Eurycipitia distanti Reuter 1905: 4 Caulotops rufoscutellatus Carvalho 1948: 530 Carvalho and Froeschner 1987: 148 Carvalho 1954: 424 Caulotops rufoscutellatus major Carvalho 1948: 532 Carvalho and Froeschner 1987: 48 Henry 1985: 321 Caulotops distanti Carvalho 1957: 94 Henry 1985: 321 Henry and Wheeler 1988: 265 Schuh 1995: 540 Wheeler 2001: 152 Figueroa-Castro et al. 2019: 642 Caulotops distanti major : Carvalho 1957: 94 Revision and Phylogeny of the Eccritotarsine Plant Bug Genus Caulotops Bergroth, with Descriptions of Four New Genera and 14 New Species (Hemiptera Heteroptera: Miridae: Bryocorinae) Associated with Agave (Agavoideae Asparagaceae) and Related Plant Genera Henry, Thomas J. Menard, Katrina L. Zootaxa 2020 2020-05-08 4772 2 201 252 (Reuter) Henry & Menard 2020 Reuter [151,511,153,180] Insecta Miridae Agaveocoris Animalia Hemiptera 18 219 Arthropoda species distanti comb. nov.  ( Figs. 24–28, 83–85, 124)      Eurycipitia distanti Reuter 1905: 4(original description), 1908: 155 (note); Carvalho 1952(as typeof genus in error).     Caulotops rufoscutellatus Carvalho 1948: 530(original description);  Carvalho and Froeschner 1987: 148( typelist). Synonymized by  Carvalho 1954: 424.     Caulotops rufoscutellatus major Carvalho 1948: 532(original description, n. ssp.);  Carvalho and Froeschner 1987: 48( typelist). Synonymized by  Henry 1985: 321.     Caulotops distanti Carvalho 1957: 94(catalog), 1985: 496 (description, synonymy);  Henry 1985: 321(key, note, host);  Henry and Wheeler 1988: 265(catalog);  Schuh 1995: 540(catalog);  Wheeler 2001: 152(habits, injury); Schuh 2002–2013 (online catalog);  Figueroa-Castro et al. 2019: 642(host, distribution).     Caulotops distanti major: Carvalho 1957: 94(catalog).   DIAGNOSIS.  Agaveocoris distanti( Figs. 24–28), as we define it, is the most variable species of the genus. It is distinguished by the head, pronotum, and legs ranging from pale yellowish brown to red or reddish orange and the hemelytra from dull brown to dark brown tinged with red and usually with a crescent-shaped red to reddish-orange mark on each corium. It can be separated from all other species by its relatively small size, the length of the second antennal segment subequal to the interocular width, the pale yellow to reddish-orange femora lacking or having only a few tiny indistinct spots, and the shape of the right paramere. This species keys to couplet 9 with  A. roseus, n. sp.based on the small size, the dark hemelytra often tinged or marked with red or reddish orange, and the femora lacking distinct spots. It differs from  A. roseusin lacking or having only indistinct marking on the frons and calli, and the corium usually with a distinct red or reddish-orange wedge-shaped mark and hemelytra with a bluish sheen, whereas  A. roseushas the hemelytra more uniformly dark rosy red and the apex of the right paramere is more strongly turned upward.   REDESCRIPTION. Male(brown color form, figs. 24, 25): (n = 5): Length from apex of head to cuneal fracture 2.30–2.62 mm; length from apex of head to apex of membrane 3.33–3.65 mm; widest point across hemelytra 1.38– 1.62 mm. Head: Width across eyes 0.93–1.04 mm; interocular width 0.56–0.62 mm. Labium: Length 1.31–1.52 mm, extending well past hind coxae to abdominal segment III or beyond. Antenna: Segment I length 0.39–0.45 mm; II, 0.54–0.66 mm; III, 0.50–0.51 mm; IV, 0.45–0.48 mm. Pronotum: Median length 0.54–0.64 mm; posterior width 1.12–1.28 mm.  Male(reddish-orange color form, fig. 27) (n = 5): Length from apex of head to cuneal fracture 2.53–2.75 mm; length from apex of head to apex of membrane 3.04–3.68 mm; widest point across hemelytra 1.50–1.60 mm. Head: Width across eyes 0.96–1.01 mm; interocular width 0.58–0.62 mm. Labium: Length 1.38–1.54 mm. Antenna: Segment I length 0.40–0.48 mm; II, 0.58–0.66 mm; III, 0.48–0.56 mm; IV, 0.35–0.48 mm. Pronotum: Median length 0.61–0.67 mm; posterior width 1.18–1.34 mm. General coloration pale grayish yellow to red or reddish orange. COLORATION. Head:yellowish to red, frons and vertex usually with only indistinct brown transverse stripes, more distinct on some specimens; clypeus and labrum dark brown. Labium: Pale yellowish brown, segment III and IV darker brown. Antenna: Segment I pale yellowish brown, with a distinct fuscous sub-basal ring; remaining segments pale brown to dark brown. Pronotum: Pale yellowish brown tinged with orange in pale forms to red or reddish orange in darker forms; calli concolorous with remainder of pronotum in pale forms to heavily infuscated in darker forms. Mesoscutumand s cutellum: Bright yellow or yellowish orange in pale forms to red in dark forms; lateral sclerites of thorax same coloration as posterior pronotum in both forms. Hemelytron: Yellowish blue-gray in pale forms to dark purple in dark forms, often with a weak bluish sheen, corium usually with a crescent-shaped red or reddish-orange mark to more broadly tinged with reddish orange to rosy red of corium; membrane clear to translucent smoky brown, veins dark brown. Ventral surface: Thorax yellowish brown to dark reddish brown, mesosternum shiny brown to dark brown; abdomen pale yellowish brown in pale forms to dark brownish or reddish orange in dark forms, margins of segments often darker brown in both color forms, apex of genital tubercle pale yellow with apex dark to uniformly dark. Legs: Coxae pale yellow to reddish orange; femora pale yellow to reddish orange, without or with only a few indistinct spots; tibiae pale yellowish brown or reddish orange; tarsomeres pale brown; claws darker brown. SURFACE AND VESTITURE. Hemelytron has weak bluish sheen; rest of surface and vestiture characters as in generic description. STRUCTURE. Interocular width greater than twice width of one eye, first antennal segment length shorter than interocular distance; length of antennal segment I approximately three fourths the length of segment II, segment II narrower than I, length of segment II less than or equal width interocular distance, antennal segments III and IV half width of segment I, individually subequal in length to segment I, apex of labium extending past meta-coxae; posterior margin weakly concave; remaining characters as in generic description.  Male genitalia: Endosoma ( Fig. 83) as in generic description. Left paramere ( Fig. 84) small and C-shaped. Right paramere ( Fig. 85) elongate, shallowly L-shaped, with a short upturned apex.  Female(brown color form, fig. 26) (n = 5): Length from apex of head to cuneal fracture 2.37–2.85 mm; length from apex of head to apex of membrane 3.42–3.92 mm; widest point across hemelytra 1.49–1.73 mm. Head: Width across eyes 0.99–1.10 mm; interocular width 0.59–0.66 mm. Labium: Length 1.26–1.57 mm. Antenna: Segment I length 0.42–0.45 mm; II, 0.51–0.67 mm; III, 0.48–0.54 mm; IV, 0.45–0.53 mm. Pronotum: Median length 0.54– 0.67 mm; posterior width 1.17–1.31 mm. Female (reddish-orange color form, fig. 28). (n = 5): Length from apex of head to cuneal fracture 2.64–2.76 mm; length from apex of head to apex of membrane 3.84–4.00 mm; widest point across hemelytra 1.60–1.76 mm. Head: Width across eyes 0.99–1.02 mm; interocular width 0.61–0.64 mm. Labium: Length 1.46–1.57 mm. Antenna: Segment I length 0.45–0.48 mm; II, 0.59–0.64 mm; III, 0.45–0.51 mm; IV, 0.37–0.38 mm. Pronotum: Median length 0.64–0.74 mm; posterior width 1.26–1.42 mm. Similar to male in size, shape and coloration. Genitalia as in generic description.   HOSTS.This species has been recorded from  Yuccasp.,  Yucca giganteaLem.[as jr. synonym  Yucca elephantipesRegel ex Trel] [ Agavoideae: Asparagaceae] ( Henry 1985) and  Agave cupreataTrel. & Berger[ Asparagaceae] and  A. viviparaL. ( Figueroa-Castro et al. 2019). From specimen data, this species has been taken on  A. fourcroydesLem.,  A. cerulataTrel.[unknown subspp. subcerulata],  A. murpheyiGibson, and  A. viviparaL. [as jr. synonym  Agave angustifoliaHaw.]. A long series from Colombiawas collected on  Furcraeasp. [ Asparagaceae]. One specimen, taken on  Opuntiasp. [ Cactaceae], clearly is an incidental record.   DISTRIBUTION.Described from Venezuelaas  Eccritotarsus distanti( Reuter 1905)and as  Caulotops rufoscutellatus( Carvalho 1948)and later reported from Costa Rica, Mexicoas  C. rufoscutellatus major( Carvalho 1948)and Florida in the United States( Henry 1985, Henry and Wheeler 1988). New country records are Colombia, and El Salvador. New state records for Mexicoare Chiapas, Guerrero, Oaxaca, Puebla, San Luis Potosi, Tamaulipas, and Yucatan. New U. S.state records are Arizonaand Texas.   TYPE MATERIAL EXAMINED.  Holotype ♂(  Caulotops rufoscutellatusCarvalho): Label 1, “Los Teques, Venez 3- 14-38, C. H. Ballou, No. 21”; 2, “on Fourcroya humboltiana”; 3, “ ♂“; 4 (red), “Typus”; 5 (red), “Type No. 59185 USNM”; 6, (handwritten), “ Caulotops rufoscutellatus n. sp., JCMCarvalho, 1948” ( USNM).   Holotype ♂(  Caulotops rufoscutellatus majorCarvalho): Label1, “On Agave, Merida, Yucatan, EJHambleton, 9-13-46-16924”; 2, “ ♂”; 3 (red label), “Typus”; 4 (red label), “Type No. 59186 USNM”; 5 (handwritten), “Caulo- tops rufoscutellatus major n. sp., JCMCarvalho 1948” ( USNM).  OTHER SPECIMENS EXAMINED.   COLOMBIA: Boyacá: 5♂♂, 28♀♀(3 nymphs), Tinjacá, hotel Ráquira Silvestre, ~  4.7 kmSW de Tinjacá, 05.5485°N, 73.6772°W,  2300 m,  25–26 Oct. 2017, D. Forrero, ex  Furcraeasp. ( Asparagaceae) ( PUJB).   COSTA RICA: San Jose: 1 ♀, San Ignacio de Acosta,  7 June 1988, C. RodriquesG. ( USNM). EL SAL- VADOR:  16♂♂, 8♀♀, B. S. Crandall, ’48, ex henequen [  Agave fourcroya] leaf ( USNM).   MEXICO: Chiapas: 8♂♂ 19♀♀,  3 mi.S La Trinitaria,  20 July 1973, Mastro& Schaffner( TAMU 691-717; 726-729).   Guerrero: 48♂♂, 15♀♀, LaMinilla, Queztalpa,  14 April 2015, P. Figueroa Castro, ex  Agave angustifolia( USNM).   Oaxaca: 42♂♂ 51♀♀,  11.3 mi.SE Totolapan.  21 July 1974, Clark, Murray, Ashe, Schaffner( TAMU 537-579; 609-660; 718-721);  1♂♂,  2.7 mi.NW El Cameron,  21-22 July 1974, Clark, Murray, Ashe, Schaffner( TAMU 580); 27♂♂ 26♀♀, 2♂♂+  ♀in copula,  14 mi.N Miahuatlan,  16 July 1974, Clark, Murray, Ashe, Schaffner( TAMU 581-608; 661-687; 689-690; 722-25);  1♀, Mortega, El Cameron,  22 July 1976,  Opuntiasp.( TAMU 688).   San Luis Potosi: 1♂, Valles,  18 Jan. 1909, brush land ( TAMU 004).  Puebla: 14♂♂ 18♀♀,  10.5 miE Azumbilla,  15 Apr. 1979, T. P. Friedlander& J. C. Schaffner( TAMU 422-453).   Tamaulipas: 1♀,  7 milesS of Antiguo Morelos,  18 Mar. 1975, Clark& Schaffner( TAMU 023).   Yucatan: 1♀, Merida, Yucatan, E. J. Hambleton  13 Aug. 1946, on  Agave( USNM); 6♂♂ 11♀, 1♂♂+  ♀in copula,  19.5 miN Uxmal,  1 Aug. 1980 Schaffner, Weaver, Friedlander, ( TAMU 005-022).   UNITED STATES: Arizona: Pima Co.:  3 ♂♂, 33♀♀, Sonora Desert Museum,  16 May 1990, W. A. Jones, ex  Agave cerulata subcerulataand  Agave murpheyi USNM);  8♂♂, 12♀♀, Pima Co., Sonora Desert Museum,  4 Oct. 1990, W. A. Jones, breeding on  Agave murpheyiGibson( USNM).   Florida: 5♂♂, 10♀♀, Orange Co., Apopka,  9 Aug.1984, P. G. Gibson, ex  Yucca elephantipes[a jr. synonym of  Y. giganteaLem.]( USNM).   Texas: 4♂♂, 5♀♀, Cameron Co., Brownsville,  19 Oct. 1944, ex  Yuccasp. ( USNM).   VENEZUELA: Aragua: 1 ♀, ElLimon,  450 m,  24 May 1982, on Agavaceae( USNM).   DISCUSSION.  Agaveocoris distantiis the most widespread and variable of all the species in the genus. It has multiple color forms across the Neotropics and into the southern United Statesthat share similar body sizes and structures and indistinguishable genitalia, particularly the shape of the right paramere, but are highly variable in color. While most species of the genus seem to be restricted to one or two host species,  A. distantifeeds on many different species, including two species of  Yucca. The original description of  A. distanti( Reuter 1905)described this species as being primarily pale, but we have found populations that contain both pale specimens and darker specimens with the head and pronotum red and reddish orange, and hemelytra often with bluish sheen. The junior synonym  C. rufoscutellatus majorconforms to the paler color forms and  C. rufoscutellatusrepresents the distinctive red to reddish-orange color form. This species would benefit from a molecular DNA analysis to verify species limits in light of the many color variations we now consider belonging to one species. 2611439359 USNM C. H. Ballou 19 220 2 2 holotype 2611439431 USNM Label Merida, Yucatan 19 220 2 2 Merida, Yucatan holotype 2611439337 2017-10-25 2017-10-26 2017-10-25 PUJB D. Forrero Colombia 2300 5.5485 Raquira Silvestre 7 -73.6772 Tinjaca 19 220 33 28 5 Boyaca 2611439450 [286,1322,1376,1403] 1988-06-07 USNM C. Rodriques Costa Rica San Ignacio de Acosta 19 220 1 1 San Jose 2611439312 [264,1128,1412,1439] USNM Crandall Costa Rica 19 220 24 8 16 San Jose 2611439325 1973-07-20 TAMU Mastro & Schaffner Mexico 3 mi. S La Trinitaria 19 220 TAMU 691-717 27 19 8 Chiapas 2611439424 2015-04-14 USNM P. Figueroa Castro Mexico Queztalpa La 19 220 63 15 48 Guerrero 2611439439 1974-07-21 TAMU Clark & Murray & Ashe & Schaffner Mexico 11.3 mi. SE Totolapan. 19 220 TAMU 537-579 93 51 42 Oaxaca 2611439405 1974-07-21 1974-07-22 1974-07-21 TAMU Clark & Murray & Ashe & Schaffner Mexico 2.7 mi. NW El Cameron 19 220 TAMU 580 1 1 Oaxaca 2611439430 1974-07-16 TAMU Clark & Murray & Ashe & Schaffner Mexico 14 mi. N Miahuatlan 19 220 TAMU 581-608 1 1 Oaxaca 2611439408 1976-07-22 TAMU Mexico El Cameron Mortega 19 220 TAMU 688 1 1 Oaxaca 2611439413 1909-01-18 TAMU Mexico Valles 19 220 TAMU 004 1 1 San Luis Potosi 2611439386 1979-04-15 TAMU T. P. Friedlander & J. C. Schaffner Mexico 10.5 mi E Azumbilla 19 220 TAMU 422-453 32 18 14 Puebla 2611439361 [265,1320,1700,1728] 1975-03-18 TAMU Clark & Schaffner Mexico 7 miles S of Antiguo 19 220 TAMU 023 1 1 Tamaulipas 2611439314 1946-08-13 USNM E. J. Hambleton Mexico Merida 19 220 1 1 Yucatan 2611439392 1980-08-01 TAMU Schaffner & Weaver & Friedlander Mexico 19.5 mi N Uxmal 19 220 TAMU 005-022 1 1 Yucatan 2611439396 [957,1406,1772,1799] Pima Co United States of America Arizona 19 220 1 Arizona 2611439317 1990-05-16 USNM W. A. Jones United States of America Sonora Desert Museum 19 220 36 33 3 Arizona 2611439345 1990-10-04 USNM W. A. Jones United States of America Pima Co. Sonora Desert Museum 19 220 20 12 8 Arizona 2611439401 1984-08-09 USNM P. G. Gibson United States of America Orange Co. Apopka 19 220 15 10 5 Florida 2611439351 1944-10-19 USNM United States of America Cameron Co. Brownsville 19 220 9 5 4 Texas 2611439372 [263,1229,1952,1979] 1982-05-24 USNM Venezuela 450 El 19 220 1 1 Aragua