Laemosaccus bimaculatus Hespenheide, 2019

Hespenheide, Henry A., 2019, A Review of the Genus Laemosaccus Schönherr, 1826 (Coleoptera: Curculionidae: Mesoptiliinae) from Baja California and America North of Mexico: Diversity and Mimicry, The Coleopterists Bulletin (MIMICRY AND LAEMOSACCUS In an earlier paper (Hespenheide 1996), I presented the hypothesis that species of Laemosaccus of the L. nephele group with red humeral spots on the elytra were Batesian mimics of members of the Chrysomelidae in the subfamily Clytrinae. There is no evidence that Laemosaccus species are distasteful, and what is either L. nephele and / or L. obrieni have been reported as prey items of birds (Beal 1912). In Cave Creek Canyon, Cochise County, Arizona, 21 forms (species and “ subspecies ”) of Clytrinae were hypothesized to be the primary models of 22 species of mimics in the families Anthribidae (one species), Bruchidae (two species), Buprestidae (four species), Chrysomelidae, subfamily Cryptocephalinae (three species), Coccinellidae (six species), Curculionidae, subfamily Baridinae (one species), and Laemosaccus (five species). Of these, the coccinellids and the cryptocephaline chrysomelids are probably distasteful Mullerian co-mimics. Ecologically, the species of Laemosaccus co-occurred with their clytrine models on both desert legumes and canyon oaks, although more clytrine species occurred in the desert and more Laemosaccus species occurred in the canyons. Species of clytrines showing the mimetic pattern are common throughout Mexico (Bellamy 2003, who renamed the Mexican buprestid genus Acherusia Laporte and Gory, 1837 as Mimicoclytrina Bellamy to reflect their resemblance to clytrines), but decline in numbers of species and in the proportion of the clytrine fauna through Central America to Panama (Hespenheide 1996, fig. 2). Laemosaccus seems to follow a similar pattern. Mimicry is more common in large faunas, especially in wet tropical areas (Hespenheide 1986, 1995); because the largest clytrine fauna is in Mexico, the clytrine mimicry complex is also larger there (Hespenheide 1996). This complex has more members than I first enumerated and deserves further study. The evolution of mimicry produces resemblances between unrelated species (Laemosaccus and other putative mimics, with clytrines and perhaps other Chrysomelidae and Coccinellidae as models; see Hespenheide 1976, 1996) and selects against the divergence of related species. In Batesian mimicry - hypothesized to be the form of relationship between Laemosaccus and clytrines - the selection for precision of mimicry is stronger on the mimic (Laemosaccus), so that resemblances among them should be closer, regardless of ancestry. Close morphological resemblances based on ecology rather than ancestry may be termed mimetic homoplasy (Hespenheide 2005) and can make recognition of species difficult (as in Laemosaccus) or complicate phylogenetic analyses. I have speculated (Hespenheide 1996) that the sympatric “ subspecies ” of the clytrine models (Moldenke 1970) may in fact be reproductively isolated sibling species. It will be interesting to see whether or not genomic studies show the closeness of relationships among Laemosaccus species that the morphology suggests) 73 (4), pp. 905-939 : 918-920

Laemosaccus bimaculatus Hespenheide , new species

Zoobank.org/ urn:lsid:zoobank.org:act:9D7CF6AF-B97C-4EB9-AF4B-B938661A91E2 ( Figs. 10 , 21 )

Description. Holotype Male. Length 3.3 mm, width 1.5 mm ( Fig. 10 ). Robust, subcylindrical in cross section, slightly obovate, broadly rounded behind, more narrowly so in front, black except each elytron with large red-orange spot on anterior 3/4 from lateral margins nearly to suture and broadly rounded behind; pronotum and elytra glabrous, thorax and abdomen ventrally with punctures each with a silvery seta, head with inconspicuous setae on rostrum, denser and conspicuous behind lower 3/4 of eyes, setae more slender and semi-erect on basal half of femora, hair-like and erect on tergite 8; tergite 7 with small, scale-like setae on basal margin, otherwise glabrous. Head hemispherical, 0.55 mm wide, rostrum rounded-terete, coarsely punctate, 0.5 mm long, antennae inserted at middle. Pronotum gibbous, convex at base, constricted before anterior margin, 1.1 mm long, 1.25 mm wide, broadest at basal third, with lateral margins very shallowly arcuate, slightly convex in lateral view, coarsely, evenly punctate, punctures rounded and separate, with distinct medial carina on basal 3/4. Elytra distinctly wider than pronotum at base, 2.0 mm long, 1.5 mm maximum width, elytral striae subequal to intervals, striae coarsely punctate, intervals carinate, interval 3 weakly toothed on middle third, interval 5 weakly toothed on apical half. First abdominal ventrite very slightly concave along midline with setae less conspicuous. Profemora with broad, abruptly acute ventral tooth beyond middle. Genitalia as in Fig. 21 ; aedeagus 1.00 mm long.

Allotype Female. As male but rostrum subcylindrical, polished, very finely, inconspicuously punctate; tergite 7 weakly convex, coarsely punctate, glabrous except at basal margin.

Specimens Examined. Holotype: Arizona: Cochise Co., Pinery Cyn., Chiricahua Mts., below Onion Saddle , 7200’, 31°56’N 109°16’W, 25.06.1999, H. A. Hespenheide, on leaves of Quercus gambelli Nutt. (USNM) . Allotype: Arizona: Cochise Co., Chiricahua Mts. , E. Turkey Creek, 6.5 mi. W Portal, 6400’, 31°54-55’N 109°15’W, 14.07.1981, W. P. Weaver, Jr., oak ( USNM) . Paratypes: USA: Arizona: Box Cyn., Santa Rita Mts. , 2.06.1968 (1, ASUHIC) ; Sta Rita Mts , 17.07.1932, R. H. Beamer (1, SEMC) ; Cochise Co., Canelo Hills, Parker Cyn. , 30.07.1979, R. L. Westcott (1, WFBM) ; Chiricahua , 20.06.?, H. S. Barber (1, USNM) ; Chiricahua Mts. , 17.06.1908, V. Owen (2, LACM) , 01.07.1908, V. W. Owen (1, CASC) ; 1908, [E. S. Ross] (1, CASC) ; Chiricahua M., 24.07.1955, D. J. & J. N. Knull (1, OSU) ; Chiric. Mts. , “24.6,” “1.7” (2, USNM) ; Cave Ck. Cyn., Chiricahua Mts., 6 mi. WSW Portal, Greenhouse Tr , 6000’, 31°53’N 109°14’W, 26.06.1984, H. A. Hespenheide (1, CHAH) ; Cave Ck. Cyn., Chiricahua Mts., Herb Martyr Dam , 5800’, 31°52’N 109°14’W, 23.05.1997, H.A. Hespenheide, Quercus emoryi (1, CHAH) ; Chiricahua Mts., Herb Martyr Dam , 03.06.1989, W. F. Barr, on oak (1, WFBM) ; Cave Ck. Cyn., Chiricahua Mts., Sunny Flat , 5100’, 31°53’N 109°10’W, 22.05.1981, 28.05, 01.06.1982, 22, 23, 26.05.1985, H.A. Hespenheide, on Quercus (7, BMNH, CHAH) , 27.05.1997, H. A. Hespenheide, Quercus emoryi (1, CHAH) , 22.05.1995, L. M. LaPierre (1, CHAH) , 27.05.1997, H. A. Hespenheide, Quercus emoryi (1, CHAH) ; Cave Ck. Cyn., Chiricahua Mts. , 5 mi. WSW Portal, 5900’, 31°54’N 109°13- 14’W, 24.06.1999, H. A. Hespenheide (1, CHAH) ; Cave Ck. Cyn., Chiricahua Mts. , 6 mi. WSW Portal, 6500’, 31°54’N 109°14-15’W, 25.05.1997, H. A. Hespenheide, Quercus arizonicus (1, CHAH) ; Cave Ck. Cyn., Chiricahua Mts. , 6 mi. W Portal, 6700’, 31°55’N 109°15’W, 28.06.2001, H. A. Hespenheide, on Quercus hypoleucoides (1, CHAH) ; same data as Holotype (6, BMNH < CHAH), same data as Holotype but 26.06.2001, 30, 31.05.2009 (4, CHAH, TAMU) , same data as Holotype but 26.06.2001, Quercus arizonicus (1, CHAH) ; Chiricahua Mountains, Onion Saddle , 25.07.1987, W. F. Barr, beating Quercus (1, WFBM) , 04.06.1989, W. F. Barr, on Quercus (1, WFBM) , 11.06.1991, W. F. Barr, on oak (1, WFBM) ; Chiricahua Mts., Onion Saddle , 7600’, 31°56’N 109°16’W, 26.06.2001, H. A. Hespenheide, on Quercus hypoleucoides (1, CHAH) ; Huach Mts. , C. Schaeffer (4, BYU) , Carr Cyn., Huachua [sic] Mts. , 07.07.1930, J. O. Martin (1, CASC) ; Miller Can, Huachu. Mts. , 20.06, 15, 20, 29.07.1907, H. A. Kaeber (6, USNM) ; Huachuca Mts. , 21.7, 24.08.[19]05, Shoemaker (2, USNM) , 10.08.1905 (1, CASC) , 29.06.1984, W. B. Warner (1, FMNH) ; Huachuca M., 20.07.1937, D. J. & J. N. Knull (1, OSU) , 18.08.1936, J. N. Knull (1, OSU) ; Carr Cyn., Huachuca Mts. , 24- 25.07.1982, N. M. Downie (1, FMNH) , 07.1905, C. W. Leng (1, BYU) ; Copper Cn., Huachuca Mts. , 5.07.1977, S. McCleve (1, TAMU) ; Palmerlee , 30.6, 1, 10, 21.1907, H. A. Wenzel (10, USNM, OSU) , C. Schaeffer (1, BYU) ; [Coconino Co.], Oak Creek Canon, 6000 ft., Aug. F. H. Snow (1, USNM) , Oak Cr. Can. , 1.08.1938, D. J. & J. N. Knull (1, OSU) ; Williams , 24.7, Barber & Schwarz (1, USNM) ; Gila Co., Globe, Lwr. Six Shooter Cyn. , 24.06.1985, W. F. Barr (1, WFBM) ; Gila Co., Pinal Mts., Lwr. Ice House Cyn. , 24.06.1985, W. F. Barr, Quercus (1, WFBM) , 5 mi. SW Globe , 3.07.1941, B. E. White (1, CASC) ; [Maricopa Co.], Phoenix, C. Palm (1, AMNH) ; Wickenburg , 20.08.38, D. J. & J. N. Knull (1, OSU) ; Mohave Co., Hualpai Mts. , 5500-7000ft., ca. 10 mi. SE Kingman, 9.06.1958, J. M. Burns, S. N. Burns (1, EMEC) ; Pima Co., Organpipe Cactus National Monument, 10 mi. on Ajo Mountain Drive, 2400 ft., [no date], L. Bezark, K. Kitayama, J. Cope, R. Morrison, ex: Encelia sp. (1, EMEC) ; Santa Cruz Co., Madera Cn., 5500’, 13, 14.06.1961, C. D. Johnson (1, TAMU) , Sta. Rita Mts., Hopkins Rd. , km 11, 3.06.2006, N. Franz, C. W. O’ Brien, on mistletoe on oak (1, ASUHIC) ; Patagonia Mts. , 15.07.52, D. J. & J. N. Knull (1, OSU) ; Yavapai Co., Prescott , 15.06.1947, B. E. White, on oak (3, CASC) , 25.07.1950, D. J. & J. N. Knull (1, OSU) , “19.6,” Barber & Schwarz (1, USNM) , 28.06, 01.07.19, E. Schiffel (2, CASC) , Prescott N.F., 20.07.1937, D. J. & J. N. Knull (1, OSU) ; Hwy. 69, 3.9 mi. S jct. Hwy. 169, 4.06.1995, W. B. Warner, oak scrub (1, ASUHIC) . Colorado: Gunnison Co., Black Canon, 30.0741, H. S. Barber (2, USNM) ; Montezuma Co., 26.07.38, D.J.&J.N. Knull (1, OSU) . New Mexico: “N. M.” (1, MCZ) ; Sierra Blanca Mt. , 7.1963, Dr. Lenczy (1, USNM) ; Porvenir, A. Fenyes (1, CASC) ; Grant Co., Silver City , 07.1973, Dr. Lenczy (1, USNM) , Pinos Altos , ca.7000’, 8/ 10.07.2002, J. & A. Rifkind, P. Gum (1, WFBM) ; [Grant/Sierra Cos.], nr. Kingston, Mimbres Mts., Emory Pass , 8,178 ft, 19.06.1948, H. S. Barber (1, USNM) ; Lincoln Co., 2.7 mi. N Alto , 7200’, 19.06.1982, G. A. P. Gibson, sweeping oak (1, CMNC) ; Ruidosa , 31.07.1983, W. F. Barr (1, WFBM) ; Otero Co., Cloudcroft , Wickham (1, USNM) , “6/11/02,” Knaus (1, ASUHIC) , 12.07.1984, W. F. Barr, Quercus (1, CMNC; 1, WFBM), 10.06.1902, L. S. Slavin (3, CASC) ; 3 km W Cloudcroft , 2400 m, 24.08.1992, H. & A. Howden (1, CMNC) ; 15.2 mi. E Cloudcroft, James Canyon Cpgd. , 6800’, 24.07.1982, R. S. Anderson, sweeping oak-locust (1, CMNC) ; [Sandoval Co.], Jemez Mts. , 3.07, J. Woodgate (2, CASC) ; [San Miguel Co.], Las Vegas HS, 3, 9, 12.8, Barber & Schwarz (3, USNM) ; Santa Fe Co., Canada de los Alamos , 8 mi. SSE Santa Fe, 2360 m, 35°36’N 105°51’W, 23.06.2000, H. A. Hespenheide, Quercus (1, CHAH) . Texas: Brewster Co., Big Bend National Park, Lost Mine Trail , 5600-6850’, 15.07.1982, G. A. P. Gibson (2, CMNC) ; BBNP, (lower) Emory Peak Trail , ca. 7,050’, 29°14’57”N 103°17’47”W, 21.06.2004, E. G. Riley-51 (5, TAMU) ; BBNP, Laguna Meadow-Colima Trails , 6,500 7,000’, 29°14’29”N 103°18’36”W, 2.08.2003, E. G. Riley (13) (1, TAMU) ; BBNP, (lower) Pinnacles Trail , ca. 5,900’, 29°15’32”N 103°17’52”W, 21.06.2004, E. G. Riley-48 (1, TAMU) ; Panther Pass , Big Bend N. P., 19.07.1984, W. F. Barr, on Quercus (1, WFBM) ; Big Bend State Park , 12.07.1941, B. E. White (1, CASC) ; Chisos Mts. , 18.07, J. W. Green (2, CASC) , 8.07.1955, D. J. & J. N. Knull (1, OSU) ; Culberson Co., Pine Springs , 12.06.1971, C. O’ Brien. night, Quercus (1, ASUHIC) ; Jeff Davis Co., 18 mi. W Ft. Davis , 8.07.1961, R. L. Westcott, hostplant: Quercus (2, ASUHIC) ; Davis Mts. , 3 mi. SE Ft. Davis, 25.06.1964, E. M. Fisher (1, CHAH) ; Jeff Davis Co., 20.06.1952, 6.07.1953, D. J. & J. N. Knull (5, OSU) ; Davis Mts. , 2.07.1940, D. J. & J. N. Knull (3, OSU) , 09.07, J. W. Green (1, CASC) ; Davis Mt. , 11.07.1958, W. F. Barr (5, CMNC, TAMU, WFBM) ; Madera Cyn. , Davis M., 11.06.1968, R. & L. Hamilton (2, CHAH) ; Davis Mts., nr. Sawtooth Mt. , 10.07.1970, D. J. & J. N. Knull (1, ASUHIC) ; 1 mi. E McDonald Observ., 04.07.1987, R. Morris (1, CMNC); 16 mi. W Ft. Davis , 7.07.1961, R. L. Westcott (2, LACM) ; 18 mi. W Ft. Davis , 8.07.1961, R. L. Westcott (4, LACM, UCDC) . Mexico: Chihuahua: Canon Prieto nr. Primavera , 6500-6800 ft., 2.07.1947, Cazier, D. Rockefeller Exp. (1, AMNH) . Durango: 5 mi. NE Donato Guerro , 6100’, G. Brothers, on Acacia (1, ASUHIC) , 9 mi. W Durango , 9.06.1967, [A. Moldenke], on Quercus undulata (1, USNM) .

Hosts. Adults have been collected on leaves of Q. gambelli , Quercus emoryi Torr. , Quercus arizonica Sarg. , Quercus hypoleucoides A. Camus , Quercus undulata Torr. , and undetermined species of oaks as well as on Encelia sp. (Asteraceae) , on mistletoe on oak, and a dubious record on Acacia sp. (Fabaceae) .

Etymology. This species is named for the two red-orange elytral spots characteristic of the L. nephele group of species.

Discussion. I originally had thought that specimens listed here might represent two different species, but lack of consistent differences and similarity of male genitalia led my conclusion that they represent a single, widely distributed, morphologically variable species. Laemosaccus bimaculatus is a medium to large species with large, red posthumeral spots that usually touch the anterior margins of the elytra on elytral intervals 3–6 (infrequently 2–6 or 4–6) and extend beyond the middle. The pronotum can be very gibbous and is finely punctate. The male metasternum may be uniformly and densely setose, or sparsely setose in a postero-medial area with longer, semierect, slender setae. Some specimens have a narrow line of setae on the basal half of the inner margins of the eyes. Males vary in size from 2.85 to 5.20 mm (mean = 3.76 mm, n = 61); females vary from 2.90 to 5.80 mm (mean = 4.05 mm, n = 79).

Figures

Figs. 7–12. Laemosaccus species. 7–8) L. andersoni, dorsal and lateral habitus, respectively; 9) L. arizonensis, dorsal habitus; 10) L. bimaculatus, dorsal habitus; 11–12) L. burkei, dorsal and lateral habitus, respectively.

Figs. 19–24. Laemosaccus species, dorsal and lateral views of aedeagus and paramere. 19) L. andersoni; 20) L. arizonensis; 21) L. bimaculatus; 22) L. burkei; 23) L. clytrinoides; 24) L. howdenae.