Agaveocoris distanti (Reuter) Henry & Menard, 2020

Henry, Thomas J. & Menard, Katrina L., 2020, 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, Zootaxa 4772 (2), pp. 201-252 : 219-220

Agaveocoris distanti (Reuter) , new combination

( Figs. 24–28 , 83–85 , 124 )

Eurycipitia distanti Reuter 1905: 4 (original description), 1908: 155 (note); Carvalho 1952 (as type of genus in error).

Caulotops rufoscutellatus Carvalho 1948: 530 (original description); Carvalho and Froeschner 1987: 148 (type list). Synonymized by Carvalho 1954: 424.

Caulotops rufoscutellatus major Carvalho 1948: 532 (original description, n. ssp.); Carvalho and Froeschner 1987: 48 (type list). 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. roseus in 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. roseus has 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. Mesoscutum and 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 Yucca sp., Yucca gigantea Lem. [as jr. synonym Yucca elephantipes Regel ex Trel ] [ Agavoideae : Asparagaceae ] ( Henry 1985) and Agave cupreata Trel. & Berger [ Asparagaceae ] and A. vivipara L. ( Figueroa-Castro et al. 2019). From specimen data, this species has been taken on A. fourcroydes Lem. , A. cerulata Trel. [unknown subspp. subcerulata], A. murpheyi Gibson , and A. vivipara L. [as jr. synonym Agave angustifolia Haw. ]. A long series from Colombia was collected on Furcraea sp. [ Asparagaceae ]. One specimen, taken on Opuntia sp. [ Cactaceae ], clearly is an incidental record.

DISTRIBUTION. Described from Venezuela as Eccritotarsus distanti ( Reuter 1905) and as Caulotops rufoscutellatus ( Carvalho 1948) and later reported from Costa Rica, Mexico as 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 Mexico are Chiapas, Guerrero, Oaxaca, Puebla, San Luis Potosi, Tamaulipas, and Yucatan. New U. S. state records are Arizona and Texas.

TYPE MATERIAL EXAMINED. Holotype ♂ ( Caulotops rufoscutellatus Carvalho ): 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 major Carvalho ): Label 1, “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 km SW de Tinjacá, 05.5485°N, 73.6772°W, 2300 m, 25–26 Oct. 2017, D. Forrero, ex Furcraea sp. ( Asparagaceae ) ( PUJB) . COSTA RICA: San Jose: 1 ♀, San Ignacio de Acosta , 7 June 1988, C. Rodriques G. ( 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♀♀, La Minilla, 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, Opuntia sp. ( TAMU 688 ) . San Luis Potosi: 1♂, Valles , 18 Jan. 1909, brush land ( TAMU 004 ) . Puebla: 14♂♂ 18♀♀, 10.5 mi E Azumbilla , 15 Apr. 1979, T. P. Friedlander & J. C. Schaffner ( TAMU 422-453 ) . Tamaulipas: 1♀, 7 miles S 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 mi N 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 subcerulata and Agave murpheyi USNM ); 8♂♂, 12♀♀, Pima Co., Sonora Desert Museum , 4 Oct. 1990, W. A. Jones, breeding on Agave murpheyi Gibson ( USNM) . Florida: 5♂♂, 10♀♀, Orange Co., Apopka , 9 Aug.1984, P. G. Gibson, ex Yucca elephantipes [a jr. synonym of Y. gigantea Lem. ]( USNM) . Texas: 4♂♂, 5♀♀, Cameron Co., Brownsville , 19 Oct. 1944, ex Yucca sp. ( USNM) . VENEZUELA: Aragua: 1 ♀, El Limon, 450 m, 24 May 1982, on Agavaceae ( USNM) .

DISCUSSION. Agaveocoris distanti is 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 States that 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. distanti feeds 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 major conforms to the paler color forms and C. rufoscutellatus represents 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.

Figures

FIGURES 21–31. Agaveocoris spp. 21, A. dimidiatus, n. sp., male (dorsal aspect). 22, A. dimidiatus, n. sp., male (lateral aspect). 23, A. dimidiatus, n. sp., female (dorsal aspect). 24, A. distanti (Reuter), male (brown color form, dorsal aspect). 25, A. distanti (Reuter), male (brown color form, lateral aspect). 26, A. distanti, female (brown color form, dorsal aspect). 27, A. distanti, male (reddish-orange color form, dorsal aspect). 28, A. distanti, female (reddish-orange color form, dorsal aspect). 29, A. marginalis, n. sp., male (dorsal aspect). 30, A. marginalis, n. sp., male (lateral aspect). 31, A. roseus, n. sp., female (dorsal aspect).

FIGURES 80–93. Male genitalia of Agaveocoris spp. A. dimidiata, n. sp.: 80, endosoma. 81, left paramere. 82, right paramere. A. distanti, n. sp.: 83, endosoma. 84, left paramere. 85, right paramere. A. marginalis, n. sp.: 86, endosoma. 87, left paramere. 88, right paramere. A. roseus, n. sp.: 89, right paramere. 90, left paramere. A. rostratus, n. sp.: 91, endosoma. 92, left paramere. 93, right paramere.

FIGURES 119–128. Male genital tubercles of Agaveocoris spp. 119, A. agavis. 120, A. barberi. 121, A. barrerai, n. sp. 122, A. bimaculatus, n. sp. 123, A. dimidiatus, n. sp. 124, A. distanti. 125, A. roseus, n. sp. 126, A. rostratus, n. sp. 127, A. schaffneri, n. sp. 128, A. scutellatus, n. sp.