, corresponds to a new species (Nogueira in litt.).

Although the type material of Melipona crassipes tenuis Ducke, 1916 was not found, based on how species has been traditionally interpreted over the years by Ducke himself as Trigona tenuis, and also by Friese, Moure and Camargo, we conclude that the species described as Melipona crassipes tenuis Ducke, 1916 correspond to Trigona latitarsis Friese, 1900 (as junior synonyms) as interpreted in the current article. This conclusion is based on the type series of T. latitarsis and on the original description published by Ducke (1916), where he refers to the thin and elongated metasoma “Abdomen muito tenue” (p. 46), the main character that defines this species gaving rise its name, and which separates it from all the other species of Scaura.

The taxonomic confusion must have been generated by a misinterpretation of the type locality mentioned by Friese in the original description, cites only "Saõ Paulo", referring to São Paulo de Olivença city mentioned by Ducke (1916: 28), and not São Paulo State (southeastern Brazil). Given that Scaura tenuis does not occur in the state of São Paulo, but only in the Amazon region from Colombia, Ecuador, Peru, Suriname, French Guiana, Bolivia and Brazil (Fig. 29), we can conclude that the locality “Saõ Paulo” written on the label of some syntypes and mentioned by Friese in the original description refers to a small town in the Amazonas State, known as “São Paulo de Olivença”, and not the state of São Paulo, as it has been interpreted since 1900 by different Meliponini taxonomists, including Silveira et al. (2002), Camargo and Pedro (2002), Melo and Costa (2004), Camargo and Pedro (2013) and Oliveira et al. (2013).

Figures 26–28. 

Labels of type material deposited at the Entomological Collection of the HNHM. Lectotype of Trigona latitarsis Friese, 1900 (26) Lectotype of Trigona heideri Friese, 1900 (27A, B) Paralectotype Trigona kohli Friese, 1900 (28).

Figure 29. 

Map of South America showing the known geographic distribution records for Scaura latitarsis (Friese, 1900), based on the type material studied here, plus analysis of additional material analyzed from Amazon region and part of the Brazilian Cerrado; Bolivia: El Beni; Brazil: Acre, Amazonas, Pará, Rondônia, Roraima, Mato Grosso; Colombia: Amazonas, Meta; Ecuador: Napo; French Guyana: Cayenne; Peru: Huánuco, San Martín, Madre de Dios, Pasco; Suriname.

It is important to note that the Amazon locality now known as São Paulo de Olivença was founded as a village in 1689 (where a mission was carried out by Spanish missionaries), and, in this period, was known as “São Paulo Apóstolo”. However, in 1708, this village came under the dominion of Portuguese missionaries and had its name changed to “São Paulo dos Cambebas”. With its elevation to city status in 1817, it then received the name of “São Paulo de Olivença”. With the creation of the Solimões district, on June 13, 1884, São Paulo de Olivença became a well-known city in the region, for having been chosen as the administrative center of this new district. Thus, in 1900, when the species in question was described, the village was known and visited by researchers and religious people (BVA 2014). This definition of the type locality as São Paulo de Olivença is a very important fact, and may modify the interpretation of many of syntypes of different species described by Friese.

Another important and relevant fact, which provides strong support for this new interpretation of Scaura latitarsis, can be seen in the original description, when Friese describes the male exemplar as having yellow scape (Fig. 14). This is not the case in those male specimens of the species from São Paulo State that have been interpreted as T. latitarsis (1/3 basal is yellowish brown). Even if the material type of Friese (syntypes) comprised of specimens of two species (that were being interpreted individually and S. tenuis and S. latitarsis), the information provided by the original description leaves no doubt about the validity of the taxon correspondence that had been interpreted as “S. tenuis” as being the same as “T. latitarsis”, according to the original description (compared with syntypes studied here). Even if additional specimens belonging to the syntypes series be found in the future, and these prove that the “T. latitarsis” type series is composed of more than one species, it would not invalidate the new lectotype from Budapest (HNHM), as Friese clearly included males from Budapest Museum in the type series.

Friese emphasized in the original description that the yellow scape occurred only in the male. However, the ventral side of the scape is also yellow, in workers, although the color is restricted to the basal third. As the male scape is completely yellow with a strong tonality, it may have been this difference which Friese wished to emphasize in his description. Consequently, all specimens must have been collected in São Paulo de Olivença, although it was written on the label only “S. Paulo” (as noted in specimens deposited in HNHM). Additional important evidence comes from the fact that Friese does not mention Jundiaí in the description, as he does for other species described in the same article, and whose type material type was collected in Jundiaí (e.g. Trigona schrottkyi – pages 386–387 as mentioned previously). Therefore, the only encountered specimens found that we are confident belong to the type series are those of the HNHM, based on the description and data of the labels affixed to the specimens.

A similar case occurs with the Amazonian species Trigona heideri Friese, 1900 (= Ptilotrigona lurida (Smith, 1854)) described by Friese under the number 39, in the same paper (pages 389 - 390), one page after he described Trigona latitarsis Friese, 1900 (number 34, page 388). The species Trigona heideri was described by Friese based on 14 workers from Brazil (Brasilia: Para (Ducke, 29. Septbr. - 11. Novbr. 1899); S. Paulo, Obidos, Fonteboa, Amazonas), Columbia and Peru (Vilcanota), and the specimens were also deposited also at the HNHM.

Based on the type material from HNHM the specimen chosen by Claus Rasmussen (27.III.2006, in litt.), and designated as lectotype of Trigona heideri Friese, 1900, is a specimen from “S. Paulo”, and, like Trigona latitarsis Friese, 1900 has the same handwritten label in black ink (with the same handwriting) (Fig. 27A, B). For this species, Camargo and Pedro (2013) had also assumed that the type locality "S. Paulo" is in fact São Paulo de Olivença (Amazonas, Brazil), as this is an Amazonian species, and one that has never been recorded in São Paulo State (Southeastern Brazil).

An additional and similar case with Trigona kohli Friese, 1900 (=Trigona pallens (Fabricius, 1798)), described by Friese (1900) in the same article (number 29 on page 387), whose paralectotype was designated by Claus Rasmussen from the HNHM type series (Fig. 28). This too is an Amazonian species never recorded in São Paulo State (southeastern Brazil).

Carefully comparing the provenance labels of Trigona latitarsis Friese, 1900 (here designated as lectotype) (Fig. 26), Trigona heideri Friese, 1900 (lectotype designated by Claus Rasmussen) (Fig. 27 A, B) and Trigona kohli Friese, 1900 (paralectotype designated by Claus Rasmussen) (Fig. 28), it is easily seen that the three specimens have the same handwritten label in black ink, written by the same letter, which leads us to assume that they were collected in the same locality and had the labels written by hand by the same person. Also the number 631 appears on observed in the label of Trigona latitarsis (Fig. 26) and Trigona heideri (Fig. 27 A, B), which seems to be the some information about the locality of collection (in this case “São Paulo”), date, or collector.

Based on all information discussed here, we propose the male specimen in HMHN carrying the label written “S. Paulo” to be considered as lectotype of Trigona latitarsis Friese, 1900, replacing the specimen from Jundiaí designated by Melo and Costa (2004) (lectotype invalidated) and Melipona crassipes tenuis Ducke, 1916 and are to be considered a junior synonym of Trigona latitarsis Friese, 1900.

The authors are grateful to Dr. Frank Koch, curator of the Hymenoptera Collection of the Museum für Naturkunde Berlin (Berlin, Germany) and to Dr. Zoltán Vas, curator of the Hymenoptera Collection of the Hungarian Natural History Museum (HNHM, – Budapest, Hungary), for allowing a visit to the collections, made by the second author, and for loaning us the type series and photographs for study. The authors are also indebted to all institutions supporting our laboratories and making this publication possible, especially the financial support for the trip to Europe provided by to the FFO, to study the type-material described here: Food and Agriculture Organization of the United Nations (FAO), Global Environment Facility (GEF), United Nations Environment Programme (UNEP), Fundo Brasileiro para a Biodiversidade (FUNBIO), Ministério do Meio Ambiente (MMA), Fundação de Amparo à Pesquisa do Estado da Bahia (FAPESB), Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) and Universidade Federal da Bahia (PRODOC Project). The authors are also grateful to Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES), by the scholarship to the first author and to Instituto Nacional de Pesquisas da Amazônia (INPA), for making possible this publication. MLO is scholarship of CNPq, Brazil (306100/2016–9). Referees are thanked for suggestions that improved the current article. Adrian Barnett helped with the English.