Stingless robber bees of the genus Lestrimelitta in Colombia (Hymenoptera: Apidae:

Stingless bees (Apidae: Meliponini) are culturally and economically important bees and they represent a major component of the Colombian melittofauna. However, the bee fauna of Colombia is still in early stages of exploration and species identification is often difficult or impossible. We revised the species of the cleptobiotic stingless bee genus Lestrimelitta in Colombia and recognized the following eleven species: L. glabrata , L. guyanensis , L. rufa , L. rufipes , and L. spinosa , which are recorded for the first time for the country; L. huilensis , L. opita , L. piedemontana , and two new species, L. diminuta , sp. n. and L. galvisi sp. n. We also confirmed the presence of L. limao in the Colombian Amazon and provide new geographical records with an updated key to species from Central America and northern South America.


INTRODUCTION
Stingless bees (Hymenoptera: Meliponini) are social, honey-making bees that represent about 2.5 % of the global melittofauna. Yet, they are among the most ecologically, economically, and culturally significant bees in tropical and subtropical areas of the world. Indigenous and non-indigenous human populations use stingless bees and their products (honey, pollen, propolis, etc.) for diverse purposes including food, medicine, and crafts (Gonzalez et al. 2018). However, the identity of many species of stingless bees is problematic and species identification is often difficult or impossible. The purpose of this work is to address the taxonomy of the stingless bee genus Lestrimelitta (Apidae: Meliponini) in Colombia. These bees are obligate robbing or cleptobiotic bees widely distributed in the Neotropical region, which despite having nests of their own, they steal food and nest materials from other stingless bees, such as species of Nannotrigona, as well as honey bees (Portugal-Araújo 1958, Sakagami et al. 1993. Lestrimelitta consist of 24 species, with its greatest diversity occurring in Brazil Pedro 2007, Gonzalez andGriswold 2012). Until recently, records of this genus in Colombia consisted of L. limao (Smith, 1836) only.
However, taxonomic studies during the last two decades documented morphological features reliable in species identification and demonstrated the existence of multiple species under this name (e.g., Ayala 1999, Oliveira 2002, Melo 2003, Oliveira and Marchi 2005, Marchi and Melo 2006, Camargo and Pedro 2007, Gonzalez et al. 2010, Gonzalez and Griswold 2012. Thus, previous records of L. limao in Colombia remain doubtful and some of them might correspond to unrecognized species. As an attempt to reveal the identity of the species of Lestrimelitta from Colombia, Gonzalez et al. (2010) and Gonzalez and Griswold (2012) recognized and described three new species: L. huilensis Gonzalez and Griswold, 2012, L. opita Gonzalez andGriswold, 2012, andL. piedemontana Gonzalez andRasmussen, 2010. However, because their work did not include an exhaustive study of material from Colombia, other species are thus likely to occur. Herein we confirm the presence of L. limao in Colombia, record five additional species of this genus for the country, and describe two other species as new. Thus, this work increases to eleven the number of species of Lestrimelitta in Colombia. We also provide new geographical records and an updated key to species from Central America and northern South America.

MATERIAL AND METHODS
Morphological terminology and measurements (e.g., interalveolar, alveolocular, ocellocular distances, etc.) follow Michener (2007), except for torulus instead of antennal alveolus. We prefer to use torulus because it is in broader application across Hymenoptera. Measurements were taken using an ocular micrometer on a Leica® MZ12 stereomicroscope. We took photomicrographs with a Leica MC 170HD camera attached to a Leica M205A stereomicroscope. We used the abbreviations S, T, and OD for metasomal sterna and terga, and median ocellar diameter, respectively. The symbol for female and the word itself refers to the worker caste, not the queen.
To map the distribution of each species, we used the geographical coordinates associated with specimen labels and, for records that did not have exact geographical information, we used Google Earth (Google, Mountain View, CA, USA) to acquire their coordinates. We assembled 36 occurrence records for all species and generated maps using Sim-pleMappr (Shorthouse c2010). We reproduced label data as appearing on the labels attached to the same specimen. We separated information on different labels by a single slash (/) and indicated annotations to clarify information in square brackets. To avoid repetition of label data, we used "ut supra" (as above) instead of "idem" to indicate the same information in the preceding label. The Latin term "idem" is more broadly used to replace the name of a male author in academic texts, although it has also been used in taxonomic works to avoid repetition of label data.

SYSTEMATICS
Diagnosis: This species resembles L. nana Melo, 2003 in the small body size (head width, ~1.6 mm), pro-and mesotibiae without erect setae on outer surfaces, mesoscutellum flattened and not covering metanotum medially, and mesotibial spur normal (Fig. 1). It can be separated from that species by the following combination of features: propodeal spiracle elongate (5.9× longer than wide), pronotal lobe and mesoscutum with erect setae, mesepisternum with decumbent setae, terga with erect or semi-erect setae at least laterally, and mandible with a single tooth on apical margin. In L. nana, the propodeal spiracle is much broader, 3.0× longer than wide, the mesoscutum, mesepisternum, and terga are glabrous (except on T6), and the mandible is bidentate. Lestrimelitta diminuta shares with L. monodonta Camargo and Moure, 1989 the mandible with a single apical tooth, the propodeal spiracle elongate, and vertex, mesoscutum, mesepisternum, and terga with setae. However, L. monodonta is larger (head width: 2.2 mm vs. 1.6 mm), the setae on mesepisternum are long and erect (short and decumbent in L. diminuta), the mesotibial spur is reduced (long in L. diminuta, 1.2× mesobasitarsal width basally), and the mesoscutellum is elevated, not flattened and partially covering metanotum medially (mesoscutellum flattened, not covering metanotum medially in L. diminuta).
Integument smooth and shiny between minute, sparse punctures as in other species of the genus; sterna weakly lineolate-imbricate.
Etimology. The specific epithet, diminuta, refers to the small body size of this species.
Comments. This species is known from the type locality in the Colombian Amazon (Fig. 2). Marchi and Melo (2006) indicated that some specimens of L. monodonta from Pará (Tucuruí and Jacareacanga) and Maranhão (Urbano Santos), Brazil, are smaller than are specimens from other localities. These authors did not mention the size of these specimens, but they might be confused with L. diminuta. However, Marchi and Melo (2006) were not able to find significant differences in other features besides body size with the typical specimens of L. monodonta. Thus, small specimens of L. monodonta can be separated from L. diminuta by the same features indicated in the key to species. Diagnosis. This species can be easily recognized by the following combination of features ( Fig. 3): propodeal spiracle elongate; vertex, preoccipital margin, anterior margin of mesoscutum, and terga with erect, long setae; propodeum laterally with short, sparse, simple setae; and mesotibial spur reduced. This species runs to L. huilensis in the key to species of Lestrimelitta from Central America and Northern South America by Gonzalez and Griswold (2012). It differs from that species in the mesotibial spur reduced (1.3× mesobasitarsal width basally in L. huilensis) and mesepisternum and disc of mesoscutum with short but distinct, decumbent setae (glabrous or nearly so in L. huilensis). In addition, both species are geographically separated and inhabiting different ecosystems. Lestrimelitta galvisi occurs in tropical rain forests of the Choco bioregion whereas L. huilensis in dry tropical forests along the Magdalena valley in Colombia.
Etymology. This species is a patronymic honoring Germán Galvis Vergara, friend and colleague from the Department of Biology, Universidad Nacional de Colombia, Bogotá, in recognition for his efforts in documenting and protecting the biodiversity of Colombia.
Additional material examined. Two females with the same data as holotype but with catalogue numbers 14439 and 14422 (not designated as paratypes). One of them is a callow specimen judging by its body yellow coloration (14439). The other specimen has a normal adult coloration but all legs and metasoma are missing. Both specimens are in LABUN. tially covering metanotum medially; propodeal spiracle elongate, about 4.7× longer than wide; mesotibial spur reduced; metatibia about 3.0× longer than broad.
Integument smooth and shiny between minute, sparse punctures as in other species of the genus; terga and sterna weakly lineolate-imbricate.

Lestrimelitta opita
huilensis along the Magdalena valley, reaching as far north as Sucre, in the Caribbean region of the Colombia (Fig. 4).
Specimens of this species have been collected from 170 to 1765 m. Gonzalez and Rasmussen, 2010 Lestrimelitta piedemontana Gonzalez and Rasmussen, in Gonzalez et al. 2010: 320 [♀].

Lestrimelitta rufa
Comments. This species is currently known from Bolivia, Brazil, and Ecuador (Gonzalez and Griswold 2012). A specimen from Peru in the SEMC, with the following label data, has been identified as L. limao: Peru. Junin: Dept. Villa-Oxapampa Rd., 1050 m, 10°48'36" S, 75°19'54" W, 18 OCT 1999, R. Brooks, PERU 1B99 053, ex. Under stones along creek / SM0144188. However, this specimen belongs to L. rufa and thus, this species is newly recorded from Peru. The records provided here expand the geographical range of the species. Lestrimelitta rufa occurs in the Amazon region of Colombia (Fig. 2). (Friese, 1903) Trigona (Lestrimelitta) limao rufipes Friese, 1903 Comments. This species is widely distributed in Brazil (Marchi and Melo 2006). Gonzalez et al. (2010) recorded it from the Province of San Martín, in northern Peru. The record from Colombia in the Orinoquia region represents the northwestern-most locality for this species (Fig. 2). 2006), only a few have assessed their taxonomy. Thus, species identification remains difficult and in many cases impossible. To date, only the Colombian species of Duckeola Moure (Gonzalez and Nates-Parra 2004), Geotrigona Moure (Gonzalez andSepúlveda 2007, Gonzalez andEngel 2012), Parapartamona Schwarz (Gonzalez and Nates-Parra 1999), Paratrigona Schwarz (Gonzalez and Vélez 2007), and Oxytrigona Cockerell (Gonzalez 2007, Gonzalez andRoubik 2008) have been revised. Such works, including ours, have documented several new species and clarified the identity and distribution of many others.

Lestrimelitta rufipes
In comparison to other stingless bee genera, specimens of Further works on the Colombian species of Lestrimelitta should try to elucidate the identity of the species from the Caribbean coast that we have tentatively identified as L.
guyanensis. In addition, the species status of L. piedemontana requires revision, as this species might be conspecific with L. rufa given that both species are morphologically similar (Gonzalez et al. 2010). Lestrimelitta spinosa Marchi and Melo, 2006 Lestrimelitta spinosa Marchi and Melo, 2006: 25 [♀].

DISCUSSION
The true diversity of stingless bees in Colombia is at most a guess, with available literature records differing in the number and identity of the species occurring in the country (e.g., Nates-Parra 2006, Pedro 2007, Ascher andPickering c2018). Although many studies summarize various aspects of the nesting biology, management, and local uses of stingless bees in Colombia (see Nates-Parra 8(7