Anthophora (Micranthophora) columbariae Timberlake and Cockerell, 1937

Orr, Michael C., Pitts, James P. & Griswold, Terry, 2018, Revision of the bee group Anthophora (Micranthophora) (Hymenoptera: Apidae), with notes on potential conservation concerns and a molecular phylogeny of the genus, Zootaxa 4511 (1), pp. 1-193 : 38-40

Anthophora (Micranthophora) columbariae Timberlake and Cockerell, 1937

( Figs. 39F , 44F , 49F , 54F , 59F , 64F , 69F , 74F )

Anthophora (Micranthophora) columbariae Timberlake and Cockerell in Cockerell, 1937: 9 (holotype: ♂; Morongo, CA, USA; coll. T.D.A. and W.P. Cockerell, 22 Apr. 1937, at Salvia columbariae ; AMNH).

Diagnosis. Males may be distinguished from all species but A. salazariae by the following character combination: galea never reaching past rear of midtrochanter in repose and weakly tessellate at most, shiny, without striations; labral rim rounded; mandible with inferior tooth small, running roughly parallel to main blade ( Fig. 49F ); scape immaculate; and T7 with strong medial longitudinal carina and a pair of vaguely triangular projections medially ( Fig. 59F ). Females may be distinguished from all species but A. salazariae by the following character combination: galea smooth to weakly tessellate, without striations or long setae; mandible with inferior tooth a thin ridge running near-parallel to main blade ( Fig. 39F ); clypeus divided by a very slight, often impunctate, vertical line along median ( Fig. 39F ); scutum shiny, with impunctate areas; tergal setae relatively sparse, giving a darker appearance without a microscope; T1–T4 with appressed setae throughout, but sometimes with very thin, easily overlooked black setal patches basally. From A. salazariae , this species may be separated with care and reference material according to couplet 11 (females) or couplet 16 (males).

Male description. Head: Facial maculations creamy to pale yellow-green. Galea reaching past rear of forecoxa in repose, usually obviously; light brown to amber; smooth or at most weakly tessellate, with strong reflections. Mandible with inferior tooth a slight ridge, sitting parallel to main blade ( Fig. 49F ); maculated from base to start or middle of inferior tooth. Labral rim rounded; fully maculated. Clypeal maculation a weak band along rim that is frequently absent, clearly not reaching lateral sides, almost never filling third of clypeal height medially ( Fig. 49F ); punctation significantly sparser within maculation than without, sometimes impunctate. Paraocular area immaculate. Supraclypeal area immaculate. Scape immaculate. Mesosoma: Scutum shiny; with distinct impunctate areas. Tegula transparent, tan to light brown. Basitibial plate almost always present; usually weak, somewhat obscure. Metasoma: T1 roughly covered in appressed setae; T2–T4 with appressed setal bands of variable size, typically filling half or less of terga; T5–T6 without appressed setae. Terga smooth to weakly tessellate between punctures, usually with strong reflections; apparent through appressed setae. Tergal rims somewhat translucent, medium brown to tan. Male T7 ( Fig. 59F ) with strong medial longitudinal carina; lateral projections absent; medial projections moderately long, triangular. Male S5 ( Figs. 38A, 38C ) setal arrangement unremarkable, without large, distinctive patches, at most longer along rim. Male S6 ( Fig. 38A ) clearly medially emarginate. Male S7 ( Fig. 64F ) with setae fully banding S7 around midpoint of length; widening from midpoint to tip, apical half widest at pointed lateral tips; apical half resembling cat head, rim broadly emarginate between lateral tips. Male S8 ( Fig. 69F ) lateral projections distinct; medial projection narrowing to tip, apically emarginate. Male genital capsule ( Fig. 74F ) with outer corners, where gonocoxite tips curve inward, marked by distinct and sharp ventral point, best seen in profile; from corner to apex, in profile, gonocoxite with low secondary bump but relatively unmodified overall; tip, in profile, rounded and weakly curved ventrally; gonostylus tip position slightly exceeding that of gonocoxite, measured from above along primary axis of latter. Pubescence: See Fig. 54F . Appears medium to dark gray overall. All white, except as follows: leg inner faces orangish; tergal basal zones dark; T5–T6 with simple setae predominantly dark, except sometimes light in apical third.

Female similar to male, except: Head: Galea often darker brown. Mandible with inferior tooth more produced, sometimes slightly angled off main blade ( Fig. 39F ). Clypeal maculation less often absent. Mesosoma: Basitibial plate circular distally, teardropped overall. Metasoma: T1–T4 usually covered in appressed setae, sometimes absent near base; T5 without appressed setae. T4 impunctate rim of short length medially, often barely visible. Anal fimbria strongly triangular, filling third to almost half of T5 length medially. Pubescence: See Fig. 44F . Vertex and scutum often with dark intermix; hindbasitarsal brush dark brown; tergal basal zones dark, if present; T5 simple setae predominantly dark; anal fimbria dark brown.

Distribution. See Fig. 11 . Present in USA: California; Mexico: Baja California Norte. Found most commonly in mid-high elevation xeric and mesic areas. Ecoregional habitance is as follows, based on 211 specimen records: 6 ecoregions total: Mojave Desert (115=55%), California coastal sage and chaparral (56=26.8%), California montane chaparral and woodlands (18=8.1%), California interior chaparral and woodlands (10=4.3%), Sonoran Desert (8=3.8%), Baja California desert (3=1.4%), NULL (1≤1%).

Phenology. Recorded from March–June, September, almost exclusively spring, most common in April–May. Univoltine, single September record from San Luis Obispo (CA) possibly mislabeled, opportunistic emergence, or indicative of altered phenology in less-known area of range.

Nesting biology. Unknown.

Floral specialization. Reported as a potential specialist on Fabaceae and Salvia (Lamiaceae) ( Moldenke & Neff 1974). Appears possibly true; known associations generally support this premise but suggest use of Lamiaceae more broadly; almost all floral associations for females are with these two plant families. However, female visits to Fabaceae are only known from three independent collection events where series were taken, while females have been collected off Lamiaceae at 15 collection events; it seems likely that Lamiaceae is the true host. Primary recorded visits by 66 females include: Family Lamiaceae (59%), Fabaceae (29%); Genus Salvia (49%), Lotus (26%), Salazaria (11%); Species Salvia columbariae (36%), Salazaria mexicana (11%). 117 total floral records from eight families include: Asteraceae (7 ♂ 4 ♀): Chaenactis fremontii 1 ♂ 2 ♀, Chaenactis sp. 2 ♂, Chaenactis stevioides 1 ♀, Ericameria sp. 4 ♂ 1 ♀, Boraginaceae (4 ♂ 3 ♀): Cryptantha intermedia 2 ♂, Cryptantha muricata 1 ♂, Eriodictyon trichocalyx 1 ♂, Phacelia minor 3 ♀, Fabaceae (5 ♂ 19 ♀): Lotus scoparius 2 ♂, Lotus sp. 3 ♂ 17 ♀, Lupinus arizonicus 1 ♀, Lupinus sp. 1 ♀, Krameriaceae (1 ♂): Krameria sp. 1 ♂, Lamiaceae (31 ♂ 39 ♀): Salazaria mexicana 8 ♂ 7 ♀, Salvia columbariae 15 ♂ 24 ♀, Salvia mellifera 1 ♂ 3 ♀, Salvia sp. 7 ♂ 5 ♀, Malvaceae (1 ♂): Sphaeralcea ambigua 1 ♂, Rosaceae (1 ♀): Adenostoma sp. 1 ♀, Zygophyllaceae (2 ♂): Larrea sp. 1 ♂, Larrea tridentata 1 ♂.

Comments. Anthophora columbariae is exceptionally difficult to separate from A. salazariae , as many characters have rare exceptions. Reference material should be used wherever possible.

Specimens examined: MEXICO: Baja California Norte: El Rosario, 10-12 mi ESE: 2 ♂ 1 ♀, 17 Mar 1984, W.J. Pulawski. USA: CALIFORNIA: Los Angeles County: 1 mi S of Littlerock [= Little Rock] - Palmdale Dam, San Gabriel Mountains: 1 ♂, 24 Apr 1965, C.L. Hogue; 2 mi S of Pearblossom: 1 ♂, 25 Apr 1977, R.R. Snelling; 3 ♂ 7 ♀, 1–2 May 1977, R.R. Snelling; 1 ♀, 15 May 1983, collector unknown; Lake Caldwell, 1.2 mi W: 1 ♀, 16 May 1986, K.W. Cooper; Littlerock: 2 ♂, 11 Apr 1936, collector unknown; 1 ♂, 29 Apr 1949, C.D. MacNeil; Littlerock [= Little Rock] - Palmdale Dam: 6 ♂ 5 ♂, 2 May 1970, R.R. Snelling; Pearblossom, 2 mi S: 1 ♂, 1–2 May 1977, R.R. Snelling; Pearblossom, 2 mi W: 1 ♀, 12 May 1956, collector unknown; Pearblossom, 4 mi S: 1 ♂, 14 Apr 1960, R.R. Snelling; Valyermo, 4 mi SE: 1 ♂, 13 Apr 1960, R.R. Not not Use Snelling; 1 ♂, 13 Apr 1960, R.R. Snelling; Valyermo, San Gabriel Mountains: 2 ♀, 21 Apr 1940, R.M. Bohart; 1 ♂, 13 Apr 1960, R.M. Bohart; Westwood Hills: 1 ♂, 3 Jun 1935, E.G. Linsley; Monterey County: Arroyo Seco, The Lakes: 1 ♀, 26 Apr 1996, T.L. Griswold; King City, 9 mi E: 1 ♂, 1 Apr 1959, D. Burdick; 2 ♂, 1 Apr 1959, D.J. Burdick; 1 ♀, 1 Apr 1959, collector unknown; Riverside County: 9 mi NE of Palm Springs: 1 ♀, 19 Apr 1957, collector unknown; Banning, 4 mi S: 1 ♂ 1 ♀, 23 May 1976, T.L. Griswold; Banning, Rt. 243: 1 ♀, Mar 1982, K.W. Cooper; Cottonwood Spring, 6 mi S: 1 ♂, 11 Apr 1963, R.L. Langston; Gavilan Hills, southeast of Lake Mathews, near Gavilan Peak: 1 ♂, 22 Apr 1932, P.H. Timberlake; 1 ♂, 27 Mar 1933, C.M. Dammers; 1 ♀, 3 May 1937, collector unknown; 1 ♂, 27 Apr 1938, P.H. Timberlake; 1 ♀, 2 Jun 1938, P.H. Timberlake; 1 ♂ 1 ♂, 20 Apr 1939, P.H. Timberlake; 1 ♂, 14 Apr 1940, P.H. Timberlake; 1 ♀, 18 Apr 1940, P.H. Timberlake; 2 ♂ 1 ♀, 8 May 1950, P.H. Timberlake; 1 ♂, 17 May 1951, E.I. Schlinger; Gavilan, The: 1 ♂, 18 May 1951, E.I. Schlinger; 1 ♀, 7 May 1957, P.H. Timberlake; Hemet: 1 ♀, 10 May 1936, E.G. Linsley; Hemet, 3 mi S, Diamond Valley: 2 ♂, 25 Apr 1980, K.V. Krombein; Joshua Tree National Monument: 1 ♂, 14 Apr 1969, P.F. Torchio; 1 ♂, 14 Jun 1969, P.F. Torchio; Joshua Tree National Park; Arch Rock, 2.5 air km SE: 3 ♂, 25 Mar 2016, M.C. Orr; Joshua Tree National Park; Barker Dam, 1.7 air km SE; Queen Valley: 1 ♂, 20 Apr 2016, M.C. Orr; Joshua Tree National Park; Belle Campground: 2 ♂ 4 ♂, 21 Apr 2016, M.C. Orr; Joshua Tree National Park; Cottonwood Visitor Center: 1 ♀, 19 Apr 2016, M.C. Orr; Joshua Tree National Park; Hall of Horrors: 1 ♂ 3 ♀, 21 Apr 2016, M.C. Orr; Joshua Tree National Park; Key's View, 1.5 air km NE: 1 ♂, 21 Apr 2016, M.C. Orr; Joshua Tree National Park; Skull Rock: 4 ♀, 21 Apr 2016, M.C. Orr; Millard Canyon: 1 ♀, date unknown, collector unknown; Mount San Jacinto: 1 ♀, 26 Apr 1990, K.W. Cooper; Mountain Center, 2.8 mi W: 1 ♀, 29 Apr 2013, Z.M. Portman; Palm Springs: 1 ♀, 1–10 Apr 2013, F.D. Parker, T.D. McIntyre; Palm Springs, 9 mi NE: 1 ♀, 19 Apr 1957, M.D. Stage; Palm Springs, Araby Wash: 2 ♀, 14–21 Apr 2015, F.D. Parker, T.D. McIntyre; Pinon Flat: 1 ♂, 18 Apr 1962, C.D. MacNeill, Rentz, Brown, J.C. Lundgren; Radic, 5 mi N: 1 ♂, 7 May 1983, K.W. Cooper; Santa Rosa Mine Road, 1.5 mi W Perris: 3 ♂ 1 ♀, 17 Apr 1938, P.H. Timberlake; 1 ♂ 1 ♀, 27 Apr 1938, P.H. Timberlake; 2 ♂ 5 ♀, 4 May 1938, P.H. Timberlake; Split Rock, near 29 Palms: 1 ♀, 7 May 1948, P.H. Timberlake; Temecula: 1 ♂, 24 Apr 1951, E.I. Schlinger; The Gavilan, S entrance: 1 ♂, 16 Apr 2016, M.C. Orr; San Benito County: Pinnacles Natl. Mon., Chalone Creek Picnic Area: 2 ♂, 1 Jun 1998, F.D. Parker; San Bernardino County: 5 mi N of Adelanto, vic Adelanto Garbage Dump: 1 ♀, 4 May 1966, S.I. Frommer; 7.5 mi S Twentynine Palms (= 29 Palms): 1 ♂ 2 ♀, 7 May 1948, P.H. Timberlake; Big Morongo Valley: 1 ♀, 28 Mar 1984, K.W. Cooper; Deep Creek, near Mojave River Forks, Lower Mojave River: 1 ♀, 10 May 1937, P.H. Timberlake; Hesperia, 5 mi SE: 2 ♀, 1 May 1961, J.G. Rozen, R. Schrammel; Hesperia, 9.5 mi SE: 1 ♂ 2 ♀, 29 Apr 1986, K.W. Cooper; Joshua Tree National Park; Quail Spring Valley, sandy burm: 1 ♂ 1 ♀, 25 Mar 2016, M.C. Orr; 1 ♂ 3 ♂, 20 Apr 2016, M.C. Orr; Morongo: 1 ♂, 22 Apr year unknown, collector unknown; Morongo Valley: 2 ♂, 21 Apr 1957, R.R. Snelling, M.D. Stage; 1 ♀, 25 Apr 1957, R.R. Snelling, M.D. Stage; 1 ♀, 12 Apr 1960, R.R. Not not Use Snelling; 1 ♀ 3 ♀, 12 Apr 1960, R.R. Snelling; 1 ♀, 12 Apr 1960, collector unknown; Morongo Valley, San Bernardino Co: 1 ♂, 18 Apr 1957, R.R. Snelling, M.D. Stage; 3 ♀, 12 Apr 1960, R.R. Snelling; Morongo [= Morongo Valley], San Bernardino Mountains: 1 ♀, 19 Apr 1937, P.H. Timberlake; 1 ♂ 1 ♀, 22 Apr 1937, P.H. Timberlake; 2 ♀, 22 Apr 1937, T.D.A. Cockerell; 4 ♂, 22 Apr 1937, W.P. Cockerell; 1 ♂, 21 Apr 1957, collector unknown; 3 ♀, 25 Apr 1957, collector unknown; 1 ♀, 26 Apr 1957, collector unknown; 3 ♀, 12 Apr 1960, R.R. Snelling; Phelan: 1 ♀, date unknown, collector unknown; Pioneertown: 1 ♂, 5–13 May 1986, K.W. Cooper; Sheep Creek, 6.5 km NNE Wrightwood: 1 ♂, 25–26 May 2005, M.E. Irwin; mouth of Deep Creek, near Mojave River Forks: 1 ♂, 5 May 1936, P.H. Timberlake; San Diego County: 7 mi SW of Boulevard: 9 ♂, 10 Jun 1956, R.R. Snelling; Boulevard, 7 mi SW: 2 ♂, 10 Jun 1956, R.R. Snelling; Culp Canyon: 1 ♂, 2 Apr 1962, E.I. Schlinger; Jacumba: 1 ♂, 17 May 1956, R.R. Snelling; San Felipe Valley: 1 ♀, 27 Apr 1946, J.L. Sperry; San Luis Obispo County: Baywood Park: 1 ♀, 15 Sep 1956, collector unknown; Black Mountain, 6 air mi NE Pozo: 1 ♀, 1 May 1962, R.L. Langston; La Panza Camp, 12 mi NE Pozo: 1 ♂, 29 Apr 1962, P.D. Hurd; McKittrick, 35 mi W: 1 ♀, 23 May 1970, P.F. Torchio, D.M. Torchio; Oso Flaco Lake: 1 ♀, date unknown, collector unknown; Santa Barbara County: Goleta: 1 ♀, date unknown, collector unknown; Ventura County: Hungry Valley, 5 mi S Gorman: 1 ♀, 6 May 1959, collector unknown.

Figures

FIGURE 11. Distribution of Anthophora columbariae. Generated with 211 specimen records from 74 locations.

FIGURE 38. Male sterna, S5 setae density and S6 rim form. Scale=0.5 mm. A) S5 with sparse setae; S6 medially emarginate, at least vaguely triangularly, evenly so (A. estebana, 72476). B) S5 setae density high, appearing tufted; S6 medially emarginate, at least vaguely triangularly, but at least slightly filled medially, sometimes tufted (A. maculifrons, 82068). C) S5 with sparse setae; S6 entire or nearly so, with transparent square of integument (A. petrophila, BBSL655748). D) S5 setae density high; S6 entire, medially knobbed (A. flavocincta, BBSL504065).

FIGURE 39. Female face of A) Anthophora abroniae (BBSL920115), B) A. albata (10234), C) A. caudata (BBSLID99019), D) A. chihuahua (BBSL502616), E) A. cockerelli (BBSL502283), F) A. columbariae (BBSL853114). Scale=0.5 mm.

FIGURE 44. Female habitus of A) Anthophora abroniae (BBSL920116), B) A. albata (BBSL513478), C) A. caudata (BBSL506712), D) A. chihuahua (BBSL502616), E) A. cockerelli (BBSL502289), F) A. columbariae (KWC0014759). Scale=1 mm.

FIGURE 49. Male face of A) Anthophora abroniae (BBSL502212), B) A. albata (BBSL504186), C) A. caudata (BBSLID98988), D) A. chihuahua (BBSL502617), E) A. cockerelli (BBSL502260), F) A. columbariae (BBSL853110). Scale=0.5 mm.

FIGURE 54. Male habitus of A) Anthophora abroniae (BBSL502223), B) A. albata (10046), C) A. caudata (BBSLID92552), D) A. chihuahua (BBSL502617), E) A. cockerelli (72472), F) A. columbariae (BBSL853157). Scale=1 mm.

FIGURE 59. Male T7 of A) Anthophora abroniae (BBSL348460), B) A. albata (9988), C) A. caudata (BBSL506708), D) A. chihuahua (BBSL502618), E) A. cockerelli (BBSL502252), F) A. columbariae (FDP80368). Scale=0.25 mm.

FIGURE 64. Male S7 of A) Anthophora abroniae (BBSL502216), B) A. albata (BBSL843758), C) A. caudata (BBSL506707), D) A. chihuahua (BBSL502617), E) A. cockerelli (BBSL502279), F) A. columbariae (BBSL502330). Scale=0.25 mm.

FIGURE 69. Male S8 of A) Anthophora abroniae (BBSL502211), B) A. albata (BBSL843758), C) A. caudata (BBSL506707), D) A. chihuahua (BBSL502617), E) A. cockerelli (BBSL502279), F) A. columbariae (BBSL502330). Scale=0.25 mm.

FIGURE 74. Male genital capsule of A) Anthophora abroniae (BBSL502216), B) A. albata (BBSL843758), C) A. caudata (BBSL506707), D) A. chihuahua (BBSL502617), E) A. cockerelli (BBSL502263), F) A. columbariae (BBSL502332). Scale=0.25 mm.