Research Article |
Corresponding author: Nathalia Flórez-Gómez ( naf63@cornell.edu ) Academic editor: Jack Neff
© 2023 Nathalia Flórez-Gómez, Bryan Danforth.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Flórez-Gómez N, Danforth B (2023) The North American bees of the genus Ptilothrix Cresson, 1878 (Hymenoptera, Apidae, Emphorini), with the description of two new species. Journal of Hymenoptera Research 95: 275-293. https://doi.org/10.3897/jhr.95.96025
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Ptilothrix Cresson is a genus of New World bees with an amphitropical distribution. Like other genera in the tribe Emphorini, Ptilothrix have narrow pollen preferences. These solitary ground-nesting bees exhibit a remarkable nesting behavior in which females carry water from ponds to facilitate the excavation of the hard soil where they nest. With 16 described species, there are few taxonomic studies and, before this work, a lack of taxonomic treatments for the species in North America. Thus, in this study we revised and recognized four species for the region: Ptilothrix bombiformis Cresson, Ptilothrix sumichrasti Cresson, Ptilothrix chiricahua Florez-Gomez & Danforth, sp. nov. and Ptilothrix zacateca Florez-Gomez & Danforth, sp. nov. We describe and illustrate males and females of the two new species. We also present diagnoses for the four species, a key to identify them, and a map of their geographic distributions.
Anthophila, Apoidea, Chiricahua, Mexico, taxonomy
The genus Ptilothrix Cresson, 1878 is a group of bees restricted to the Western hemisphere with an amphitropical distribution. In North America the genus is found from Ontario, Canada to Oaxaca, Mexico and in South America it occurs from northern Brazil to northern Patagonia, Argentina (
Within Emphorini, Ptilothrix is phylogenetically related to Diadasina and Melitomella (
The genus currently includes 16 described species (
We examined specimens deposited in the following collections:
Cornell University Insect Collection (
The morphological terminology used in this study follows that proposed by
Finally, the known distribution obtained from the specimen labels of the four species were mapped using QGIS software and topographic layers downloaded from EarthEnv. For Ptilothrix bombiformis, we additionally included data from the Global Biodiversity Information Facility (GBIF).
Body size of females from 8.6–12 mm, males 9–10.8 mm. This species is distinguished by the polished and shiny ocellocular area, with few scattered punctures close to the eye margin in both sexes. Morphologically similar to P. sumichrasti, but males differ by the yellowish pubescence on T7, and the shape of S7, S8 and the genital capsule (Figs
Ptilothrix chiricahua sp. nov. A male holotype habitus, lateral view B male holotype habitus; frontal view C male holotype dorsal view D male holotype metasoma E female paratype habitus, lateral view F female paratype habitus, frontal view G female paratype dorsal view H female paratype metasoma.
A Ptilothrix chiricahua sp. nov. male S7 B Ptilothrix chiricahua sp. nov. male S8 C Ptilothrix zacateca sp. nov. male S7 D Ptilothrix zacateca sp. nov. male S8 E Ptilothrix sumichrasti male S7 F Ptilothrix sumichrasti male S8 G Ptilothrix bombiformis male S7 H Ptilothrix bombiformis male S8.
A, B Ptilothrix chiricahua sp. nov. male genital capsule A dorsal view B ventral view C, D Ptilothrix zacateca sp. nov. male genital capsule C dorsal view D ventral view E, F Ptilothrix sumichrasti male genital capsule E dorsal view F ventral view G, H Ptilothrix bombiformis male genital capsule G dorsa view F ventral view.
Male. Total length 9.16 mm (paratypes 9–10.8 mm, n=10). Head. Integument black, except mandible brown with middle area lighter brown; apex yellowish in some paratypes. Mandible with rounded apical margin. Pubescence whitish on labrum, clypeus, around antennal sockets, frons and gena, becoming pale yellowish on vertex, whitish in some paratypes. Labrum rectangular, disc densely punctate, margin raised and impunctate. Clypeus protuberant in lateral view, with punctures densely distributed, separated by a distance less than a puncture diameter. Punctation of lower paraocular area, supraclypeal area and frons punctation as in clypeus. Inner ocular margins subparallel. Ocellocular area polished, with few scattered punctures and short pilosity closer to the eye margin, distance 0.52 mm. Antennae dark brown, scape 0.68 mm long, first flagellomere longer than broad, 1.7 times longer than its apical width. Head length 2.6 mm. Head width 3.2 mm. Gena width 0.62 mm. Lower interocular distance 1.8 mm. Upper interocular distance 2 mm. Lower interocular distance 0.8 mm. Antennocular distance 0.28 mm. MOD 0.228 mm.
Mesosoma. Integument black. Pubescence fulvous on the pronotum, scutum, scutellum and metanotum; becoming whitish towards the propodeum, mesepisternum and metepisternum; some paratypes with pubescence entirely whitish. Propodeal triangle with short appressed, whitish hairs. Scutum with homogeneously distributed punctures separated by a distance up to a puncture diameter. Mesepisternal disc with punctures densely distributed, separated by a distance up to a puncture diameter, interspaces shiny. Hypoepimeral area with close punctures on the upper area, impunctate and polished towards the scrobal groove. Tegula translucent brownish. Tuft of yellow pubescence at base of wings, membrane of wings subhyaline, venation brown. Length of forewing 8 mm. Intertegular distance 2.6 mm. Scutum length 2.35 mm. Legs. Integument dark brown, except tarsi reddish brown, with whitish pubescence. Hind coxa and femur enlarged. Hind basitarsus slightly curved, 5 times the length of the second tarsomere. Posterior tibial spurs slightly hooked at the apex.
Metasoma. Tergal integument black, sterna brown. T1 and T2 covered with whitish erect pilosity and posterior margin with a band of yellowish appressed hairs. T3–T6 with appressed short, dark hairs, margin with bands of yellowish pilosity. T7 covered with yellowish pubescence, apical margin with two pointed projections forming a medial U–shaped notch. Sternal pubescence erect, yellowish. S6 with a median tuft of hairs. Metasomal pubescence whitish in some paratypes. S7 and S8 as in Fig.
Female. Total length 10.62 mm (8.6–12 mm n=10). Head. Integument color as in males. Maxillary palpus 6–segmented, segments 1–4 with short setae on the outer margin. Mandible with rounded apical margin. Pubescence color as in males, but hairs shorter. Labrum densely punctate, apex subtriangular with margin raised, impunctate. Inner ocular margins nearly parallel. Clypeus as in male, interspaces between punctures up to a puncture diameter. Punctation in paraocular area as in males. Ocellocular area polished and with few scattered punctures and short pilosity closer to the eye margin, distance 0.56 mm. Antennae dark brown, scape 0.71 mm long, first flagellomere 1.8 times longer than its apical width (Fig.
Holotype. United States of America • ♂; New Mexico; Hidalgo Co.1 mi. S Rodeo; 23 Aug. 2002; B.N. Danforth leg.; on Kallstroemia;
Paratypes. Mexico – Sonora • 1 ♀; Rancho San Bernardino. 28 km E Agua Prieta, Ciénaga; 27 Jul. 2000; K. Toal leg.; MCUR 1519 • 1 ♀; same data as for preceding; 22 Jun. 2000; MCUR 1874 • 1 ♂; 30 km E Agua Prieta; 31°19'08"N, 109°15'04"W; 12 Aug 2001; A. Romero leg., MCUR SBV043745.
United States of America – Arizona • 1 ♂; Cochise Co. 10 mi SW Apache; 31.571°N, 109.26°W; 27 Aug. 2014; B.N. Danforth leg.;
Mexico – Sonora • 1 ♀; 30 km E Agua Prieta; 31°30'00"N, 109°28'00"W; 1 Aug 2001; E. Serrano leg., MCUR SBV069744 • 1 ♀; same data as for preceding; MCUR SBV069979 • 1 ♀; same data as for preceding; MCUR SBV070114 • 1 ♀; same data as for preceding; MCUR SBV070141 • 1 ♀; same data as for preceding; 1–15 Aug. 2001; R.L. Minckley leg.; MCUR SBV079342 • 1 ♀; same data as for preceding; MCUR SBV079339 • 1 ♀; same data as for preceding; MCUR SBV079331 • 1 ♀; Rancho San Bernardino. 28 km E Agua Prieta, Ciénaga; 27 Jul. 2000; K Toal leg.; MCUR 1329 • 1 ♀; same data as for preceding; 10 Jul. 2000; MCUR 232 • 1 ♀; same data as for preceding; MCUR 2555 • 1 ♀; same data as for preceding; MCUR 226 • 1 ♀; same data as for preceding; MCUR 182.
United States of America – Arizona • 1 ♂; Portal. Cochise Co.; 7 May 1977; R Brooks leg.; on Kallstroemia; SEM 1304613 • 1 ♂; Pima, San Luis Wash; 31.63227°N, 111.429°W; alt. 1038 m; 19 Aug. 2016; T.L. Griswold leg.;
P. chiricahua is associated to mixed Chihuahuan desert grassland habitats of Southern Arizona, New Mexico, Western Texas and Northern Mexico (Fig.
This species is named in honor to the Chiricahua Apache people whose historical homeland encompasses the area where this bee now occurs.
Female body size from 11.3–12.7 mm, male 10.5–13 mm. This species can be recognized by the overall whitish to pale yellowish pubescence, except for that on the female basitarsi, which is dark yellowish to pale brown. Differing from P. sumichrasti and P. chiricahua by the longer first flagellomere (length at least twice its apical width), presence of erect, overall longer pubescence, especially on the anterior areas of T2 and T3, more robust body and slightly larger size. Both sexes with tergal hair bands on T1–T4 in females, T1–T6 in males.
Male. Total length 12.34 mm (paratypes 10.5–13 mm, n=7). Head. Integument black, except brown middle area of the mandible, mandibular apex yellowish in some paratypes. Uniformly whitish pubescence covering all of the head except ocellocular area where hairs are sparser. Mandible with rounded apical margin. Maxillary palpus 6–segmented. Labrum rectangular, disc densely punctate, margin raised and impunctate. Clypeus slightly protuberant in lateral view, with coarse punctation. Inner ocular margins subparallel. Frons, paraocular area and vertex with close punctures, separated by a distance up to a puncture diameter. Ocellocular integument micro–sculptured and with punctures towards the eye margin, small area around the lateral ocellus polished and shiny, distance 0.63 mm. Antennae black, scape 0.8 mm, first flagellomere 2.5 times longer than its apical width. Head length 3.25 mm. Head width 3.85 mm. Gena width 0.8 mm. Lower interocular distance 2.45 mm. Upper interocular distance 2.5 mm. Interocular distance 1.03 mm. Antennocular distance 0.41 mm. MOD 0.25 mm
Mesosoma. Integument black. Overall pubescence whitish on pronotum, scutum, scutellum, mesepisternum, metepisternum and propodeum; pale yellowish on the scutum of some specimens. Scutum with close punctures regularly distributed, separated by less than a puncture diameter with shiny interspaces. Scutellar punctures even more closely spaced than those of the scutum. Disc of mesepisternum punctured as in scutum, deep punctures and very close one to each other, imbricate interspaces between punctures. Hypoepimeral area deeply and densely punctate, micro–sculptured towards the scrobal groove. Propodeal triangle with whitish short pubescence, middle line glabrous. Tegulae translucent brown. White tuft of hairs at the base of wings, membrane of wings slightly infuscate, venation brownish. Length of forewing 10.83 mm. Intertegular distance 3.25 mm. Scutum length 3.0 mm. Legs. Integument black to dark brown, except tarsi reddish brown. Pubescence whitish overall. Hind coxa and femur enlarged. Hind basitarsus slightly curved, 6 times the length of the second tarsomere. Hind tibial spurs dark brown, slightly hooked at the apex.
Metasoma. Overall integument black, margin of sterna subhyaline. T1–T4 with dense, whitish erect pubescence, posterior margin with band of white hairs. T5–T6 with dense and erect black pubescence on the anterior area, margin with a band of whitish dense pubescence. T7 covered with black pubescence, margin with two pointed projections forming a middle notch. Sterna with dense, whitish, erect pubescence. S6 with a median tuft of hairs. S7 and S8 as in Fig.
Female. Total length 12.67 mm (11.3–12.7 mm n=8). Head. Integument color as in male, pubescence as in male, but less abundant. Mandible color and shape as in males. Labrum densely punctate, with subtriangular, raised, impunctate apex. Clypeus with coarse punctures separated by a distance up to twice a puncture diameter. Punctation of paraocular area and frons as in males, puncture size smaller than those on clypeus. Inner ocular margins nearly parallel. Integument of ocellocular area as in males, distance 0.76 mm. Antennae black, scape 0.93 mm long, first flagellomere 2.3 times longer than its apical width (Fig.
Holotype. Mexico • ♂; Zacatecas; Guadalupe; 28 Jun. 1953; C. & P. Vaurie leg.; D. Rockefeller Mex exped.; >
Paratypes. Mexico – Durango • 2 ♂; 14 mi N.E of Durango; alt. 1889 m; 17 Jun. 1956; H.A. Scullen leg.;
This species is endemic to Mexico. Records are from the mountain regions in the Sierra Madre Occidental from 1800–2000 m (Fig.
This species is named in honor to the Zacatecos, the indigenous group that inhabited the state of Zacatecas, the area where the holotype was collected.
Ptilothrix sumichrasti Cresson, E. T. (1878). (Lectotype: ANSP #f: Mexico).
Body size of females from 10–10.6 mm, males 8.2–10 mm. This species is morphologically similar to P. chiricahua but is recognizable by the punctation pattern in the ocellocular area, which is micro–sculptured, with punctures towards the eye margin and a small area around the lateral ocellus polished and shiny in both sexes. Males are clearly distinguishable by the brown pubescence covering T7, in addition to the shape of S7, S8 and genital capsule (Figs
Mexico – Guerrero • 1 ♂; Iguala;
Guatemala • 3 ♀; C.A; alt. 300 m; 10 Jan.1923; >
Records of this species are from lowlands in Guatemala and Mexico in the Transmexican Volcanic Belt and the Balsas River Basin (Fig.
Emphor bombiformis Cresson, 1878.
Emphor fuscojubatus Cockerell, 1913.
Ptilothrix bombiformis Cresson, (1878). (Lectotype: ANSP #f: Kansas, Snow).
Large bees, female body size from 13.5–19 mm, males from 12.5–18.6 mm. This species is clearly recognizable by the white to pale yellow pubescence on the head and mesosoma, and entirely black metasoma, except T1 which has whitish pubescence on the lateral sides in some specimens. Differs from P. sumichrasti, P. chiricahua and P. zacateca by the absence of yellowish metasomal bands, darkened wings, entirely black scopa, and overall larger size. First flagellomere length twice its apical width, as in other Ptilothrix species except P. sumichrasti and P. chiricahua. Shape of male S7, S8 and genital capsule as shown in Figs
Ptilothrix zacateca sp. nov. A male holotype habitus, lateral view B male holotype habitus; frontal view C male holotype dorsal view D male holotype metasoma E female paratype habitus, lateral view F female paratype habitus, frontal view G female paratype dorsal view H female paratype metasoma.
United States of America – Alabama • 1 ♂; Morgan; 34.6059°N, 86.9833°W; alt 178 m; Aug.1944; GE Bohart leg.;
This species is the most widely distributed in North America, its geographical range from Ontario, Canada (
1 | Metasoma with bands of yellowish or whitish pubescence from T1–T4; wings infuscate or subhyaline; female hind tibia with white or yellow scopa, yellow or brownish on basitarsus | 2 |
– | Metasoma pubescence entirely black, without bands, T1 sometimes with yellow pubescence on lateral areas; wings darkened, female hind tibia and basitarsus with entirely black scopa | Ptilothrix bombiformis |
2 | First flagellomere up to two times longer than its apical width (Fig. |
3 |
– | First flagellomere more than two times longer than its apical width (Fig. |
Ptilothrix zacateca sp. nov. |
3 | Ocellocular area predominantly impunctate, polished and shiny (Fig. |
Ptilothrix chiricahua sp. nov. |
– | Ocellocular area micro–sculptured, with scattered punctures and a small polished area around the lateral ocellus (Fig. |
Ptilothrix sumichrasti |
The genus Ptilothrix is a species–rich group, especially in South America (
Ptilothrix bombiformis is the most widely distributed species in North America and one of the most commonly collected, it is associated to the genus Hibiscus (Malvaceae), although occasionally can visit flowers of Ipomoea (Convolvulaceae) (
We thank the Cornell University Insect Collection (