Laccophilus salvini Sharp 1882

Laccophilus salvini Sharp (Figs. 62-69, 291-292)
Laccophilus salvini Sharp, 1882a, p. 291. Holotype: male, British Museum (Natural History), Guatemala, Guatemala; Sharp, 1882b, p. 10; Zimmermann, 1920, p. 25; Blackwelder, 1944, p. 74.
DIAGNOSIS. — The combination of posterior elytral blotch, black venter, sawlike ovipositor, and file in the male and female should separate salvini from
salvini
below
Tehuantepec TO TAL LENGTH (mm) Males are shown crosshatched; females, stippled. all other North American Laccophilus. It differs from mexicanus and pseudomexicanus in its smaller size (average length, 4.15 to 4.30 mm as compared to average lengths of over 4.7 mm) and black elytral blotch, and from fuscipennis, fasciatus, and proximus by the black venter. Small females of mexicanus and large ones of salvini cannot always be reliably separated, since the blotch is sometimes weak and there is some size overlap. A mid-lateral elytral macula is almost always well-defined in salvini, but weak in mexicanus. Small specimens of /. fasciatus can be very similar dorsally, but the differently colored venter permits reliable separation. The aedeagi of proximus and salvini are similar enough to pose the question of whether they are specifically different. DESCRIPTION. — Medium (length 3.9 to 4.8 mm; width 2.3 to 2.7 mm), dark brown, irrorated species; black beneath; metacoxal file prominent in males and weakly suggested in females; prosternal process short; ovipositor sawlike. COLOR. Head: pale brownish-yellow above and beneath, slightly darker at base of pronotum between the eyes; appendages yellow except for reddishbrown mandibles. Pronotum: pale brownish-yellow. Elytra: pale brownishyellow background with irrorated pattern distinctly outlined (figure 291); strong tendency to completely suffuse and coalesce in the posterior half to form a nearly complete transverse fascia or blotch in some specimens; apex darkened, but usually a large clear area immediately anterior to apex; epipleura pale anteriorly and dark brown posteriorly. Tergite VIII: dark brown to black. Venter: prosternum, its process, prolegs, and mesolegs pale brownish-yellow; hind legs and postcoxal processes reddish-brown, darker at edge and on tarsi; mesothorax, metathorax, and metacoxal plates black; abdominal sternites from yellowish-brown to reddish-black with all degrees of intermediacy; males usually darker than females. Genitalia: generally reddish-brown with varying degrees of yellow. ANATOMY. Microreticulation: weakly double on head, pronotum, and elytra; secondary mesh discernible, but individual cellules still apparent. Head: supraclypeal seam closely parallel to margin. Pronotum: WH PW, 0.70; LP PW, 0.40. Elytra: epipleural flange small and seldom present: truncation slight. Venter: coxal file finely present in males, composed of about 28 to 30 lines; weakly present in females, but too fine to count; lobes of postcoxal process rounded laterally and projecting well beyond the midline; male and female last visible abdominal segments rounded, not truncated, and similar in outline: that of males slightly produced; female with faint groove on either side forming a marginate edge; crest fairly well-defined; asymmetrical in males; broad ridge in females; several rugae near the posterior margin; setigerous punctures thickest near the apex. Legs: male proand mesotarsi expanded in a dorsoventral plane; palettes easily visible at 20 power magnification; fifth tarsal segment of both pair of legs about one and one-half times as long as corresponding fourth; profemoral setae (5 to 6) shorter and finer than mesofemoral ones (5 to 6). Genitalia: oval plate with long acuminate tip and a well-defined median crest which has little anterior curvature; weak raised lines on either side: aedeagus evenly curved, narrowing toward tip; without distinctive knobs
or sculpturing; right paramere with blunt apex; apex of left paramere slightly asymmetrical; ovipositor with about 13 sawlike teeth.
NOMENCLATURAL NOTES. — Confusion in this taxon has been limited to it being mistaken for L. confusus in Arizona (Leech 1948a). The specimens were somewhat lighter than usual beneath; and since salvini and proximus ( = confusus) are so similar, it is not surprising that the Mexican synonym was given to those individuals.
VARIATION. — The most apparent pattern variation is the degree to which the posterior part of the elytra has darkened. Some have a fascia or blotch nearly as complete as that of L. f. fasciatus, while others have little enough to be confused with me.xicanus. The darkening is stronger in specimens from south of the Isthmus of Tehuantepec. Arizona specimens have lighter venters than other salvini and, except for the elytral pattern, are very close to proximus.
Populations from south of the Isthmus are larger than those to the north. A comparison of mean pronotal widths for males from Morelos and Las Casas, Chiapas, which are the closest samples on either side of the Isthmus, give a highly significant difference (Table 4). The samples from Comitan, Chiapas, and Guatemala agree with the Las Casas population. One could readily justify describing the northern populations as a different race, but I have not done so. The population from Arizona appears even more different than that from Central Mexico, but only ten specimens were available for comparison. The species shows considerable geographic variation and deserves additional descriptive analysis (Fig. 8). — L. salvini occurs from southern Arizona to Guatemala. At present there is a gap in the records from northern Sonora to Nayarit. Too few collections have been made on the west edge of the Sierra Madre Occidental to consider this more than a collecting gap, however. It is common in Nayarit, Jalisco, Michoacan, and Morelos, and further south in Chiapas and Guatemala. It generally is found between two and seven thousand feet (Fig. 7).
Like proximus it seems to be a pioneer species in newly formed situations at higher altitudes. It seldom occurs on the coastal lowland. Grassy margined ponds are the most common habitat.
UNITED STATES OF AMERICA. — ARIZONA. Pima County. Tucson, Bear Can., 1 4, iii. 18.46; 2 4 , 3 2, iii.26.46, J. W. Green (CAS). Catalina Mtns., 2,5,29, iv.18.— , Hubbard & Schwarz (USNM); 1 9, xi.8.34, K. Thomas (NMSU).
GUATEMALA. — Carcha (Dept. Alto V. P.), 9 6, 2 9, vi.23.66, Flint & Ortiz: Chimaltenango, 20 m. NW„ 48 4 , 40 9, viii. 20.65, P. J. Spangler: Guatemala City, 20 m. S., 10 8, 7 9, vii.7.65, P. J. Spangler (USNM).
MEXICO. — CHIAPAS. Buchil (near), 1 8, viii.31.63; Comitan, 92 8, 69 9, viii.30.63; 13 m. W., 38 8, 40 9, viii. 30.63; Ixtapa, 15 8, 6 9, viii.31.63; Ocozucuatla, 6 m. W., 1 8, ix.1.63, JRZ (NMSU). San Cristobal de las Casas, 15,19, i.25.66, P. H. Raven & D. E. Breedlove (CAS); 38 8, 17 9, viii.28.63; 5 m. E., 17 8, 7 9, viii.28.63; 20 m. W., 3 8, 1 9, viii.30.63; Tuxtla Gutierrez, 10 m. W„ 6 4,5 9, ix.1.63, JRZ (NMSU). COLIMA. Trapechi (near Colima), 2 8, 6 9, vii.29.62; 15 8, 22 9, vii.30.62, JRZ (NMSU). JALISCO. Ameca, 25 km. E., 43 8, 51 9, iii.25.64, JRZ (NMSU). Atenquique, 15 m. NE., 3 8, 3 9, xii.5.48, H. B. Leech (CAS). Autlan, 53 8, 38 9, x.26.66, A. H. Smith & JRZ (NMSU). Barra de Navidad, 1 8, iii.18.61, C. O. Morse; Colima, 29 m. NE., 3 4,19, xii.3.48, H. B. Leech (CAS). Cuidad Guzman, 5 m. N., 5 8, 1 9, vii.28.62; Guadalajara, 20 m. E., 4 8, 1 9, iii.25.64; 11 m. S., 8 8, 1 1 9, vii.30.62, JRZ (NMSU). Jiquilpan, 15-20 m. W., 1 8, 1 9, xi.30.48, H. B. Leech (CAS). La Huerta, 6 m. N., 84,89, x.25.66, A. H. Smith & JRZ (NMSU). Mazamitla, 17 m. S., 10 8, 8 9, xii.5.48, H. B. Leech (CAS); 5 km. E., 1 4,2 9, iii. 27.64, JRZ (NMSU). Tala (near), 27 4, 17 9, iii.25.64; Tamazula, 5 4,6 9, iii.27.64; Tecalitlan, 10 m. S., 6 4, 13 9, iii.27.64, JRZ; Tecolotlan, 12 4, 12 9, x.24.66, A. H. Smith & JRZ; Tehetlican (Rd. 110), 12 4, 12 9, vii.28.62; Tizapan (near), Rd. 15, 26 4, 17 9, iii.26.64, JRZ (NMSU). Tlaquepaque, 2 9, vii.— .53, N. L. H. Krauss (CAS). Union de Tula, 10 m. S., 9 4, 18 9, iii.28.64, JRZ (NMSU). La Venta de Mochitiltic, 5 4, 7 9, vii.20.55, R. B. Selander (CAS). MEXICO. Near Tonatico, 14, 2 9, viii.29.62. JRZ (NMSU). MICHOACAN. Carapan, 15 m. S„ 2 6 , xii.7.48, H. B. Leech (CAS). Jacona, 1 9, viii. 11.57, D. R. Lauck (USNM). Jiquilpan, 8 km. E., 114, 10 9, iii.2.64; 11 m. W., 2 4, vii.28.62; Uruapan, 2 4, 19, iii.26.64; 10 m. S., 10 4, 16 9, iii.26.64, JRZ (NMSU). Zamora, 9 m. W, 3 3,19, xii.6.48, H. B. Leech (CAS); 2 9, vii.28.62, JRZ (NMSU). MORELOS. Acatipaca, 4 4,3 9, viii.29.62; Cuautla (near), 10 4, 8 9, viii.28.62, JRZ (NMSU). Cuernavaca, 14 m. S., 2 4 , xii.8.48, H. B. Leech (CAS); 1 9, iv.15.46, J. D. Pallister (AMNH); 2 4,29 (UMMZ). Puente de Ixtla, 34,29, viii.29.62; Temixco, 3 4,2 9, viii.29.62, JRZ (NMSU). NAYARIT. Ixtlan del Rio, 34 4, 21 9, ix.22.53; San Bias, 14,2 9, ix.17-21.53, B. Malkin (CAS); 5 m. E, 2 4, 1 9, vii. 3 1.62, JRZ (NMSU). Sierra de Zapotan, 1 4, xi.— .42, Eugenio Paredes (CAS). Tepic, 1 4, ix.15-17.53; 3 4, 3 9, ix.21-24.53; 19 m. SE., 14,7 9, ix.24.53, B. Malkin (CAS); 2 9, xi.27.48; 15 m. SE., 16,6 2, xi.28.48; 25 m. SE., 6 6, 11 2, xi.23.48, H. B. Leech; 20.5 m. NW., 10 6, 3 2, xi.24.48, H. B. Leech (CAS). OAXACA. Camotlan (near), 3 S, 4 2, ix.3.64; Huajuapan, 2 km. S., 4 6, 2 2, ix.4.64, JRZ (NMSU). PUEBLA. Amatitlan, 9 m. N., IS, xii. 10.48, H. B. Leech (CAS). Puebla(?), 1 S, vi.26.57, D. R. Lauck (USNM). SONORA. Aribaba (near Moctezuma), 16,12, xii. 15.62, JRZ (NMSU). Hermosillo, 3 2, vii.9-16.53, B. Malkin (CAS). Near Rancho Uriquepa, 1 2, vi.26.56, F. N. Young (UMMZ). ZACATECAS. Fresnillo, 16 m. NW., Rio Trujillo, 6600 feet, 1 2, vi.29.54, R. H. Brewer (CAS). This polytypic species is composed of three races in North America. It appears to be part of a complex that extends to southern South America. Laccophilus chilensis Sharp bears strong resemblance to this group. As yet, no intergrade zones have been found between the North American races. One suspected intergrade has been seen between mexicanus and atristemalis. It is a female, however, and does not permit a reliable determination of intergradation. The races are very much alike, and individuals of the two races mentioned above can be separated only on the basis of male genitalia. The southern race, oaxacensis, does show more differentiation, and details of pattern and size permit good separation between it and mexicanus. The ranges of the southern races are, apparently, completely allopatric with a gap of about 200 miles.
DESCRIPTION. — Medium to large (length, 4.2-5.4 mm; width, 2.4-3.1 mm) species, brown above and dark brown or black beneath; metacoxal file present; prosternal process short; ovipositor sawlike. COLOR. Head: it and its appendages yellow or light brown, darkening to light reddish-brown near the pronotum: mandibles also darkening toward tip. Pronotum: yellow to yellowishor reddish-brown. Elytra: light yellowish-brown background with relatively even irrorations, anterior fingerlike detail of maculosus group generally lacking in mexicanus and atristemalis, but evident in oaxacensis: some tendency for suffusion and anastomosing of dots in posterior half and near the medium suture; dots frequently forming chains; suture dark brown; pattern limits ill-defined: epipleura pale anteriorly, dark in constricted posterior part. Tergite VIII: dark brown to black. Venter: prosternum, its process, fore and middle legs light brownish-yellow with reddish tinge; hind legs of about the same color with much stronger reddish highlights; mesosternum, metasternum. metacoxal plates, except postcoxal processes, very dark brown or black; abdominal segments variable from brownish-yellow to black; females with abdominal segments frequently pale except for the black hind margin of first visible segment, but other segments tend to be suffused with some dark brown or black.
Genitalia: oval plate, aedeagus, and parameres with varying amounts of dark reddish-brown or yellowish-brown; ovipositor reddish-brown. ANATOMY. Microreticulation: finely meshed and only weakly double at best (finer and less impressed than in maculosus) on head, pronotum and elytra. Head: supraclypeal seam closely parallel to margin. Pronotum: WH'PW, 0.67 to 0.68; LP/PW, 0.40 to 0.41. Elytra: epipleural flange rare in atristernalis and mexicanus, but common in oaxacensis; truncation of elytral apices slight. Venter: coxal file prominent in males, composed of from 25 to 30 lines, laterally 10 or 15 more which are much weaker and not distinctly part of the file; a faint trace of file apparent in females; prosternal process with well-defined crest; lobes of postcoxal processes rounded and laterally projecting well beyond the midline; male last visible ventral abdominal segment slightly produced with or without rugae along hind margin; asymmetrical curving ridge on middle forming depression on left side or both sides; female last segment either sub triangular or rounded; slight crest in posterior half of segment. Legs: male proand mesotarsi noticeably enlarged in a dorsoventral plane, easily observed at 20 power magnification; fifth tarsal segment on proand mesolegs from one and threequarters to twice as long as fourth; profemoral setae (5 to 6) shorter and finer than mesofemoral ones (5 to 7). Genitalia: oval plate with acuminate tip and well defined crest which anteriorly curves to the left; numerous raised lines on either side of the crest; aedeagus of complex sculpture and abruptly constricting near the apex; right paramere with small distinct conical apex; ovipositor with 13 to 15 sawlike teeth.
NOMENCLATURAL NOTES. — The biological situation in L. mexicanus seems to be a relatively uncomplicated problem, but the nomenclatural condition has been one of continuous confusion. This is understandable, since the species lacks the pattern and colors that might lead one to easily recognize and define the geographical races. Aube described mexicanus in 1838 from a single specimen from Mexico. No further locality information was given. I have not seen his type. It is apparently not at the Institute Royal des Sciences Naturelles de Belgique in Brussels or in the Paris Museum. It appears to be lost; but since the writer has not visited the European museums that might contain the type, a neotype has not been designated in this paper. I have seen a female specimen from Guanajuato (Brussels Museum) compared by Regimbart, with the type, and I have also examined a male in the Paris Museum from the collection of Regimbart; from Durango, Mexico; both are what have been considered mexicanus. Without examination of the type, there cannot be certainty about the identity of mexicanus. The specimens from Guanajuato and Durango belong to a race that extends to south of Mexico City and north into the southwestern United States. Two other races occur in North America, however.
Crotch (1873) described atristernalis from California. Sharp and Horn synonomized it with mexicanus. Leech (1948) maintained that mexicanus specimens from Oaxaca were different from the California and Lower California specimens. He mentions the elytral apices as pale in those from Oaxaca and dark in the ones from north of there. The male aedeagus is also different and supports Leech's observations. The specimens from Oaxaca are being described as a new subspecies. The material from California is heterogeneous, however, since there are two races in that state. The one in southern and Lower California is what is here considered to be typical mexicanus. The one from northern California, Oregon, Idaho, Nevada, and Utah is being retained as atristernalis. I have not been able to find Crotch's type of atristernalis, and California has proven to be too vague a locality. The type locality is being restricted to Woodside, San Mateo, California; and the type locality for the nominate race is being restricted to Guanajuato, Guanajuato, Mexico.
Aube's mexicanus could have been either of the two races that occur in Mexico; but since there is no certain way of establishing which one he had in hand, it seems better to describe as new the race that has the more restricted distribution and retain mexicanus for the race that is common over most of the country. There appears to be no confusion of mexicanus with any other species of North American Laccophilus.