taxonID	type	description	language	source
03CD2A013179BD043BE0F9019E4DA809.taxon	materials_examined	Type species: Knowltonia biramosa Fisher, by original designation. Gender: feminine.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A013179BD043BE0F9019E4DA809.taxon	description	Description. Head with clypeus broadly, shallowly, triangularly to arcuately emarginate; eyes separated on vertex by 1.2 to 2.2 times the eye width at middle of head. Antenna with 11 antennomeres, none being laterally compressed; first antennomere elongate and expanded apically; second antennomere short and globular; third antennomere of male elongate, strongly, sometimes acutely expanded on one side or weakly bifurcate at apex, of female elongate, expanded apically with the outer angle distinctly acute to right-angled; antennomeres 4 – 10 of male densely setose, biramous or slightly to strongly asymmetric, of female short with ventral lobes moderately to widely separated, truncate to slightly rounded apically; antennomere 11 of male forked or asymmetric at apex, of female elongate-oval, usually truncate to subtruncate, or sometimes vaguely acute at apex. Pronotum with front margin truncate to feebly bisinuate and broadly, shallowly arcuate at middle; disk subflattened, laterally convex or swollen, lateral margins ventrad, widely and distinctly indicated, entire or disappearing shortly before front angles, not visible from above. Elytra without or with vague longitudinal carinae, lacking foveae and pubescence; basal margin of each elytron strongly, evenly angulate; lateral margins finely, indistinctly serrate; sutural margins terminating in a tiny acuminate spine or a right-angled tooth; apices broadly rounded, not attaining apex of abdomen, exposing in part 1 or 2 abdominal terga. Legs with profemoral tooth elevated, acute to very narrowly rounded at apex, not serrulate; foretibia of male unmodified. Underside with prosternum flattened at middle, swollen at sides, front margin subtruncate to broadly arcuate, narrowly reflexed or with a reflexed lobe at middle; abdomen with lateral margins entire, ventrites without distinct lateral callosities, fifth ventrite lacking an elevated submarginal ridge.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A013179BD043BE0F9019E4DA809.taxon	discussion	Discussion: Fisher (1935) described Knowltonia and related it to Chrysobothris. However, he did not discuss any distinguishing character except to say “ .. differs from all the known genera of Buprestidae in having strongly biramose antennae. ” He assumed that the single specimen before him was a male. In his paper describing C. alleni, Cazier (1938) discussed Knowltonia and regarded it as a synonym of Chrysobothris, although we consider his reasoning faulty because it was partly based on incorrect association with such species as C. prasina Horn and C. platti Cazier (the latter presently in Sphaerobothris Semenov-Tian-Shanskij & Richter), and he lacked sufficient material for such a diagnosis. Fisher (1942) formally synonymized Knowltonia under Chrysobothris. His rationale was that he had overlooked Horn’s (1886) male antennal figure for C. atrifasciata LeConte, and that “ .. more material of Chrysobothris has been available for study and this peculiar character of the males has been found in atrifasciata, biramosa, and alleni .. ” To our way of thinking that is no reason for synonymy; it simply enlarges the genus Knowltonia. We are unaware of further published data under the genus Knowltonia, though Bellamy and Volkovitsh (1997) briefly mentioned it. For some years we and some of our colleagues have considered that it should be recognized at least to subgeneric level. Meanwhile those authors elevated the formerly Old World subgenus C. (Sphaerobothris) to genus rank, including in it the North American C. platti and C. ulkei LeConte. Superficially, those species resemble species of Knowltonia but are readily separated by their antennae, which are like those found in most species of Chrysobothris, with no sexual difference and the third antennomere not at all angulate apically. Although the facies of Sphaerobothris seems to distinguish it from Chrysobothris, nevertheless the apparent primary character used by Bellamy and Volkovitsh (1997) to separate them, the width on the vertex between the eyes, is not reliable. In Chrysobothris, this distance was stated to be “ equal or less than width of eye ”. We examined 25 North American species and found the width to vary from 0.31 – 2.08 times the width of the eye. We found this to range from 1.20 – 2.00 in 12 specimens of S. platti. The character does not appear to be sexual, and it may prove useful in separating some species, at least in Chrysobothris and Knowltonia. We found problems with other characters given in the key to separate Sphaerobothris from Chrysobothris, and subgenera of the former; however, that is beyond the scope of this paper. Nevertheless, we mention it to help justify our position with regards to Knowltonia, which is clearly separable, in a key or otherwise, based on the unique antennal structure of the male. To our knowledge, Fisher’s (1935) statement about that character remains true for New World species; however, two species in the unrelated Australasian genus Castiarina Gory and Laporte have a similar antennal modification. Members of the genus Knowltonia are moderately elongate and convex beetles, most being various shades of metallic green dorsally (Fig. 1), often with cupreous or, more rarely, bluish overtones; others vary from coppery to dark brown. The elytra usually have violaceous black markings, but may be immaculate. Ventrally, the color varies from light green to dark brown. All species of Knowltonia have been associated with the plant genus Atriplex, from which they have been reared and / or collected. However, there has been considerable misidentification of these beetles. Males are easily distinguished and separated from those of Chrysobothris and Sphaerobothris by their unique antennae. Other than by facies, which does not always readily separate them from the latter genus, females can almost always be separated from those genera by the shape of antennomere 3. We examined 117 specimens of Knowltonia and found antennomere 3 to be distinctly dilated ventrally (Fig. 6), usually produced triangularly or as a blunt tooth. Though variable, usually the antennae can be considered serrate from antennomere 3; however, on one specimen of K. calida (Knull) this is not so distinct, approaching the odd species of Chrysobothris in form. Antennomere 11 usually is distinctly narrower and more elongate than antennomere 10, not at all laterally compressed and usually more or less truncate at apex where sometimes it is distinctly acute. One specimen each of 57 species of Chrysobothris and two species of Sphaerobothris was examined, representing most if not all the presumed groups of the former found in North America. In all but three species of Chrysobothris the third antennomere is either essentially parallel-sided or slightly swollen apically. Of those three, said antennomere is indistinctly, in two cases more or less acutely, expanded ventrally (Fig. 9), but in no case would we term the antennae serrate from antennomere 3. Antennomere 11 is quadrate, rarely elongate-quadrate, almost always distinctly rounded apically, usually not narrower or more elongate than antennomere 10. The outer antennomeres, including antennomere 11 are laterally compressed in most species of Chrysobothris. Based on material examined, although most represents K. atrifasciata and K. calida, we suspect that additional taxa exist. However, considering general variability, the similarity of male genitalia, the lack of comprehensive samples of specimens from between known populations of the various phenotypes, and that even the limits between described species seem blurred, we feel this is not the appropriate time for descriptive work. It may be that Knowltonia is like the genus Crossidius LeConte (Coleoptera: Cerambycidae), species of which occur in the same habitats and are highly polytypic. Unfortunately, unlike most species of Crossidius, Knowltonia are not commonly collected. The following key applies almost entirely to populations in the U. S. and should be used with caution for specimens from Mexico. The user is strongly encouraged to use this key in conjunction with the species treatments to follow.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317ABD023BE0FAF79DB6ACC1.taxon	description	Wickham’s (1898) report of this species from Yuma, Arizona, Fisher’s (1942) records and Nelson’s (1959) record of it from California on A. lentiformis all refer to K. calida. The listings from Nevada by Fisher (1942) and California by Walters (1975) should instead be referred to K. biramosa. Beer’s (1949) rearing record from Atriplex sp. in Nevada was based on a misidentification of K. biramosa. We studied the specimen recorded as A. atrifasciata from Puerto Peñasco, Sonora, Mexico by Nelson et al. (1981). It is a female. Lacking the associated male, it is impossible to ascertain the species. However, we examined a male from a nearby locality and it comes closest to the green forms of C. biramosa. Without more specimens for study the listing of either species from Sonora is questionable.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317CBD003BE0FAC79A2EADE9.taxon	description	Knull (1958) distinguished this taxon from “ typical C. biramosa ” only by its dorsal green color. He made no mention of any other difference, and it is strange he overlooked the antennal character. The color no longer works well as a diagnostic character, though K. calida is uniformly deep green, sometimes with a bluish cast, while green forms of K. biramosa are lighter green, often with coppery reflections that are more notable at the base of the elytra. This species has been collected on Atriplex lentiformis (Nelson, 1959; 1962; 1966) and A. canescens (Nelson, 1960). Barr and Westcott’s (1976) treatment for this species should instead refer to K. biramosa.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317FBD003BE0FE9A9C49AF07.taxon	description	This species was described from the State of Durango, Mexico, from one specimen of unspecified sex. One of us (WFB) has long suspected that specimens collected in the U. S. attributed to this species represent a different taxon. We now consider it to be subspecifically distinct.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317FBD003BE0FD3F9B7BAB3C.taxon	description	Cazier (1951) recorded C. socialis from Sinaloa, undoubtedly based on two specimens (AMNH) lacking additional data. He considered them female, but his description of the last visible abdominal sternite and our examination clearly show them to be male. We have compared them to the holotype and consider all three specimens conspecific. We examined another specimen from Sinaloa, Estuilla Dist., Alamosa, 2 - X- 33, H. S. Gentry, CASC. There is an “ Alamos ” in that state, near the coast, but we cannot find mention of “ Estuilla ” anywhere. The only published record (Nelson et al., 1981) of this species from Sonora must be referred to the following new subspecies (see below). However, we examined two specimens of C. s. socialis from that state: Río Mayo, Arroyo Guajaray, 20 & 25 - X- 34, H. S. Gentry, CASC. The only female of this subspecies that we have seen is labeled “ Nayarit, Mexico, XI- 1917 ” (misidentified as C. trisignata Waterhouse), CASC. The elytra bear distinct, extensive coppery reflections and reduced markings. However, in every other respect, notably the color of the underside, it matches the other specimens we examined. Including the holotype, the six males we saw range in length from 10.8 – 13.5 mm, the female is 14.3 mm long.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317FBD013BE0F9C09D27A8B9.taxon	description	Specimens examined: Holotype labeled “ Ariz ” / HOLOTYPE Chrysobothris socialis apache Westcott and Barr ” (red card), and we select as allotype a specimen measuring 12.41 mm long by 5.14 mm wide, labeled “ Nogales, St Cruz Co, IX: 18.06 Ariz. / COLL’D BY F. W. NUNENMACHER / CharlesSchaeffer Collection / Shoemaker Collection 1956. ” The holotype and allotype are deposited in USNM. Paratypes as follows: “ Arizona ”, 1 F, 3 M, [no other data], AMNH, CLBC, USNM; 2 M, CharlesSchaeffer Collection / Shoemaker Collection 1956, USNM; 1 F, Henry Ulke Beetle Coll., CMNH Acc. No. 1645; 1 F, same, From Coll. of W. J. Chamberlin, CASC; 1 F, same, Van Dyke Collection, CASC. Cochise Co., 1 F, Naco, 2 - IX- 35, E. D. Ball, J. N. Knull Collection, FMNH; 2 M, Chiricahua Mts., Cave Crk. Can., Sunny Flat, 5100 ’, 31 ° 53 ’ N, 109 ° 10 ’ W, 28 - VIII- 80, at Mimosa biuncifera & 22 - VIII- 90, “ VERBENAC. ”, both H. A. Hespenheide, DSVC, FSCA; 1 F, 11 mi W Portal, 31 ° 55.636 N, 109 ° 15.350 ’ W, 6686 ’, 8 - VIII- 2000, A. Gilbert, CLBC. Pima Co., 1 F, Kitt Peak, IX- 1974, [no collector], CLBC; 1 F, 1220 m, 11 - IX- 82, Ex Baccharis, N. Schiff, Collection of R. L. Westcott, RLWE; 1 M, Madera Canyon, Santa Rita Mts., 5 - IX- 70, E. A. Kane, CSCA; 1 F, 7 - IX- 71, B. Villegas, RLWE; 1 F, 29 - VIII- 2000, J. F. Limón, on Mimosa sp., CLBC; 1 F, 8 - IX- 81, on Prosopis juliflora, W. F. Barr Collection, WFBM; 1 F, running on mesquite, Prosopis sp.; 8 - IX- 81, FSCA; 1 F, near waterfall, Proctor Ranch, 20 - IX- 64, J. N. Knull Collection, FMNH; 1 M, waterfall below Proctor Ranch rd., 16 - VIII- 70, on Anisocanthus thurberi, D. S. Verity, DSVC; 1 F, Santa Rita Mts., Box Canyon, 6 - IX- 70, F. Hovore. RLWE; 1 M, Santa Catalina Mts., Sabino Canyon, 26 - VIII- 59, G. H. Nelson, on Celtis pallida, FSCA. Santa Cruz Co., 2 F, Nogales 8 & 10 - IX- 1906, Koebele Collection, CASC; 1 M, 4.5 mi W Nogales, 9 - IX- 71, B. Villegas, Collection of R. L. Westcott. Mexico, 1 F, Sonora, 28 mi S Navojoa, 25 - X- 72, E. M. Fisher, RLWE. Comparison: Chrysobothris socialis apache differs from the nominate subspecies primarily by the ventral coloration, which is coppery bronze in the latter. Also in that subspecies, based upon only six specimens examined, the male face is entirely or predominantly copper — the punctures of the lower face may be green at the bottom — and the sides of the pronotum are distinctly and more widely reddish-copper, except near the hind angles where only one of the specimens examined shows any such color. In the only female of C. s. socialis that we examined, the face also is coppery, as contrasted to the green, blue-green or blue of C. s. apache, though more distinctly green below, and the sides of the pronotum are colored similarly to the male. Etymology: The subspecies epithet is chosen to honor the tribe of original Native Americans who inhabited the region in which this beetle lives.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317EBD0E3BE0F99F9C9CA843.taxon	materials_examined	Holotype male: length 7.97 mm, width 3.08 mm, widest at base of elytra across umbones, subequal in width across pronotum; shining black with small yellow elytral markings as in figure 3, several of which either reach or occur on margin, where the left apical — and right apical and subapical — spots are vaguely suffused with orange; setae on dorsum long, mostly black, especially on head above and anterior portion of pronotum, with some mixture of white, below white and moderately dense, more densely placed than above, especially at sides. Head shallowly convex, flattened above, coarsely densely punctate; clypeus depressed at base, front margin broadly deeply arcuately emarginate; antennae sharply serrate from 5 th antennomere, extending to procoxae. Pronotum broadly shallowly convex, shallowly transversely depressed immediately behind front margin, shallowly depressed on base at middle and obliquely so on each side anterior to deep basal foveae; punctures of disk coarse, shallower on middle, becoming deep and reticulately placed on sides; anterior margin broadly shallowly emarginate, indistinctly lobed at middle; posterior margin subtruncate; lateral margins well defined, except narrowly at base, visible from above only from apex to short distance behind, shallowly evenly arcuate; front angles subquadrate; hind angles quadrate, poorly defined. Elytra subflattened, strongly depressed at base between strongly raised 3 rd intervals, these intervals elevated for only short distance behind base; umbones extremely bold; humeral angles moderately, broadly triangularly projecting ventrad; lateral margins distinctly, moderately coarsely serrate on about apical 1 / 2; suture distinctly elevated along apical 2 / 3; punctures of discal area as in figure 3, much coarser and more closely placed laterad of strongly elevated 5 th intervals; 9 th intervals wider, strongly elevated from behind umbones to about apical 1 / 5, 10 th intervals flattened, marginal intervals slightly raised from base to about middle. Underside with front margin of prosternum having well developed blunt tooth on either side of subtruncate middle and sides transverse, ending well behind front angles of pronotum; abdomen with surface clearly visible throughout, not obscured by setae, punctures small to moderate on middle, becoming larger and more dense at sides; last visible sternite sparsely, evenly punctate, broadly rounded apically and with a very well developed preapical plate, the narrowly rounded apex of which almost attains the apical margin.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317EBD0E3BE0F99F9C9CA843.taxon	description	Specimens examined: holotype (EMEC) labeled “ 5 mi NW Ocotillo, Imperial Co. Calif, April 8 1956, R. R. Snelling / HOLOTYPE Acmaeodera audreyae Westcott and Barr. ” One male paratype (RLWE) with same data except taken on Larrea divaricata (creosote bush; family Zygophyllaceae). Variation: The paratype is 8.87 mm long, 3.34 mm wide, the only notable difference otherwise is that the preapical plate of the 5 th visible abdominal sternite does not so nearly attain the apical margin; nonetheless, it is very well developed.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317EBD0E3BE0F99F9C9CA843.taxon	discussion	Remarks: This species seems closest to A. inopinata Barr, a species that breeds in Haplopappus pinifolius (Barr, 1972). However, that beetle is less robust, not so black — usually exhibiting a slight metallic reflec-tion — and the 3 rd, 5 th and 9 th intervals are not so strongly elevated. Like A. inopinata, we suggest that the larva of A. audreyae works in one of the similar or unrelated low growing desert shrubs which at the type locality are very sparsely interspersed among the dominant plant, Larrea tridentata. It seems most likely that the association of A. audreyae with that plant is as a flower visitor, rather than it reflecting a larval host.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
03CD2A01317EBD0E3BE0F99F9C9CA843.taxon	etymology	Etymology: We take special pleasure in dedicating this species to Audrey K. Barr, loving wife of the junior author and longtime inspirational friend of the senior author.	en	Westcott, Richard L., Barr, William F. (2007): Resurrection and review of the genus Knowltonia Fisher, with description of a new subspecies of Chrysobothris Eschscholtz and new species of Acmaeodera Eschscholtz from North America (Coleoptera: Buprestidae). Zootaxa 1481: 49-58, DOI: 10.5281/zenodo.273778
