A checklist of Euglossini in Ecuador is given, including all currently described, valid species collected until 2018. The list has been assembled from museum records, fieldwork cited herein, and literature. The former species lists are nearly doubled here, with 1 Aglae, 23 Eufriesea, 68 Euglossa, 18 Eulaema, and 5 Exaerete, 115 in total with >50 new records for the country. Distribution and collection data are included, and some doubtful species are discussed. The Amazon region is the most species rich area but not necessarily a natural pattern, perhaps due to uneven sampling effort across the country. Southern Ecuador is relatively little sampled.
1. Introduction
Orchid bees are a conspicuous and a fascinating group of Neotropical bees, in which males have noteworthy behavior related to gathering aromatic compounds from orchids and other sources [1]. The bees increasingly are objects of study in different fields such as ecology [2, 3], chemical ecology [4–8], conservation [9–11], evolution [12], and taxonomy [13–17]. Such research contributes to an understanding of general bee biology and how neotropical ecosystems function.
More than 252 species have been described, and many synonymized, since early descriptions by Linneaus (1758) under the genus name Apis [18]. Several species lists have been published at regional levels [18–21] and the country level, Brazil [19, 21–28], Peru [19, 29, 30], Panama [19, 31, 32], Bolivia [33], Colombia [19, 34], Mexico [35], Nicaragua [36], Honduras [37], and Venezuela [19].
Ecuador is well known as a megadiverse country with an outstanding diversity of organisms and ecosystems and considered important for conservation proposes, e.g., the Tumbes-Choco-Magdalena region and the tropical Andes [38, 39]. At the same time, from the species records examined here and from biogeographic analyses of other apidologists, e.g., [13, 40], it is clear that the NW of Ecuador is quite similar to Central America, while Amazonian Eastern Ecuador and the Andes have some different species. Additionally, among the well-studied Ecuadorian palms, although most species are Andean compared to other regions, 60% of them are from adjacent lowlands; i.e., they are not Andean endemics [41].
Here we update the taxonomic base of knowledge on Ecuadorian euglossines. Only two publications had reported, in general, species for Ecuador. Ramírez et al. [19] report 62 species and Moure et al. [18] report 63. Recent articles treat the taxonomy of the orchid bees of Ecuador, mainly focusing on the description of new species, Euglossa samperi [42]; Euglossa natesi [43]; Euglossa williamsi [44], Euglossa celiae, and Euglossa subandina [45]. As part of our continuing study of euglossines in Ecuador and elsewhere, while not listing unnamed species here (as for Meliponini, see [46], or Euglossini, [47]) we attempt to bring euglossine taxonomy up to date, particularly in western South America, and include limited discussion of taxonomic problems.
2. Methods2.1. Material Examined
All material examined from museums came from the following institutions:
INABIO: Instituto Nacional de Biodiversidad, Quito-Ecuador.
MZUA: Museo de Zoología de la Universidad del Azuay, Cuenca- Ecuador.
NMNH: National Museum of Natural History, Smithsonian Institution, Washington DC - USA.
QCAZ: Museo de Zoología de la Pontificia Universidad Católica del Ecuador, Quito-Ecuador.
STRI: Smithsonian Tropical Research Institute (DW Roubik collection), Balboa, Panama.
Additional data came from the following institutions:
AMNH: Amercian Museum of Natural History, New York, USA.
DZUP: Departamento de Zoología, Universidad Federal do Paraná, Curitiba, Brasil.
FLMNH: Florida Museum of Natural history, Gainesville, USA.
Several locations in the country had been visited in most of the provinces of Ecuador during the last decades, mainly by DW Roubik and also by PS Padrón and RB Picón (Figure 1). Orchid bees were collected in bait traps or collected using hand nets at baiting stations, using chemical attractants such as cineole, eugenol, skatole, vanillin, PDMB (p-dimethoxybenzene), methyl salicylate, and methyl cinnamate, during the day at different periods between 08,00 and 17,00 h.
Species richness pattern observed for the Euglossini tribe in Ecuador based in 2332 collecting records.
The species list is presented by subgenus and species following the systematic arrangement proposed by Moure et al. [18], so it does not include the recently named subgenus of Euglossa, Alloglossura, which will be subject to further evaluation [17], nor a proposed name Hybomelissa for the Exaerete group containing E. frontalis [48]. The species determinations were based upon comparative study of type and other material at NMNH and in the collections of DW Roubik and RL Dressler, at STRI and at the Florida Museum of Natural History, particularly collections made by DW Roubik from French Guiana, Panama, Costa Rica,l and Ecuador. Diagnostic reference photographs also were consulted for euglossines of French Guiana (Discover Life, https://www.discoverlife.org/mp/20p?see=I_DWR, and the NMNH euglossine types, https://biogeodb.stri.si.edu/bioinformatics/roubik_bees/). We attempt to treat determinations made by others as likely correct, unless we had additional information, and acknowledge that some of the above cited reference material, or identifications, may be erroneous.
2.3. Species Richness
A total of 2322 distribution records were georeferenced, localities were plotted using DIVA-GIS 7.5 [49], and the species richness was calculated.
3. Results
The preliminary checklist presented here contains 115 species distributed among the genera, 1 species in Aglae, 68 species in Euglossa, 23 in Eufriesea, 18 species in Eulaema, and 5 species in Exaerete. The list is arranged by genera and subgenera (when formally designated), and species are listed alphabetically, with specimen label information or the literature references. For each label or collection, available information included types, number of specimens, province, locality, elevation, latitude, longitude, date of collection, collector, museum, and reference.
Checklist of Orchid Bees from Ecuador
Tribe Euglossini Latreille, 1802
Genus Aglae Lepeletier & Serville (1825)
Aglae coerulea Lepeletier & Serville, 1825
Records: 1, Napo, Portoyacu, 700 m, D. Velastegui, INABIO; 1, Pastaza, QCAZ; 1, Pastaza, Villano Taranguro, 342 m, 1 22.6670 S, 77 22 91 O, 35345, J. Naranjo, QCAZ; 1, Esmeraldas, Rìo Cananda, 400 m, 0 28 45, 77°11 51 O, QCAZ; 1, Napo, Avila Viejo, 750, QCAZ; Esmeraldas, Choco Region, [50]; 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Genus Eufriesea Cockerell (1908)
Eufriesea buchwaldi (Friese, 1923)
Records: [18].
Eufriesea auripes (Gribodo, 1882)
Records: 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Eufriesea chrysopyga (Mocsáry, 1898)
Records: [18]; 1, QCAZ; 17, Orellana, Yasuní, QCAZ; 1, Orellana, Est. Río Huiririma, 220 m, 0 38 LS 75 54 LW, QCAZ; 1, 500 m, QCAZ; 1, Morona Santiago, Wapula, 1380 m, -2.126813, -78.089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA.
Eufriesea combinata (Mocsáry, 1897)
Records: Morona Santiago, Res. Biologica El Quimi, 1671 m, McPhail trap, Julio 2017, A. Suárez-Torres. INABIO.
Eufriesea concava (Friese, 1899)
Records: Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, STRI.
Eufriesea pretiosa (Friese, 1903)
Records: Holotype, Tungurahua, Santa Inez, Ecuador, AMNH, [13]; Tungurahua, Santa Inez, [13]; Tungurahua, Santa Inez, [19, 54]; Tungurahua, Santa Ines, [18, 51].
Eufriesea pulchra (Smith, 1854)
Records: Pastaza, Puyo km 35 to Tena, 960 m, 29/02/1980, N.H. Williams NMNH; 57, Orellana, Yasuní, 280m, QCAZ; [19, 55]; 6, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Eufriesea purpurata (Mocsáry, 1896)
Records: 2, Napo, Limoncocha (forest), 25/07/1971, D. L. Pearson, NMNH; 6, Orellana, Yasuní, 250 m, QCAZ; 6, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Eufriesea superba (Hoffmannsegg, 1817)
Records: Los Ríos, Palenque Station, April, [13, 19]; Los Rios, [18, 56].
Eufriesea surinamensis (Linnaeus, 1758)
Records: 1, Guayas, Guayaquil, 43010, C. Dodson, NMNH; 2, Guayas, Guayaquil, NMNH; 14, Guayas, Guayaquil, 18/03/1960, G. Frymire, NMNH; 14, Guayas, la Lolita, 10/02/1960, G. Frymire, NMNH; 4, Los Ríos, Quevedo, 22010, C. Dodson, NMNH; Los Ríos, Montalvo, 28/02/1968, G. Frymire NMNH; Los Ríos, Vic. Quebrada, march abril 1955, E. N. O’ Rourike, NMNH; Guayas, Guayaquil, [13]; 1, 500 m, QCAZ; [18, 19].
Eufriesea theresiae (Mocsáry, 1908)
Records: [19]; Zamora, Río Zamora, [18, 51].
Eufriesea venezolana (Schrottky, 1913)
Records: Los Rios, Palenque River, [13, 18]; Guayas, Guayaquil, [13].
Genus Euglossa Latreille (1802)
Subgenus Euglossa Latreille, 1802
Euglossa amazonica Dressler, 1982
Records: 3, Orellana, Yasuní, QCAZ; 2, Orellana, Coca, 700 m, 79 02 27 W 00 18 21 S, QCAZ; 2, Morona Santiago, Chiwasa, LW 79 05 LN 00 05 18, QCAZ; 1, Morona Santiago, San Juan Bosco, 1050 m, -3,119898, -78,527502, II/2016, H. Garzón, MZUA; 1, Morona Santiago, Wapula, 1380 m, -2,126813, -78,089803, 25/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; 42, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: [19]; 1; Cañar, Ocaña, 600 m, 11-I-06, S. Padrón, MZUA; [57].
Euglossa charapensis Cockerell, 1917
Records: West of Ecuador, [58]; 1, 1601 m, QCAZ; [19]; 1, Loja, Malacatos, 1450 m, -4,224100, -79,262287, 31/XII/2016, P. Picón, S. Padrón, MZUA; 1, Morona Santiago, Wapula, 1360 m, -2,125181, -78,088571, 18/V/2017, S. Padrón, MZUA.
Euglossa chlorina Dressler, 1982
Records: 1, Pichincha, 2000 m, 79 00 00 W 00 25 25 S, QCAZ; 1, Pichincha, Patricia Pilar, QCAZ; 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa cognata Moure, 1970
Records: 6, QCAZ; 2, Napo, Yuturi Lodge, Río Napo, 359 m, 79 12 W 00 28 N, QCAZ; [19]; Orellana, Yasuní, [59]; Napo, [59]; Alto Tambo, [18, 59]; 15, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa cordata (Linnaeus, 1758)
Records: [18].
Euglossa cybelia Moure, 1968
Records: [19].
Euglossa deceptrix Moure, 1968
Records: Ibarra (?), 1897, leg. W. F. H. Rosenberg, [60]; 1, El Oro, Piñas, 960 m, 26.2.-2.3.1982, leg. N. H. Williams, det. R. L. Dressler, [60]; 2, Lita, 17.8.1987, leg. det. R. L. Dressler, [60]; 1, Paramba, 3.1897, leg. F. H. Rosenberg, det. Bembé (Eg. cordata, det. Friese) [60]; 1, ZSM, [60]; Esmeraldas, Bilsa Biological Reserve (“Bilsa” hereafter), 0°20,816' N 79°42,659' W, [47]; 1, QCAZ; 4, Esmeraldas, Mache-Chindul Ecological Reserve, August to December 2014, [47].
Euglossa despecta Moure, 1968
Records: 1, Esmeraldas, Pitzara, QCAZ; 8, Esmeraldas, Mache-Chindul Ecological Reserve, August to December 2014, [47]; 14, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa dissimula Dressler, 1978
Records: 12, Esmeraldas, Mache-Chindul Ecological Reserve, August to December 2014, [47].
Euglossa dressleri Moure, 1968
Records: [19]; 18, Esmeraldas, Mache-Chindul Ecological Reserve, August to December 2014, [49].
Euglossa gaianii Dressler, 1982
Records: 1, Guayas, San Joaquin, QCAZ.
Euglossa gibbosa Dressler, 1982
Records: 1, Holotype, Pichincha, Santo Domingo, 20/07/1967, R. L. Dressler, NMNH; 1, Allotype, Pichincha, Santo Domingo, 20/07/1967, R. L. Dressler, NMNH; 1, Pichincha, Palenque, QCAZ; 2, Pichincha, Santo Domingo - Quevedo, 220 m, 0 38 LS 75 54 LW, QCAZ; 3, Pichincha, Patricia Pilar, 389 m, 79 12 W 00 28 N, QCAZ; 2, Los Ríos, Río Palenque, 218 m, 75 52 W 00 36 S, QCAZ; 1, Esmeraldas, Tonsupa, 517 m, 01 28.434 S 77 31.868 W; QCAZ; Santo Domingo, [59]; Pichincha, [18]; Santo Domingo, [18]; 1, Manabí, Jama Coaque, 300 m, -0,107904, -80,122931, 02/XII/2016, P. Picón, S. Padrón, MZUA; [49]; 2, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa hansoni Moure, 1965
Records: 2, Pichincha, Santo Domingo, [61]; 15, Esmeraldas, Mache-Chindul Ecological Reserve, [61]; 21 Aug. 1967, [62]; 3 Jan. 1970, H. A. Hespenheide (Dressler coll.), [62]; 2, Pichincha, Santo Domingo, 22/07/1967, R. L. Dressler, NMNH; 15, Esmeraldas, Bilsa Biological Reserve (“Bilsa” hereafter), 0°20,816' N 79°42,659' W, [47].
Euglossa hemichlora Cockerell, 1917
Records: 1, Holotype, Paramba, Ecuador; NMNH, [53]; 3, Pichincha, Santo Domingo, 20/07/1967, R. L. Dressler, NMNH; 4 Feb. 1971, H, Kennedy (Dresller coll. and to be distributed, [62]; 13, Pichincha, Santo Domingo, det. Bembé (Eg. heterosticta, det. R. L. Dressler), DZUP, [60]; 14, Pichincha, Santo Domingo, 20.-22.7.1967, leg. R. L. Dressler det. Bembé, DZUP, [60]; Pichincha, Santo Domingo, 22/07/67, R. L. Dressler, [63]; Pichincha, Santo Domingo, [18, 45]; 1, Loja, Cabeza de Toro, 309 m, -80,30235, -4,294143, 26/VIII/2017, J. Falcón, J. Fajardo, P. Picón, S. Padrón, MZUA; 1, Guayas, Guayaquil Espol, 100 m, -2,14829, -79,965824, 16/XI/2016, P. Picón, S. Padrón, MZUA; 19, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: 1, D. Roubik, NMNH; 3 males, Napo, Río San Miguel, NMNH; 27/04/1988, D. Roubik, NMNH; 2, Napo, Tena Shushufindi, QCAZ; 1, Orellana, Coca, QCAZ; 4, Orellana, Yasuní; QCAZ; Napo, [18]; 1, Morona Santiago, Wapula, 1380 m, -2,126813, -78,089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; 28, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa nigropilosa Moure, 1965
Records: Bolivar, Balzapamba, NMNH; 20/07/1967, R. L. Dressler, NMNH; Tunguragua, 700 - 1400 m, NMNH; Tungurahua, Topo, 700 –1400 m, NMNH; 1, Tungurahua, Río Verde, 600 m, 78 27 18 W 00 50 24, QCAZ; 1, Cotopaxi, Las Pampas, 250 m, 00 56 S 75 24 W, QCAZ; 1, Pichincha, 250 m, 00 56 S 75 24 W, QCAZ; Bolivar, Balzapamba, 16/01/1960, C. Jativa, NMNH; Tunguragua, 700 - 1400 m, Dr. C. H. Dodson, [65]; Tungurahua, [18]; 1, Morona Santiago, El Paraíso, 1630 m, -3,299298, -78,589826, 05/XII/2015, S. Padrón, MZUA; 1, Morona Santiago, Wapula, -2,126813, -78,089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; 1, Morona Santiago, Wapula, 1380m, -2,126813, -78,089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA.
Euglossa platymera Dressler, 1982
Records: 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa pleosticta Dressler, 1982
Records: 2, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa retroviridis Dressler, 1982
Records: 2, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: 4, Pichincha, Santo Domingo, 20/07/1967, R. L. Dressler, NMNH; 1, 250 m, QCAZ; 1, Pichincha, Patricia Pilar, QCAZ; 4, Pichincha, Santo Domingo, [18, 19]; 1, Loja, Cabeza de Toro, 309 m, -80,30235, -4,294143, 26/VIII/2017, J. Falcón, J. Fajardo, P. Picón, S. Padrón, MZUA; 2, Manabí, Jama Coaque, 300 m, -0,107904, -80,122931, 02/XII/2016, P. Picón, S. Padrón, MZUA.
Euglossa variabilis Friese, 1899
Records: 1, Paramba, NMNH; April 1998, D. W. Roubik, [66]; 7, Pichincha, Santo Domingo, DZUP, [60]; Guayas, 1, Guayaquil, leg. Buchwald, (Eg. cordata, det. Alfken 1904), det. Dressler, [60]; 1, El Oro, 16 km SSE Buenavista, 16.4.1989, leg. Dressler, Whitten and Williams, det. Bembé, [60]; 1, Los Rios, Río Palengue Station, 5.4.1972, leg. C. H. Dodson, MC, det. Bembé, [60].
Subgenus Euglossella Moure, 1967
Euglossa apiformis Schrottky, 1911
Records: Zamora, 5-7/III/1982, N. H. Williams 89 [handwritten on the underside], [45]; Zamora Chinchipe, [18].
Euglossa bigibba Dressler, 1982
Records: 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ; 1, Orellana, Yasuní, March 2018, D.W. Roubik.
Euglossa cyanura Cockerell, 1917
Records: [18]; 2, Esmeraldas, Mache-Chindul Ecological Reserve, August to December 2014, [47].
Euglossa granti Cheesman, 1929
Records: 1, Imbabura, Lita, 320 m, QCAZ; 16 Aug. 1972, Dressler, 1982; 13-27 April 1998, D. Roubik; coll. No 33.
Euglossa perfulgens Moure, 1967
Records: 1, Orellana, Yasuní, 220 m, 0 38 LS 75 54 LW, QCAZ.
Records: 1, Paratype, Esmeraldas, km 17 Lita-Alto Tambo, 730 m, 19/05/1976, J. Cohen, NMNH; Esmeraldas, [18].
Euglossa occidentalis Roubik, 2004
Records: 1, Holotype, Napo, Yasuní National park, 13-27 April 1998, D. Roubik, QCAZ, [68]; 12, Paratypes, Orellana, Parque Nacional Yasuní, Sept. 2001, E. Baús D. Roubik, QCAZ, [68]; 3, Paratypes, Napo, Yasuní National park, 13-27 April 1998, D. Roubik, QCAZ, [68]; 16, Paratypes, Orellana, PUCE SC Yasuní, 250m, 76°24’19” W 00°40’32 S, 18- 23 Feb. 2001, D. Roubik & E. Báus, QCAZ, [68]; Orellana, Parque Nacional Yasuní, nov. 1998, E. Báus D. Roubik, QCAZ, [68]; Orellana, Shushufindi Yasuní, 500 m, 76°30’W 00°38’ S, 3 Aug. 1999, F. Palomeque, QCAZ, [68]; 2, Paratypes, Orellana, Loreto Cotapino, 640 m, 22 May 1999, F. Palomeque, QCAZ, Donoso, et al., 2009; Napo, Talag, 600 m, W 77°54’ S01°03’, 12 Jun 99, H. Zumárraga, QCAZ, [68]; 2, Paratypes, Orellana, Parque Nacional Yasuní, dic. 2001, E. Báus D. Roubik, QCAZ, [68]; 17, Paratypes, Orellana, Parque Nacional Yasuní, dic. 2002, E. Báus D. Roubik, QCAZ, [68]; Orellana, E.C. Yasuní, 250 m, 00°40’S 76°23’W, 20 Nov 1999, L. Torres, [66]; Paratype, Orellana, Yasuní Universidad Estación, 36923, D. Roubik, NMNH; 18 Jan. 1990, [43]; 1, Orellana, Yasuní, 250 m, QCAZ; Napo, [18]; 15, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa orellana Roubik, 2004
Records: 1, Holotype, Napo, Yasuní National Park, 13-27 April 1998, D. Roubik, QCAZ, [68]; 132, Paratypes, Orellana, Yasuní National Park, 13-27 April 1998, D. Roubik, QCAZ, [68]; 7, Paratypes, Orellana, Tena Shushufindi Yasuní, 500 m, 76°30’ W 00°38’S, 03 Aug. 1999, F. Palomeque, QCAZ, [68]; Napo, Tena Misahualli Jatun Sacha, 550 m, 77°30’W 01°03’S, 23 Oct. 1999, P. Carrera, QCAZ, [68]; 5, Paratypes, Napo, E.C. Yasuní, 250 m, LW78°58’ LS00 56, 22 Apr. 1998, F. Palomeque, QCAZ, [68]; 2, Paratypes, Napo, Loreto, 9 Aug. 1991, D. Roubik, QCAZ, [68]; 189, Paratypes, Orellana, PUCE SC Yasuní, 250 m, 76°24’19” W 00°40’32 S, 18-23 Feb. 2001, D. Roubik and E. Baus, QCAZ, [68]; Pichincha-Napo, Taracoa S. Abedravo, 18-V-84, QCAZ, [68]; 2, Paratypes, Napo, Yuturi Lodge Río Napo, 0°32’54”S 76°2’18” W, 270 m, 20 Mar. 1999, R. Brooks, QCAZ, [68]; 108, Paratypes, Orellana, Parque Nacional Yasuní, dic. 2002, E. Baus D. Roubik, QCAZ, [68]; 49, Paratypes, Orellana, Parque Nacional Yasuní, Sept. 2001, E. Baus D. Roubik, QCAZ, [68]; 47, Paratypes, Orellana, Yasuní Nat. Park Catholic Univ. Station, Aug 7-17 2004, D. Roubik, QCAZ, [68]; 5, Paratypes, Orellana, Yasuní Universidad Estación, 27/04/1988, D. Roubik, NMNH; 100, Paratypes, Orellana, Yasuní, QCAZ; Pastaza, Tewaeno 32km sw of Tzapino, 19/05/1976, J. Cohen, NMNH; Napo, Yasuní National Park, [18]; 2, Morona Santiago, Wapula, 1380 m, -2,126813, -78,089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; 1, Morona Santiago, Wapula, 1360 m, -2,126813, -78,089803, 03/VI/2016, C. Maldonado, S. Padrón, MZUA; 14, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa rufipes Rasmussen & Skov, 2006
Records: Napo, [18]; Pastaza, [18]. Río Hollín, D. W. Roubik, 2018, QCAZ.
Euglossa tiputini Roubik, 2004
Records: 1, Holotype, Orellana, Yasuní National Park, [19]; 2, Paratypes, Napo, Hacienda Ila, D. Velastigui, [19]; Paratypes, Napo, Hacienda Ila, 12-26-68, D. Velastegui, QCAZ, [68]; Napo, Talag, 400 m, 28 Dic. 1993, O. Torres, QCAZ, [66]; Napo, Yasuní National Park, [18].
Euglossa viridifrons Dressler, 1982
Records: Paramba, 13–27 April 1998, D. Roubik, QCAZ, [66]; 1, Napo, Limoncocha, Feb. Apr. (1998, 2001), D. Roubik, [66]; Napo, [18]; 2, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Subgenus Glossurella Dressler, 1982
Euglossa augaspis Dressler, 1982
Records: 4, Esmeraldas, 20 m, LW 78 59 39 LN 00 47 30, QCAZ; 1, Morona Santiago, Wapula, 1380 m, -2,126813, -78,089803, 26/II/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; 11, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: [19]; 1, Manabí, Jama Coaque, 300 m, -0,107904, -80,122931, 02/XII/2016, P. Picón, S. Padrón, MZUA; 65, Esmeraldas, Mache-Chindul Ecological Reserve, August to December 2014, [47].
Euglossa fuscifrons Dressler, 1982
Records: 1, Holotype, Napo, Vera Cruz, 03/II/1969, D. Velastegui, NMNH, Dressler 1982; 2, Orellana, Loreto Cotapino, 220 m, 0 38 LS 75 54 LW, QCAZ; 1, Napo, Talac, 938 m, 77 69 LW 0128 LS, QCAZ; [19, 51]; Napo, [18]; 1, Morona Santiago, Wapula, 1380 m, -2,126813, -78,089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; Napo, Río Hollin, 14.I.2012, [17]; 31, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa gorgonensis Cheesman, 1929
Records: Esmeraldas, Mache-Chindul Ecological Reserve, [59]; 1, Manabí, Jama Coaque, 300 m, -0,107904, -80,122931, 02/XII/2016, P. Picón, S. Padrón, MZUA.
Records: 1, Holotype, Esmeraldas, Bilsa Biological Station, 500 m, 00° 21’ N 79° 44’ W, leg. Santiago Ramírez, QCAZ, [19]; 13, Paratypes, Esmeraldas, Bilsa Biological Station, 500 m, 00° 21’ N 79° 44’ W, leg. Santiago Ramírez, [42]; Donoso, et al., 2009; Esmeraldas, Bilsa Biological Station, 00°21'N 79°44’W, 500 m [18].
Euglossa turbinifex Dressler, 1978
Records: 1, Pichincha, Santo Domingo, 6 Aug. 1968, C. Dodson (Dressler coll.); 2, Pichincha, Santo Domingo, 16 - 18 Feb. 1969, N. H. Williarns (Dressler coll.), [69]; 1, Pichincha, Santo Domingo, emerged from a nest collected by C. Dodson, no date (Dressler coll.), [19, 69].
Euglossa williamsi Hinojosa-Díaz & Engel, 2011
Records: 1, Holotype, Napo, September 1987, Dressler, Wille, Whitten, Williams, FLMNH, [44]; December 26, 1968, [66]; Napo, [18].
Subgenus Glossuropoda Moure, 1989
Euglossa intersecta Latreille, 1817
Records: 53, QCAZ; 88, Orellana, Yasuní, QCAZ; 5, Napo, Tena Misahualli Jatun Sacha, 640 m, QCAZ; 71, Orellana, Yasuní, 804 m, QCAZ; 3, Sucumbios, Cuyabeno, 350 m, QCAZ; 4, Orellana, Estación Chiruisla, QCAZ; 36, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Euglossa rugilabris Moure, 1967
Records: 12, Orellana, Yasuní National Park, Feb., Apr. (1998, 2001), D. Roubik, [66]; Orellana, Yasuní National Park, 13-27 April 1998, Roubik, NMNH; 2, QCAZ; 12, Orellana, Yasuní National Park, [18, 70]; 14, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: 1, Carchi, El Placer, 500 m, 76 30 W 00 38 S, QCAZ; 1, Pichincha, Endesa, 610 m, 77 48 W 00 54 S, QCAZ; 1, Pichincha, San Miguel de los Bancos, 1500 m, 78 17 42 W 01 23 54 S, QCAZ; 1, Pichincha, Santo Domingo, QCAZ; 4, Esmeraldas, Río Canandé, 250 m, 00 56 S 75 24 W, QCAZ; [18, 19].
Eulaema meriana (Olivier, 1789)
Records: 13, Napo, Limoncocha, 14/06/1977, D. L. Vincent, NMNH; Napo, Lago Agrio 2 km N, 26 de Septiembre 1975, A. Langley, NMNH; Pastaza, Puyo, 8 de febrero 1976, Spangler, et al., NMNH; Napo, Limoncocha, 16/08/1972, E. W. Stiles, NMNH; Dressler, 1979; 1, Napo, Tarapa, QCAZ; 1, Campanacocha, QCAZ; 1, Esmeraldas, QCAZ; 86, Orellana, Yasuní, QCAZ; 1, Napo, Yuturi Lodge, Río Napo, QCAZ; 5, Napo, Loreto, QCAZ; 1, Esmeraldas, Quininde, QCAZ; 1, Pichincha, Pedro Vicente Maldonado, QCAZ; 1, Los Ríos, Quevedo, QCAZ; 1, Sucumbios, Cuyabeno, QCAZ; 1, Pastaza, Loracachi, QCAZ; 1, Los Ríos, San Carlos, QCAZ; 1, Napo, Lago Agrio, QCAZ; 45, Napo, 600 m, 1° 02 8 77° 40 W, QCAZ; 1, Orellana, Est. Río Huiririma, 500 m, QCAZ; 1, Pastaza, Via Tena Puyo, 1800 m, QCAZ; 33, QCAZ; [18, 19]; Pichincha, Nanegalito- Reserva Inti Llacta, 1800 m, [73]; 1, Azuay, Sarayunga, 530 m, -3,315218, -79,582313, 10/XI/2016, P. Picón, S. Padrón, MZUA; 1, Morona Santiago, San Juan Bosco, 1050 m, -3,119898, -78,527502, II/2016, H. Garzón, MZUA; 4, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: Napo, Limoncocha, 15 de Junio 1977, P. J. Spangler D. R. Givens, NMNH; 1, Napo, Tena Talag, 269 m, 76 11 S 00 01 S, QCAZ; 1, Napo, Coca Palmoriente, QCAZ; 1, Morona Santiago, Macas, 1000 m, 77 23 W 00 15 N, QCAZ.
Eulaema sororia Dressler & Ospina-Torres, 1997
Records: 1, Holotype, Esmeraldas, Oeste de Lita, 14-18 Agosto 1987, [19, 79]; Esmeraldas, Oeste de Lita, 14-18 Agosto 1987, Dressler, Hills, Whitten, Williams, Leg. KU, [80]; Esmeraldas, [18].
Genus Exaerete Hoffmannsegg (1817)
Exaerete dentata (Linnaeus, 1758)
Records: 2, Guayas, Guayaquil, NMNH; 1, A. Cartyright Donor, NMNH; San Rafael, 100 m oeste, F. Campos, NMNH; 1, Purchased Kny Scheerer Co NY, NMNH; [18, 19]; 2, [80].
Exaerete frontalis (Guérin, 1844)
Records: 1, Cotopaxi, Guasanda, La Maná, 500 m, QCAZ; 1, Orellana, Coca, QCAZ; 1, Napo, Tena Shushufindi, 500 m, QCAZ; 1, Napo, Archidona, QCAZ; 1, Pichincha, P. V. Maldonado Río Pizar, 280 m, QCAZ; 1, Esmeraldas, Borbon, 25 m, 78 58 W 1 4 N, QCAZ; 8, Orellana, Yasuní, 250 m, QCAZ; Los Ríos, Río Palenque Biological Station Quevedo, 14/03/1979, Joseph J. Anderson, NMNH; 1, Napo, Limoncocha, 16/08/1972, E. W. Stiles, NMNH; [18, 19, 59]; 1, Morona Santiago, Wapula, 1380 m, -2.126813, -78.089803, 26/XII/2016, S. Padrón, P. Picón, C. Maldonado, J. Falcón, MZUA; 1, Morona Santiago, San Juan Bosco, 1500 m, -3,119898, -78,527502, 18/VIII/2016, S. Padrón, H. Garzón, MZUA; 1, Manabí, Jama Coaque, 300 m, -0,107904, -80,122931, 02/XII/2016, P. Picón, S. Padrón, MZUA; 6, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
Records: 1, Tena, Shushufindi, Yasuní, 500 m, QCAZ; 1, Guayas, Guayaquil, QCAZ; 2, Napo, Tena Talag, 750 m, QCAZ; 3, Pichincha, Santo Domingo, QCAZ; 1, Esmeraldas, km 6 Lita San Lorenzo, QCAZ; 1, Pichincha, Puerto Quito, QCAZ; 1, Loja, Malacatos, QCAZ; 1, Los Ríos, San Carlos, 100 m, 79 26 6 W 1 6 42 S, QCAZ; 1, Orellana, Loreto Cotapino, 640 m, QCAZ; 1, Los Ríos, Palenque, QCAZ; 1, El Oro, Buenaventura, Piñas, 620 m, S. Padrón, QCAZ; 1, Napo, Avila Viejo, 750 m, QCAZ; 1, Napo, Archidona, 77 50 00 W 0 55 0 S, QCAZ; 26, Orellana, Yasuní, 250 m, QCAZ; 1, Pichincha, Santo Domingo, QCAZ; 1, Los Ríos, Quevedo, 4 de abril/1960, C. Dosdson, NMNH; 1, Esmeraldas, San Lorenzo, 24/03/1979, Jos. J. Anderson, NMNH; 1, Los Ríos, Río Palenque Biological Station 40 km Quevedo, 14/03/1979, J. Anderson, NMNH; Pastaza, Puyo (22 km W), 28373, D. R. Givens, NMNH; Pastaza, Santa Clara, 30/06/1976, P. M. Turner, NMNH; [19]; Orellana, Yasuní, [18, 59, 80]; 4, Loja, Malacatos - Ceibopamba, 1500 m, -4,167369, -79,300962, 02/III/2017, P. Picón, S. Padrón, MZUA; 1, Loja, Malacatos, -4,198158, -79,262703, 03/XI/2016, P. Picón, S. Padrón, MZUA.
Exaerete trochanterica (Friese, 1900)
Records: [19]; 1, Orellana, Yasuní, Julio 2017, D.W. Roubik, QCAZ.
4. Discussion
With 115 species reported for Ecuador, our results almost double the number known from previous work [19] (62) and [18] (63 species). In a medium elevation area of north central Peru [81] there may be similar species richness, but as we suggest below, the highest species diversity occurs, as in many groups, in NE Ecuador.
In Ecuador, the genus Euglossa is the most diverse, with 68 species, followed by Eufriesea 23, Eulaema 18, Exaerete 5, and Aglae 1 species. Fifty-two new species records are added for the country, compared to the most recent species list, in [18]. The new species records are distributed in four genera. Thirty-nine new species records are added for the genus Euglossa, 4 new species records for Eulaema, 9 new species records for Eufriesea, and 2 new species records for Exaerete (Table 1). There are at least 10 more undescribed species.
New species records added for Ecuador.
Genus
Species
Euglossa
amazonica Dressler, 1982
analis Westwood, 1840
avicula, Dressler, 1982
bidentata Dressler, 1982
championi Cheesman, 1929
charapensis Cockerell, 1917
chlorina Dressler, 1982
cybelia Moure, 1968
deceptrix Moure, 1968
despecta Moure, 1968
dissimula Dressler, 1978
dressleri Moure, 1968
gaianni Dressler, 1982
hansoni Moure, 1965
heterosticta Moure, 1968
iopyrrha Dressler, 1982
maculilabris Moure, 1968
magnipes Dressler, 1982
modestior Dressler, 1982
platymera Dressler, 1982
pleosticta Dressler, 1982
retroviridis Dressler, 1982
securigera Dressler, 1982
variabilis Friese, 1899
bigibba Dressler, 1982
granti Cheesman, 1929
perfulgens Moure, 1967
singularis Mocsáry, 1899
allosticta Moure, 1969
asarophora Moure & Sakagami, 1969
flammea Moure, 1969
imperialis Cockerell, 1922
augaspis Dressler, 1982
moratoi Nemésio & Engel, 2012
dodsoni Moure, 1965
gorgonensis Cheesman, 1929
laevicincta Dressler, 1982
turbinifex Dressler, 1978
intersecta Latreille, 1817
Eulaema
nigrita Lepeletier, 1841
chocoana Ospina-Torres & Sandino-Franco, 1997
marcii Nemésio (2009)
tenuifasciata (Friese, 1925)
Eufriesea
auripes (Gribodo, 1882)
combinata (Mocsáry, 1897)
concava (Friese, 1899)
duckei (Friese, 1923)
magrettii (Friese, 1905)
opulenta (Mocsáry, 1908)
pulchra (Smith, 1854)
purpurata (Mocsáry, 1896)
venezolana (Schrottky, 1913)
Exaerete
trochanterica (Friese, 1900)
lepeletieri Oliveira & Nemésio, 2003
Finally, for the monotypic genus Aglae, here we added new distribution records for its conspicuous species A. coerulea Lepeletier & Seville, 1825, in Napo, Pastaza, and Esmeraldas Provinces, extending its known distribution to the Eastern Andes in Ecuador. Previously it was reported for Ecuador by [50] from the Chocó Region in western Ecuador.
Our results show a clear richness pattern (Figure 1), with a concentration of species in the Amazon region. As noted by previous authors [13, 19, 40, 66] the northwest is similar to Mesoamerica and Panama. The Andean and inter-Andean region, between the Cordillera Occidental and Oriental, may hold endemic euglossines, but the overall similarity to Peru and Colombia does not permit such a view at present. The lower to medium elevation Andes region may, like the palms [41], hold the most species. For now, most Ecuadorian euglossines have been found east of the Andes, in the lowlands. This is perhaps the result of more intensive sampling there, especially at the Estación Biológica Yasuní. It is also clear that some areas are poorly sampled, especially the southern part of Ecuador. In those areas, such as the Cordillera del Condor, Cerro Plateado, Zapotillo, and Zumba Regions, more new species records for the country will likely be found, as is now the case in Yasuní forest, in the northeast.
Any single reported collection should, for the time being, be viewed with some skepticism and needing replication and confirmation. The single recorded specimens that differ from the gender of the designated holotype, for example, the male of Eufriesea combinata or Eufriesea theresiae [51], require further field work to validate. We see a need to emphasize that some species are quite difficult to identify and distinguish, likely due to mimicry, and phylogenetically close relatives that first separated by climate changes during glaciations, either on both sides of the Andes or in higher elevation areas, compared to lowlands. There are reasonably cryptic sibling or close relatives. For example, Euglossa chalybeata and E. piliventris, previously thought to be native to Ecuador, have yet to be found there, but in their place Euglossa orellana, E. occidentalis, and E. lugubris were described [65]. Similarly, Kimsey [13] pointed out the synonymy of Eufriesea andina with its senior synonym, Ef. magrettii, and the color polymorphism of the males in both this species and Ef. venezolana, which had confused previous taxonomic work. We find also very slight differentiation between two Eulaema and subgenus Apeulaema, although some of the reported males in Ecuador may in fact match those of French Guiana—the type area for Eulaema cingulata—and we have examined two from Eastern Ecuador that are different, one indistinguishable externally from Eulaema of Panama, which is now tentatively given the name Eulaema marcii, but which was described from Eastern Brazil. Further study of genitalic details would repay the effort.
Conflicts of Interest
The authors declare that there are no conflicts of interest regarding the publication of this paper.
Acknowledgments
The authors would like to thank the Universidad del Azuay and its Research Funding Program 2016-2017 for the support to the fieldwork and the museum visits of Pablo S. Padrón and Ruben P. Picón. Pablo S. Padrón wishes to thank Cristina Lalama for help databasing museum specimens and also José Falcón and Henry Garzón for help during fieldwork. The authors are also grateful to Albaro Barragán, Brian Harris, and Santiago Villamarín for allowing them to examine specimens from the collections under their care at the QCAZ, NMNH, and INABIO, respectively. Finally they want to thank Ministerio del Ambiente del Ecuador for the Collecting Permits.
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