Melanoliarus, Hendrix & Bartlett, 2025

Hendrix, Solomon V. & Bartlett, Charles R., 2025, Reclassification of the Planthopper Genus Melanoliarus Fennah, 1945 (Hemiptera: Fulgoromorpha: Cixiidae), primarily North of Mexico, with Notes on American Pentastirini, Zootaxa 5619 (1), pp. 1-87 : 22-60

Melanoliarus s.l. ( Melanoliarus sensu Holzinger et al. 2002 )

Diagnosis. Small to midsized pentastirine cixiids, body weakly compressed, wings held tectiform; color usually uniformly deep brown to black, with carinae and legs often paler; body often coated with waxy pruinosity. Head generally narrower than pronotum. Vertex usually narrow (longer than wide), disc concave, weakly convex anteriorly, projecting past eyes; median carina obscure or absent; subapical transverse carina distinct and diagonal (usually meeting medially before head apex); head apex transversely carinate, usually bearing dorsoapical fovea between head apex and subapical transverse carina; hind margin angulately notched. Frons and clypeus collectively rhomboid in shape (truncate dorsally at fastigium). Frons narrowest at fastigium, expanding ventrally and broadest near frontoclypeal margin, lateral margins sinuate, lateral carinae slightly foliate, median carina distinct, forked just below fastigium (defining a frontal fovea that is usually contiguous with fovea formed at the apex of the subapical transverse carina of the vertex). Frons with distinct median ocellus just above frontoclypeal suture. Frontoclypeal suture usually inflected into frons, defining an excavation of variable shape. Lateral ocelli distinct at anterioventral margin of eyes. Antennae short, scape ring-like, pedicel ovoid bearing many sensory pits, flagellum long and bristlelike with bulbous base. Pronotum narrow with hind margin angulate, bearing a median carina, dorsolateral carinae (extending from anterior margin near midline, strongly arched laterally into paradiscal region), and usually lateral carinae (extending transversely between eye and tegula). Mesonotum bearing five carinae (intermediate carinae sometimes obscure). Hind tibiae bearing 2–5 lateral teeth (usually 4–5), apical ornamentation usually 6(5+1)-7-7, platellae absent. Tegulae prominent. Forewings transparent or with few markings (rarely fuscous), but variable within and between species; wings elongate and often weakly spatulate (leading margin weakly convex), apex rounded; wing veins setaceous with tubercles (setal bases) often pigmented darkly; forewing branching pattern most often RA 2-branched, RP 3-branched, MP 5-branched, CuA 2-branched (reduced in some genera, most often with RP 2-branched), not anastomosing (no closed procubital cell, viz. Emeljanov 1996). Male pygofer bilaterally symmetrical (or nearly so) with distinct medioventral lobe (in ventral view) between elongated lateral lobes of the pygofer (medioventral process usually much shorter than lateral lobes). Gonostyli not connected basally. Aedeagal complex articulated dorsally to pygofer with weak, membranous connective to anal tube (vs. Mnemosyne ). Aedeagus with endosoma angled or curved left from ventral view. Anal tube expanded, ovate, and usually hood-like in form.

Distribution. Nearctic and Neotropical (including Antillean) regions (southern Canada to south temperate South America).

I. Generic Reclassification of Melanoliarus

Genus Melanoliarus Fennah, 1945 (s.s.)

Figs. 5–6 , 32 , 34 , 41A–B , 44A

Oliarus (Melanoliarus) Fennah, 1945: 141 .

Melanoliarus Fennah, 1945 , taxonomic status revised by Emeljanov (2001: 122) and Holzinger et al. (2002: 129).

Type species. Oliarus (Melanoliarus) maidis Fennah, 1945 , by original designation.

Diagnosis. Small dark species (~ 3–5 mm). Maculae of the frons absent.Carinae of mesonotum dark, concolorous with disc. Forewing veins pale, tubercles concolorous and obscure. Forewings generally clear, unmarked, with RP 2-branched. Male terminalia, in ventral view, with pygofer narrow, medioventral process weakly pentagonal. Periandrium processes reduced to short projections. Anal tube with apex of caudoventral margin prolonged into acuminate process, angled ventrad in lateral view.

Description. Males 3.2–5.1 mm, body dark brown to black.

Head. Head only slightly narrower than pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex somewhat narrow, longer than wide at midline; disc dark brown to black, carinae light brown to yellow, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae narrowly angled (acutely angled at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black, carinae light brown to yellow (carinae of clypeus usually darker than frons), maculae of frons absent; median carina prominent, frontal fovea well-defined, distinct from dorsoapical fovea; frontoclypeal suture angulate (i.e., frons triangularly excised at ventral margin). Rostrum short, not reaching caudal margin of posterior trochanters.

Thorax. Disc of pronotum brown to black, dorsal carinae light brown to yellow, lateral carinae (between eye and tegula) concolorous to disc. Disc of mesonotum uniformly dark brown to black; carinae concolorous to disc (rarely paler); intermediate carinae weak to nearly obscure. Forewings ( Fig. 6 ) transparent, unmarked; veins pale; tubercles usually inconspicuous and concolorous to veins (except Melanoliarus viequensis (Caldwell)) , absent or very sparsely present on CuP; vein branching pattern RA 2-branched, RP 2-branched, MP 5-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia( Fig. 44A )with narrow pygofer; bilaterally symmetrical; medioventral process weakly pentagonal, sharply acute at apex, half to approximately ¾ length of lateral lobes; lateral lobes in ventral aspect broad, usually rounded apically (sometimes pointed). Gonostyli exceeding caudal margin of pygofer in ventral view, strongly curved cephalad and thickened subapically. Aedeagal complex ( Figs. 41A–B ) with narrow periandrium bearing numerous short processes; sinistral, dextral and median processes reduced to short (sometimes hook-like) protuberances, bearing a subapical process (varying in orientation and length). Dorsum of periandrium usually without any processes, sometimes with recurved process at apex. Endosoma ( Figs. 41A–B ) moderately narrow, directed sharply left (in ventral view) and cephalad (subgenus Melanoliarus ) or caudad (subgenus Amplectiarus subgen. nov.). Endosoma with few processes; apical process of endosoma present (subgenus Melanoliarus ) or absent (subgenus Amplectiarus subgen. nov.); if present, long and sharply angled cephalad at approximately 90º from apex of endosoma, narrow and acute. Anal tube bilaterally symmetrical or slightly asymmetrical, apex angled approximately 90º in lateral view and sharply acute, prolonged into acuminate process (conspicuous in ventral view).

Distribution. Primarily present in the Antillean Islands, as far north as Florida and Texas. There appear to be numerous undescribed Antillean and Central American species.

Etymology. The genus name is derived from the Greek word melas (μέλας, black) as the prefix “ melan -”, added to the genus name “ Oliarus ” (the Latin name for the island of Antiparos in the Cyclades island group in the Aegean sea) ( Dmitriev, 2022). The name is masculine in gender.

Remarks. Members of Melanoliarus are recognized by their small size, dark coloration and the ventral spine on the anal tube, which can be seen in ventrocaudal view without dissection. In terms of the terminalia, Melanoliarus acicus is atypical for the genus as it has a broad and highly elongated process originating from the base of the ventral endosoma directed left (in ventral view) that sharply curves right around the endosoma. The endosoma also lacks the long spine-like apical process common in the subgenus and is twisted caudad. However, the species shares the general form of the pygofer, the reduced sinistral and dextral processes on the ventral periandrium, and the distinctive form of the anal tube. These differences serve as the basis for the new subgenus, Amplectiarus subgen. nov.

“ Melanoliarus sensu stricto ” will be used to refer to the genus as here defined, including both the nominate subgenus and Amplectiarus subgen. nov., excluding the numerous species in Melanoliarus incertae sedis.

Subgenus Melanoliarus Fennah, 1945

Figs. 5 , 32 , 34 , 41A , 44A

Diagnosis. Small dark species (~ 3–5 mm). Maculae of the frons absent. Carinae of mesonotum dark, concolorous with disc. Forewing veins pale and tubercles concolorous and obscure. Forewing generally clear, unmarked, with RP 2-branched. Male terminalia, in ventral view, with pygofer narrow, medioventral process weakly pentagonal. Periandrium processes reduced to short projections. Endosoma in ventral view curved left near to 90º, often with apical process strongly angled cephalad. Anal tube with apex of caudoventral margin prolonged into acuminate process, angled ventrad in lateral view.

Distribution. Mostly Antillean, north to Florida. Present in Belize.

Remarks. In some respects, Melanoliarus (Melanoliarus) is similar to the genus Nivcentia . Both groups are rather small pentastirines commonly found in the Caribbean with an endosoma sharply angled to the left and an apical spine of the endosoma sharply angled cephalad ( Hendrix & Bartlett, 2024a). However, Nivcentia has strongly produced sinistral, medioventral, and dextral processes originating from the periandrium and lacks the distinctive ventral spine of the anal tube found in Melanoliarus . Nivcentia species are usually paler in coloration, somewhat larger, and have more darkly-marked tubercles on the forewings than Melanoliarus .

Caldwell & Martorell (1951) synonymized Oliarus cinereus with M. complectus , a nomenclatural act that was either overlooked or omitted by Mead & Kramer (1982). We found a single male syntype at USNM (here designated as lectotype, see ‘type material examined’ under subgenus Melanoliarus ). Our examination of the Oliarus cinereus lectotype supports the assertion of Caldwell and Martorell that cinereus is a synonym of complectus .

Melanoliarus lunatus ( Fabricius 1803: 54) was originally described from “Americae Infulis” (American Islands), subsequently reported as South America and the Isle of Pines (i.e., the Cuban Isla de la Juventud; e.g., Fabricius 1803, Stål 1869, Heidemann and Osborn 1917). We interpret “Americae Infulis” to refer to the Caribbean Islands due to the species’ morphology agreeing with other members of Melanoliarus (Melanoliarus) , an assertion supported by Caldwell & Martorell (1951: 140) which states that “ complectus or one of the closely related forms may prove to be lunata Fabricius which was described from the West Indies in 1803” and the unpublished notes of F.W. Mead, in which he considered the species near to complectus , maidis , and ‘ cinerea ’ [sic] which he considered to form a species complex with lunatus based on study of the wing structure and female anal tube. Two syntype specimens of this species are in the Fabricius collection at the University of Copenhagen (ZMUC, Fig. 32 ; a female bearing a Mead lectotype label [catalog NHMD918145] and a specimen missing its abdomen [NHMD918131]). Mead never published this lectotype designation. These specimens appear to belong to Melanoliarus (Melanoliarus) and, as suggested by Caldwell & Martorell (1951) could reasonably prove to be a senior synonym of one of the other Caribbean species. According to Zimsen (1964), three more specimens are located at the Zoological Museum of Kiel University, Kiel, Germany (DE-ZMK). We are reluctant to designate a lectotype for this species until the specimens at Kiel can be examined; however, we consider it to belong to the nominate subgenus.

Species Composition

Melanoliarus (Melanoliarus) castro Bahder & Bartlett, 2023 .

Melanoliarus (Melanoliarus) complectus ( Ball, 1902) .

= Oliarus cinereus Wolcott, 1921:18 ; syn. by Caldwell & Martorell, 1951: 138; synonymy here affirmed.

Melanoliarus (Melanoliarus) lunatus ( Fabricius, 1803) .

Melanoliarus (Melanoliarus) maidis Fennah, 1945 .

Melanoliarus (Melanoliarus) viequensis (Caldwell, 1951) .

Subgenus Amplectiarus new subgenus

Figs. 6 , 41B

Type species. Oliarus acicus Caldwell, 1947 .

Diagnosis. Small dark species (~ 4–5 mm). Maculae of the frons absent.Carinae of mesonotum dark, concolorous with disc. Forewing veins pale and tubercles concolorous and obscure. Forewing generally clear, unmarked, with RP 2-branched. Male terminalia, in ventral view, with pygofer narrow, medioventral process weakly pentagonal. Periandrium processes reduced to short projections. Endosoma directed left then caudad, with broad process originating ventrally at aedeagal joint, directed left (in ventral view) and twisted around dorsal surface of endosoma. Anal tube with apex of caudoventral margin prolonged into acuminate process, angled ventrad in lateral view.

Description. Males 4.1–5.1 mm, body black.

Head. Head only slightly narrower than pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex somewhat narrow, longer than wide at midline; disc black, carinae light brown to yellow, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae narrowly angled (acutely angled at juncture), dorsoapical fovea weakly defined. Eyes without markings. Disc of face (frons+clypeus) black, carinae light brown to yellow (carinae of clypeus usually darker than frons), maculae of frons absent; median carina prominent, frontal fovea well-defined, distinct from dorsoapical fovea; frontoclypeal suture angulate (i.e., frons triangularly excised at ventral margin). Rostrum short, not reaching caudal margin of posterior trochanters.

Thorax. Disc of pronotum generally black, dorsal carinae light brown to yellow, lateral carinae (between eye and tegula) concolorous to disc. Disc of mesonotum uniformly black; carinae concolorous to disc; intermediate carinae weak. Forewings transparent, unmarked; veins pale; tubercles usually inconspicuous and concolorous, to veins absent or very sparsely present on CuP; vein branching pattern RA 2-branched, RP 2-branched, MP 5-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia with narrow pygofer; bilaterally symmetrical; medioventral process weakly pentagonal, sharply acute at apex, half-length to approximately ¾ length of lateral lobes; lateral lobes in ventral aspect broad, usually rounded apically (sometimes pointed). Gonostyli exceeding caudal margin of pygofer in ventral view, strongly curved cephalad and thickened subapically. Aedeagal complex ( Fig. 41B ) with narrow periandrium bearing numerous short processes; in ventral view with sinistral, dextral, and median processes reduced to short protuberances; with short subapical process. Dorsum of periandrium usually without any processes, sometimes with recurved process at apex. Endosoma somewhat narrow, directed sharply left (in ventral view) then caudad. Endosoma ( Fig. 41B ) with single basal process originating at aedeagal joint, directed left (from ventral view) before curving right around dorsum of endosoma. Anal tube bilaterally symmetrical or slightly asymmetrical, apex angled approximately 90º in lateral view and sharply acute, prolonged into acuminate process (conspicuous in ventral view).

Distribution. Southern Texas to Mexico.

Etymology. The genus name is derived from the Latin word amplexus (embracing/encircling) + Oliarus , a reference to the form of the endosoma. The name is masculine in gender.

Remarks. This subgenus is similar to the nominate subgenus externally and in portions of the male terminalia (pygofer, gonostyli, and anal tube). However, the form of the endosoma is unlike that of Melanoliarus (Melanoliarus) , warranting the segregation of a new subgenus. The two groups also appear to be geographically isolated as presently understood.

Species Composition

Melanoliarus (Amplectiarus) acicus (Caldwell, 1947) , new combination.

Genus Amiarus new genus

Figs. 7–10 , 37 , 41C–D , 44B–C

Type species. Oliarus kindli Bourgoin, Wilson & Couturier, 1998 .

Diagnosis. Medium-sized (~ 5–6 mm); light brown to black. Head in lateral view with acute apex. Face narrow, maculae of frons reduced or absent; frontoclypeal suture angulate (defining triangular area). Mesonotal carinae orange to concolorous with mesonotum. Wing veins pale, tubercles concolorous; CuP without tubercles; wings generally unmarked or marked with transverse dark pigmentation at anterior and basal thirds of tegmina. Male pygofer with medioventral process small and elongate; acuminate to triangular. Periandrium with three conspicuous elongate ventral processes. Endosoma narrow and apically lobed (bearing 1–2 subapical processes). Anal tube somewhat bilaterally asymmetrical in dorsal aspect; with prominent ridge surrounding epiproct and paraproct in lateral view.

Description. Males 4.5–6.2 mm, body light brown to black.

Head. Head narrower than pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex broad, longer than wide or long as wide at midline; disc yellow to black, carinae light brown to yellow, lateral maculae absent; lateral margins subparallel (or weakly converging anteriorly); median carina weakly present to obscure (or absent); subapical transverse carinae somewhat broadly angled (convexly rounded at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) yellow to black, carinae light brown to yellow maculae of frons absent; median carina prominent, frontal fovea weakly to well-defined, distinct from or merged with dorsoapical fovea; frontoclypeal suture angulate (i.e., frons triangularly excised at ventral margin). Rostrum somewhat short, not attaining caudal margin of posterior trochanters.

Thorax. Disc of pronotum light brown to black, dorsal carinae light brown to yellow, lateral carinae (between eye and tegula) yellow or concolorous to disc. Disc of mesonotum uniformly light brown to black, carinae usually concolorous to disc (except A. deserticola comb. nov. in which carinae paler); intermediate carinae often weak. Forewings ( Figs. 8 , 10 ) transparent, generally unmarked, often tinted yellow (except A. dimidiatus comb. nov. and A. (Umbrarus) humilis comb. nov. in which forewings have transverse dark pigmentation at anterior and basal thirds); veins pale; tubercles usually inconspicuous, concolorous to veins, absent (or sparsely present) on CuP; vein branching pattern RA 2-branched, RP (1–2)-branched, MP 5-branched, CuA 2-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia ( Figs. 44B–C ) with broad pygofer, medioventral process (in ventral view) small and elongate, acuminate or triangular; medioventral process less than half length of lateral lobes; lateral lobes approximately symmetrical, pointed at apex in ventral view. Gonostyli exceeding caudal margin of pygofer in ventral view, thick at base, narrowed just before curving and expanding subapically. Aedeagal complex ( Figs. 41C–D ) with broad periandrium, bearing several processes; venter of periandrium with three prominent elongated processes (sinistral, medioventral and dextral); sinistral process long and slender, usually curving left; medioventral process long and slender, apically arcuate (often surpassing endosoma), arched leftwards (sometimes curved cephalad over endosoma); dextral process long and slender, often leaning right; periandrium in dorsal view with (subgenus Umbrarus subgen. nov.) or without (subgenus Amiarus ) long slender mediobasal process extending length of endosoma; apex of periandrium in dorsal view with spine-like median process, distally hooked right, either short (e.g., A. humilis comb. nov., A. oryzicola comb. nov.), elaborated into prominent twisted process (e.g., A. deserticola comb. nov.), or intermediate (e.g., A. kindli comb. nov.). Endosoma somewhat narrow, directed left (ventral view); slightly inflated distally after aedeagal joint, then narrowed before prominent membranous apical lobe (directed cephalad); bearing long, slender dorsal process (originating near aedeagal joint, often not conspicuous until subapical region) and slender, elongate, apical process. Anal tube somewhat bilaterally asymmetrical in dorsal aspect, ovate, longer than wide, hood-like; with prominent ridge surrounding epiproct and paraproct (prominently visible in ventral view as two protuberances; Figs. 44B, C ).

Distribution. Canada to Argentina .

Etymology. The genus name derived from a truncation of “America” joined to a truncation of Oliarus , a reference to the group’s widespread presence throughout North and South America. The name is masculine in gender.

Remarks. Amiarus gen. nov. is widespread throughout North and South America. While described species in the nominate subgenus are restricted to South America, we have studied specimens from Mexico, Guatemala, and Costa Rica also belonging to the subgenus.

This genus is difficult to identify from habitus and is often dependent on examination of the male terminalia, although this can be done without dissection. In ventral view, the two protuberances on the anal tube (which are parts of a semi-circular ridge) separate this group from most others. Perhaps most distinctive about the general habitus is the moderate body length of the genus coupled with the pale wing veins with concolorous tubercles, in addition to the lack of tubercles on the CuP vein (somewhat abnormal for American Pentastirini of this size). Additionally, the rostrum is shorter than in other groups of comparable size (failing to attain the caudal margin of the posterior trochanters, except in a few specimens of A. (U.) humilis comb. nov.).

Based on photographs of the female type specimen of the Brazilian Cixius fulvus Walker, 1858 , this species appears to belong to this genus and we place it here as Amiarus fulvus comb. nov. The species A. deserticola comb. nov. is unusual for the group (in comparison to the other described species), in having a much narrower vertex and unusual coloration, but the characters of the terminalia unequivocally agree with those of the genus.

Subgenus Amiarus

Figs. 7–8 , 37 , 41C , 44B

Diagnosis. Medium-sized (~ 5–6 mm); light brown to black. Head in lateral view with acute apex, bearing apical transverse carinae. Face narrow, maculae of frons reduced or absent; frontoclypeal suture angulate (defining triangular area). Mesonotal carinae orange to concolorous with mesonotum. Wing veins pale, tubercles concolorous; wings generally unmarked or marked with transverse dark pigmentation at anterior and basal thirds of tegmina. Male pygofer with medioventral process small and elongate; acuminate to triangular. Periandrium with three conspicuous ventral processes, lacking prominent mediobasal process on dorsum, but with hook-like process at dorsal apex. Endosoma narrow, directed left and apically lobed (bearing 1–2 subapical processes).Anal tube somewhat bilaterally asymmetrical in dorsal aspect; with prominent ridge surrounding epiproct and paraproct in lateral view.

Distribution. Southern Mexico to Argentina ; Trinidad.

Species Composition

Amiarus (Amiarus) deserticola ( Campodonico, 2018) , new combination.

Amiarus (Amiarus) dimidiatus ( Berg, 1879) , new combination.

Amiarus (Amiarus) fulvus ( Walker, 1858) , new combination.

Amiarus (Amiarus) kindli ( Bourgoin, Wilson & Couturier, 1998) , new combination.

Amiarus (Amiarus) oryzicola ( Bourgoin, Wilson & Couturier, 1998) , new combination.

Subgenus Umbrarus new subgenus

Figs. 2A , 9–10 , 41D , 44C

Type species. Flata humilis Say, 1830 .

Diagnosis. Small planthoppers (~ 4–5 mm); dark brown to black. Head narrower than pronotum. Maculae of vertex small if present. Face black with carinae dark brown to orange; maculae of frons absent; frontoclypeal suture angular (defining triangular region). Forewings smoky, heavily marked, often deeply embrowned, entirely black or marked with transverse dark pigmentation at distal and proximal thirds; veins pale or dark, RP 2-branched. Male pygofer with medioventral process triangular, about half (or more) length of lateral lobes; lateral lobes obtuse at apex in ventral view. Ventral periandrium with three long digitiform processes. Dorsum of periandrium with long, slender median process. Endosoma broad, directed left (in ventral view), with apical process; apically lobed.

Description. Males 4.1–5.0 mm, body dark brown to black.

Head. Head narrower than pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex broad, longer than wide or long as wide at midline; disc dark brown to black; carinae dark brown to yellow, lateral maculae absent; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae somewhat broadly angled (convexly rounded or truncate at juncture), dorsoapical fovea weakly defined (nearly obscure). Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black, carinae light brown to concolorous to disc, maculae of frons absent; median carina prominent, frontal fovea well-defined, merged with dorsoapical fovea (apical transverse carina of vertex obsolete at merger); frontoclypeal suture angulate (i.e., frons triangularly excised at ventral margin). Rostrum length variable, usually not reaching caudal margin of posterior trochanters, though sometimes slightly exceeding margin.

Thorax. Disc of pronotum dark brown to black, dorsal carinae yellow or concolorous to disc, lateral carinae (between eye and tegula) concolorous to disc. Disc of mesonotum uniformly dark brown to black, carinae concolorous to disc; intermediate carinae weak. Forewings ( Fig. 10 ) fumose and heavily marked, either entirely embrowned or marked with transverse dark pigmentation at distal and proximal thirds; veins pale or dark; tubercles dark, absent or sparse on CuP; vein branching pattern RA 2-branched, RP (1–2)-branched, MP 5-branched, CuP 2-branched. Legs dark brown, without banding.

Abdomen. Male terminalia ( Fig. 44C ) with broad pygofer; medioventral process (in ventral view) triangular, half (or more) length of lateral lobes; lateral lobes bilaterally symmetrical, obtuse at apex in ventral view. Gonostyli exceeding caudal margin of pygofer in ventral view, thick at base, narrowed just before curving and expanding subapically. Aedeagal complex ( Fig. 41D ) with broad periandrium giving rise to digitiform processes; in ventral view, periandrium venter with three prominent processes (sinistral, medioventral, and dextral); sinistral process long and slender, apex strongly curved left; medioventral process long and slender, arched left; dextral process slender, straight, leaning right; in dorsal view; dorsum of periandrium with long, slender process originating mediobasally (mediodorsal process), nearly surpassing length of endosoma; apex of dorsal periandrium with spine-like median process, hooked right. Endosoma broad, directed left (in ventral view), apically lobed; endosoma bearing a dorsolateral process and a narrow subapical process on dorsum. Anal tube bilaterally symmetrical in dorsal view, broadly ovate (longer than wide), hood-like.

Distribution. Transcontinental U.S. and Canada, as far south as Arizona.

Etymology. The subgenus name is derived from the modern Latin word umbra (shadow) + Oliarus , a reference to the darkly-marked forewings. The name is masculine in gender.

Remarks. The strongly and distinctly patterned wings, on a nearly black body, make Umbrarus subgen. nov. one of the most distinctive Nearctic pentastirines. Specimens, particularly in vivo, can be waxy-pruinose, causing the body to take on a bluish cast.

Species Composition

Amiarus (Umbrarus) humilis ( Say, 1830) , new combination.

Genus Antilliarus new genus

Figs. 1B , 11– 12 , 41E , 44D

Type species. Oliarus slossoni Van Duzee, 1912 .

Diagnosis. Medium-sized planthoppers (typically ~ 6–7 mm); reddish-brown. Vertex relatively narrow; transverse carinae of vertex greatly angled, lateral maculae present. Eyes mottled with red spots. Frons broad with prominent, but diffuse, yellow maculae. Rostrum very long, exceeding hind trochanters. Mesonotum with carinae concolorous to orange. Forewing veins pale with tubercles concolorous; unmarked or with variable transverse dark bar proximally. Male terminalia with short trilobed medioventral pygofer process; narrow periandrium lacking sinistral process, but with slender dextral and subapical processes; narrow endosoma with enlarged and bulbous apex with three apical processes.

Description. Males 5.0– 7.4 mm, body reddish-brown to dark brown.

Head. Head only slightly narrower than pronotum; in lateral view rounded at fastigium, projecting anteriorly. Vertex generally narrow (excepting Antilliarus aztecus comb. nov.), longer than wide at midline; disc reddish-brown to black, carinae yellow to brown (median carina often concolorous to disc), lateral maculae usually weakly present; often bearing incomplete median carina from posterior margin; subapical transverse carinae of vertex steeply angled, carinae orange, lateral maculae present (sometimes small). Eyes mottled with many red spots. Disc of face (frons+clypeus) reddish-brown to dark brown (often lustrous), carinae yellow to brown, maculae of frons present (large, yellow, and diffuse); median carina weak to obscure (excepting A. aztecus comb. nov.), frontal fovea well-defined to obsolete, somewhat merged with dorsoapical fovea when present (apical transverse carina of vertex completely to partially obsolete at merger); frontoclypeal suture quadrangulate (i.e., frons quadrangularly excised at ventral margin). Rostrum very long, exceeding caudal margin of posterior trochanters.

Thorax. Disc of pronotum color reddish-brown to black, dorsal carinae yellow to brown, lateral carinae (between eye and tegula) yellow or concolorous to disc. Disc of mesonotum reddish-brown to black with median portion (within lateral carinae) often paler than lateral portions, carinae yellow to concolorous to disc; intermediate carinae weak. Forewings ( Fig. 12 ) transparent, usually unmarked or marked with variable transverse bar proximally (sometimes also darkened along trailing margin); veins pale; tubercles obscure, concolorous to wing veins (excepting A. aztecus comb. nov.), present on CuP; vein branching pattern MA 2-branched, MP 3-branched, RP 5-branched, CuP 2-branched. Legs generally uniformly yellow to pale brown, without banding.

Abdomen. Male terminalia ( Fig. 44D ) with broad pygofer; medioventral process (in ventral view) very small, usually somewhat trilobed or sagittate (lateral lobes of medioventral process much shorter than median lobe if trilobed); medioventral process much less than half length of lateral lobes; lateral lobes bilaterally symmetrical. Gonostyli exceeding caudal margin of pygofer in ventral view, narrow at base, continuing to narrow distally before spatulately expanding subapically. Aedeagal complex ( Fig. 41E ) with narrow periandrium bearing several elongate processes; in ventral view with sinistral process absent but with prominent, slender subapical process present on left side, directed left; median process slender, directed right; dextral process long and slender; additional small process directed left at apex of periandrium; periandrium dorsum with prominent retrose process rising subapically. Endosoma narrow, directed left before making a 90º sinistral turn; apical region of endosoma enlarged and bulbous, bearing apical processes (usually three with at least one elongated). Anal tube slightly bilaterally asymmetrical in dorsal aspect, ovate, longer than wide, hood-like; in lateral view appearing elongate triangular.

Distribution. Primarily Neotropical, ranging from throughout the Antillean islands (as far south as Curaçao) to southern Florida; also present in Mexico.

Etymology. The genus name is an arbitrary combination derived from a truncation of Antilles joined with a truncation of Oliarus . The name is a reference to the group’s widespread distribution in the Antilles . The name is masculine in gender.

Remarks. Antilliarus gen. nov. is distinguished by the usually narrow vertex, large size, and red speckling of the eyes (which it shares with Oliaridus gen. nov. and Oliaronus ). It is also relatively large relative to other Antillean Pentastirini (i.e., larger than most Nivcentia and Melanoliarus s.s., although smaller than most Cyclopoliarus ). The genus appears to be common and widespread in the Antilles and the southern half of Florida. Based on the red speckling of the eyes and almost loop-like form of the endosoma, this genus is likely related to Oliaridus gen. nov.

Antilliarus slossoni comb. nov. in the Virgin Islands is commonly found on century plant ( Agave americana L.) (e.g., Bartlett 2000; reported as Oliarus slossonae ), but is apparently not associated with this plant elsewhere (e.g., Puerto Rico and Florida). Virgin Island specimens also appear to have a narrower vertex and lack the dark transverse band at the base of the forewings, raising the possibility that these specimens attributed to A. slossoni comb. nov. may represent multiple species or subspecies. However, Mead & Kramer (1982) described ‘puzzling’ variation in the male terminalia of specimens from Florida, so the interpretation of species variation should be approached with caution.

This group bears superficial resemblance to the genus Cyclopoliarus , which is quite similar in its rather large size, reddish-brown coloration, and prominent dark maculae on the frons. However, Cyclopoliarus males can be distinctively larger, have a very prominent acute projection at the end of the anal tube (e.g., Fennah 1945a, fig. 24), and have a distinctively different aedeagal complex with a long and arcuate apical process of the endosoma. Through examination of the illustrations of the described Cyclopoliarus terminalia ( Myers 1928; Metcalf 1938; Fennah 1945 a, 1971), Cyclopoliarus jamaicensis and C. pirata belong to Antilliarus gen. nov. and are here transferred to the genus as A. jamaicensis comb. nov. and A. pirata comb. nov. Both species have a close affinity to A. slossoni comb. nov. and are similar except for the form of the endosoma. While A. jamaicensis comb. nov. is notably smaller than the other two taxa with a significantly shorter endosoma and shorter apical processes, examination of the type material will be necessary to confirm that both of these species are not simply variants of M. slossoni . The remaining six Cyclopoliarus species appear to be a heterogeneous grouping and require further review.

While most of the specimens observed belonging to this group are from Antillean islands, the disjunct Mexican species A. aztecus comb. nov. belongs to this genus based on the form of the male terminalia. It differs in some respects from the external morphology of the Antillean members of the genus including its broad vertex, distinctive tubercles on the wing veins, and a prominent median carina of the frons, possibly warranting subgeneric status in the future.

Also belonging to the group is an undescribed species from Curaçao. The species also has a wide vertex and the endosoma is greatly reduced in length. The medioventral process of the periandrium is also quite different in both A. aztecus comb. nov. and the Curaçao species, at least from A. slossoni comb. nov., in that they are pentagonal rather than trilobed (de Haas, pers. comm.).

Species Composition

Antilliarus aztecus (Caldwell, 1947) , new combination.

Antilliaru s jamaicensis (Fennah, 1945) , new combination.

Antilliarus pirata ( Fennah, 1971) , new combination.

Antilliarus slossoni ( Van Duzee, 1912) , new combination.

Genus Bifoliarus new genus

Figs. 13–14 , 41F , 44E

Type species. Oliarus chuliotus Ball, 1934

Diagnosis. Small (usually ~ 4 mm) dark species. Lateral maculae of vertex present. Maculae of frons absent. Forewings transparent, without pigmentation, with reduced venation (RP 2-branched, MP 4-branched). Pygofer in ventral view bilaterally symmetrical (or nearly so); medioventral process somewhat sagittate, apex pointed, constricted at base. Periandrium of aedeagal complex bearing several ventral processes (obscuring sinistral and dextral processes); in dorsal view periandrium with sclerotized plate bearing broad, pointed and diverging processes.

Description. Males 3.4–4.9 mm, body dark brown to black.

Head. Head narrower than pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex narrow, longer than wide at midline; disc dark brown to black, carinae yellow to black, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae somewhat acutely angled (convexly rounded at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black (clypeus sometimes somewhat paler than frons), carinae yellow to brown; maculae of frons absent; median carina prominent, frontal fovea weakly defined, distinct from dorsoapical fovea; frontoclypeal suture arched (i.e., frons semicircularly excised at ventral margin). Rostrum short, not reaching caudal margin of posterior trochanters.

Thorax. Disc of pronotum dark brown to black, dorsal carinae yellow to orange, lateral carinae (between eye and tegula) dark yellow to concolorous to disc. Disc of mesonotum uniformly dark brown to black, carinae concolorous to disc, occasionally orange; intermediate carinae somewhat weak. Forewings ( Fig. 14 ) transparent, usually without pigmentation (but sometimes slightly fumose); broadly rounded at apex; veins pale with tubercles generally concolorous, sparsely present on CuP; vein branching pattern RA 2-branched, RP 2-branched, MP 4- branched. Legs brown proximally, smoky-yellow distally, without banding.

Abdomen. Male terminalia ( Fig. 44E ) with broad pygofer; medioventral process (in ventral view) ovate, constricted at base, apex rounded or broadly pointed; lateral lobes symmetrical or somewhat asymmetrical, apices obtuse in ventral view. Gonostyli only slightly exceeding caudal margin of pygofer in ventral view, narrow at base, continuing to narrow distally before curving outwards (perpendicular to body) and slightly thickening subapically. Aedeagal complex ( Fig. 41F ) with narrow periandrium bearing several elongated processes; venter of periandrium with four prominent processes; sinistral process short and hooked, curved left (in ventral view), swollen at basal two-thirds; median of ventral periandrium giving rise to process curving sharply left behind sinistral process; dextral process long, tapered, sometimes bifurcate; dorsum of periandrium with prominent sclerotized plate bearing two broad and pointed processes; left process pointing left (in dorsal view) or curved cephalad; right process at extreme right apex of sclerotized plate, directed right or caudad. Endosoma broad, short, and curved leftwards (in ventral view), bearing variable processes; slender process originated from base of ventral endosoma; apical process sometimes present (viz. Bifoliarus sablensis comb. nov.). Anal tube bilaterally symmetrical (or nearly so), ovate, longer than wide, hood-like, apex usually slightly concave.

Distribution. Generally eastern U.S. and adjacent Canada; as far west as western Texas.

Etymology. The genus name is derived from Bi - (latin, “two”) + folia (latin, leaf) + Oliarus ; referring to the sclerotized leaf-like plate bearing two processes on the dorsum of the periandrium; a feature unique to this genus.

Remarks. This genus is the chuliotus -group of Mead & Kramer (1982). Bifoliarus gen. nov. species are among the smallest Pentastirini in the Nearctic. Bifoliarus gen. nov. can be separated from other genera by their small size, dark body coloration, lack of maculae on the frons, and transparent, unmarked forewings with reduced venation (viz. RP 2-branched, MP 4-branched). However, the best diagnostic feature to separate Bifoliarus gen. nov. from other small pentastirines is the sclerotized plate on the dorsum of the periandrium which bears short expanded processes. This is the best feature to separate Bifoliarus gen. nov. from Vitroliarus gen. nov., along with the form of the anal tube and differing shape of the medioventral process of the pygofer (roughly sagittate in Bifoliarus gen. nov., triangular in Vitroliarus gen. nov.). Vitroliarus gen. nov. also tend to have faintly embrowned forewings, compared to clear forewings of most specimens of Bifoliarus gen. nov. Members of this genus are most often taken by sweeping or beating vegetation.

The delineation between members of Bifoliarus gen. nov. in the eastern United States is a problem deserving greater attention. Mead and Kramer (1982) briefly addressed this issue in their descriptions of Bifoliarus chuliotus comb. nov. and B. sablensis comb. nov. Unpublished notes of F.W. Mead accessed at the Florida State Collection of Arthropods (FSCA) detailed his initial uncertainty in defining a reliable diagnostic character to distinguish these species (including detailed correspondence asking for thoughts from colleagues); he arrived at the concept that B. bispinus comb. nov. has no endosomal processes, B. chuliotus comb. nov. has one endosomal process and B. sablensis comb. nov. bears two. Beyond this, the species exhibit a troubling degree of variation in the form of the processes of the periandrium. A high degree of variation is present in the species throughout their distributions, suggesting that they may be members of a larger complex of closely related species. Morphological analysis of the male terminalia will likely be insufficient to resolve this problem without molecular study.

Species Composition

Bifoliarus bispinus (Caldwell, 1947) , new combination.

Bifoliarus chuliotus ( Ball, 1934) , new combination.

Bifoliarus sablensis (Caldwell, 1951) , new combination.

Genus Cilidius new genus

Figs. 1C , 15–16 , 33 , 42A , 44F

Type species. Cixius vicarius Walker, 1851 .

Diagnosis. Relatively large (~ 5–8 mm) with a proportionately narrower head relative to the prothorax; vertex wider at posterior margin than a midline. Face relatively narrow; maculae of the frons present, but obscure. Carinae of mesonotum prominent. Forewings transparent (sometimes variable marked with fuscous, especially females), with veins pale and prominent dark tubercles. Male pygofer with large bluntly sagittate medioventral process. Ventral periandrium with sclerotized area with two processes bearing an array of spine-like processes, endosoma weakly angled left from ventral view.

Description. Males 5.4–8.2 mm, body reddish-brown to dark brown.

Head. Head narrower than pronotum; in lateral view weakly angulate or rounded at fastigium, somewhat projecting anteriorly. Vertex broad, longer than wide or as long as wide at midline; disc reddish-brown to dark brown, carinae yellow to nearly concolorous to disc, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina weakly present to obscure (or absent); subapical transverse carinae broadly angled (convexly rounded or truncate at juncture), dorsoapical fovea well-defined. Eyes sometimes with dark transverse median marking. Disc of face (frons+clypeus) reddish-brown to dark brown, carinae orange, maculae of frons present (reduced to thin lines); median carina prominent, frontal fovea well-defined, distinct from dorsoapical fovea; frontoclypeal suture quadrangulate (i.e., frons quadrangularly excised at ventral margin). Rostrum reaching or surpassing caudal margin of posterior trochanters.

Thorax. Disc of pronotum reddish-brown to dark brown; dorsal carinae yellow, lateral carinae (between eye and tegula) yellow to concolorous to disc. Disc of mesonotum reddish-brown to dark brown with median portion (within lateral carinae) often paler than lateral portions, carinae orange to concolorous to disc; intermediate carinae distinct. Forewings ( Fig. 16 ) usually transparent, sometimes with dark patterning (especially females); veins pale except where spotting or banding is present; tubercles dark and conspicuous, present on CuP; vein branching pattern RA 2-branched, RP 3-branched, MP 5-branched, CuA 2-branched. Legs with front and middle tibiae with alternating light and dark banding.

Abdomen. Male terminalia ( Fig. 44F ) with broad pygofer, approximately bilaterally symmetrical; medioventral process large (half or more length of lateral lobes) and bluntly sagittate, narrowest at base then abruptly expanded before narrowing towards a blunt apex; lateral lobes symmetrical and broad in ventral aspect. Gonostyli exceeding caudal margin of pygofer in ventral view, narrow at base, thickening distally and curving outwards (perpendicular to body) and thickening subapically. Aedeagal complex ( Fig. 42A ) with narrow periandrium bearing few processes; venter of periandrium with short and slender curved process on left (from ventral view, apparently the sinistral process), directed left then curved right; venter of periandrium (near apex) bearing two large, elongated, sclerotized processes; upper process generally straight pointing caudad, curved at apex; lower process shorter, pointing cephalad; both bearing many small spine-like processes; dorsum of periandrium with long and slender left-basad process barely exceeding length of endosoma, curved right (in dorsal view), barely surpassing length of endosoma. Endosoma slender, short, and directed cephalad, slightly bent left (in ventral view), weakly expanded distally, bearing short and thickened dorsal process. Anal tube weakly bilaterally asymmetrical, long and narrow, flap-like, apex truncate, slightly concave at midlength.

Distribution. Common in the eastern U.S. and adjacent Canada; as far west as Colorado.

Etymology. The genus name is derived from the modern Latin word cilia - (eyelash, plural) + -idius (Greek:— ίδιον, diminutive suffix); referring to the many small lash-like processes of the ventral aedeagal processes. The name is masculine in gender.

Remarks. This genus is the quinquelineatus -group of Mead & Kramer (1982). Cilidius gen. nov. is a common taxon in the eastern U.S. Wing pigmentation is highly variable in this group, ranging from completely transparent to heavily banded. The dark tubercles of the wings are highly contrasting with the veins leading to a “peppered” appearance that is often a usefully diagnostic feature. The dense arrangement of small processes on the ventral process of the aedeagal complex separates Cilidius gen. nov. from all other American genera of Pentastirini . The shape of the medioventral process of the pygofer along with the larger body length also usefully separates Cilidius gen. nov. from other eastern US Pentastirini . Cilidius gen. nov. differs from Melanoliarus s.s. in being larger and broader with a proportionately narrower head relative to the prothorax. Cilidius gen. nov. overlaps in size with Xenoliarus gen. nov. but is readily separated from this genus by the conspicuous tubercles on the wing veins and the medioventral process of the pygofer in males (greatly expanded in Xenoliarus gen. nov.).

Examination of photographs of the female holotype of Cixius testaceus ( Fig. 33 ), a species of unknown provenance, agrees with the habitus of boldly-marked females of Cilidius vicarius comb. nov. Here we designate Cixius testaceus as a new junior synonym of Cilidius vicarius . Further details are provided below (in section ‘IIa. Synonyms’).

Species Composition

Cilidius quinquelineatus ( Say, 1830) , new combination.

Cilidius vicarius ( Walker, 1851) , new combination.

= Cixius testaceus Walker, 1851 , new synonym.

Genus Galapagoliarus new genus

Fig. 17

Type species. Oliarus agrippa Fennah, 1967 .

Diagnosis. Small planthoppers (~ 4–6 mm); reddish-brown.Vertex narrow, about 1.25x as long as wide; subapical transverse carinae of vertex steeply diagonal, meeting at sharp angle (resulting in elongated triangular lateral fovea), lateral maculae present. Carinae of face pale and conspicuous, maculae absent, clypeus darker than frons. Forewings transparent, usually unmarked (or with few maculations), veins pale bearing dark tubercles; RA 1-branched, RP 2-branched. Male terminalia, in caudoventral view, with narrow inversely triangular pygofer, narrowest ventrally, widely broadening dorsad. Ventral periandrium with at least three processes (sinistral, medioventral and dextral), dorsal periandrium with several small processes. Endosoma with long, slender dorsal process and slender, elongated apical process (usually directed cephalad). Anal tube somewhat bilaterally asymmetrical in dorsal aspect, ovate, longer than wide; either obtusely rounded at apex or emarginated.

Description. Male and females 3.5–5.8 mm, body reddish-brown to dark brown.

Head. Head narrower than pronotum; in lateral view weakly angulate or rounded at fastigium, somewhat projecting anteriorly. Vertex narrow, longer than wide; disc reddish-brown to dark brown, carinae pale yellow to orange, lateral maculae present; lateral margins parallel or subparallel (or weakly converging anteriorly); median carina weakly present to obscure (or absent); subapical transverse carinae very steeply angled (acutely angled at juncture), dorsoapical fovea weakly defined or obscure. Eyes without significant markings. Disc of face (frons+clypeus) reddish-brown to dark brown (clypeus somewhat darker than frons), carinae yellow to orange, maculae of frons absent; median carina somewhat reduced (though distinctly pigmented), frontal fovea somewhat weakly defined, distinct from dorsoapical fovea; frontoclypeal suture arched (i.e., frons semicircularly excised at ventral margin). Rostrum long, surpassing caudal margin of posterior trochanters.

Thorax. Disc of pronotum reddish-brown to dark brown, dorsal carinae pale yellow to dark yellow, lateral carinae (between eye and tegula) dark yellow to concolorous to disc. Disc of mesonotum uniformly reddish-brown to dark brown, carinae orange to concolorous to disc; intermediate carinae distinct. Forewings transparent (occasionally with few markings, especially in females), veins pale bearing distinct dark tubercles, present on CuP; vein branching pattern RA 1-branched, RP 2-branched MP 5-branched, CuA 2-branched. Legs yellow to light brown, paler distally, without banding.

Abdomen. Male terminalia with narrow pygofer in ventral view, narrowest at ventral margin, widely broadening dorsad; medioventral process triangular to somewhat pentagonal, relatively short (less than half length lateral lobes), distally ornamented with ridges; lateral lobes bilaterally symmetrical, obtuse pointed apically in ventral view. Gonostyli exceeding caudal margin of pygofer in ventral view, narrow at base, curving and thickening subapically. Aedeagal complex with somewhat broad periandrium bearing numerous elongated processes; venter of periandrium with sinistral process long and slender, curved left (in ventral view); medioventral process long and slender, pointed left; dextral process long and slender, curved and pointed caudad; dorsum with variable processes including subapical process directed cephalad. Endosoma broad to narrow in shape with broad lobe directed left (in ventral view) and long, slender dorsal process; apical process slender and elongate, directed retrorsely. Anal tube somewhat bilaterally asymmetrical in dorsal aspect, ovate, longer than wide; either obtuse or emarginated at apex.

Distribution. Galápagos Islands.

Etymology. The genus name is derived from a truncation of ‘Galápagos’ combined with Oliarus . The name is masculine in gender.

Remarks. This genus is endemic to the Galápagos islands. The most distinctive feature seems to be the the male pygofer, which is narrow basally and broadens distally resulting in a flabelliform appearance unique to this group. The small size, color, and narrow vertex are also notable features. The dark tubercles of the wing veins give Galapagoliarus gen. nov. a distinctly “peppered” appearance. Males in this genus sometimes have markings on the forewings, which is somewhat atypical for the tribe.

The form of the aedeagal complex in Galapagoliarus alastor comb. nov. is atypical. The endosoma is expanded and has an additional dorsal subapical process, pointing medially. The apical process of the endosoma is greatly angled inwards, compared to the usual cephalad angle in the rest of the genus.

Fennah (1967) treated all of the Galápagos Pentastirini , except for the subsequently described Oliarus hernandezi . Unfortunately, Fennah’s (1967) illustrations of the aedeagal complex are only from dorsal view, which obscures some important features. The obligately cavernicolous Galápagos species Galapagoliarus hernandezi comb. nov. is unusual in several respects, likely due to its troglobitic nature. Hoch & Izquierdo (1996) suggested that the ancestor of this species was likely an epigean species from the islands (which they suggested was most likely an ancestor of alastor or agrippa ). Similar to G. alastor comb. nov., the endosoma is expanded with inward-pointing processes. We tentatively place this species in Galapagoliarus gen. nov., despite the substantive differences in its external morphology.

Species Composition

Galapagoliarus agrippa ( Fennah, 1967) , new combination.

Galapagoliarus alabandus ( Fennah, 1967) , new combination.

Galapagoliarus alastor ( Fennah, 1967) , new combination.

Galapagoliarus galapagensis ( Van Duzee, 1933) , new combination.

Galapagoliarus hernandezi ( Hoch & Izquierdo, 1996) , new combination.

Galapagoliarus remansor ( Fennah, 1967) , new combination.

Genus Lapsium new genus

Figs. 18–19 , 42B , 45A

Type species. Oliarus difficilis Van Duzee, 1912 .

Diagnosis. Medium-sized (~4.5–5.0 mm), reddish-brown (mesonotum sometimes paler medially). Lateral maculae of vertex distinct, maculae of frons highly reduced and diffuse. Forewings clear, tubercles distinct on veins, vein forks and some crossveins embrowned; RP 2-branched. Male terminalia with medioventral process of pygofer triangular, less than half of length of lateral lobes; lateral lobes narrow with obtuse apices; in ventral view sinistral and dextral processes of the periandrium highly reduced to small triangular protrusions; two large, slender processes on ventral periandrium, one of which is directly behind sinistral process; endosoma curved to left with apex recurved right with two long apical processes directed right-cephalad.

Description. Males 4.5–5.0 mm, body reddish-brown.

Head. Head narrower than pronotum; in lateral view weakly angulate or rounded at fastigium, projecting anteriorly. Vertex narrow, longer than wide at midline; disc reddish-brown, carinae yellow to orange, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae steeply angled (acutely angled at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) reddish-brown (margins with pale wash, interpretable as maculae of frons), carinae yellow to orange, maculae of frons absent or obscure; median carina prominent, frontal fovea well-defined, distinct from dorsoapical fovea; frontoclypeal suture quadrangulate (i.e., frons quadrangularly excised at ventral margin). Rostrum somewhat long, usually just surpassing caudal margin of posterior trochanters.

Thorax. Disc of pronotum reddish-brown, dorsal carinae yellow to dark yellow, lateral carinae (between eye and tegula) generally concolorous to disc. Disc of mesonotum reddish-brown to dark brown with median portion (within lateral carinae) often paler than lateral portions, carinae paler than to nearly concolorous to disc; intermediate carinae highly reduced, nearly obsolete. Forewings ( Fig. 19 ) generally clear, veins brown; tubercles darkly pigmented, present on CuP; veins often embrown at forks and crossveins; vein branching pattern RA 2-branched, RP 2-branched, MP 5-branched. Legs mostly light brown, without banding.

Abdomen. Male terminalia ( Fig. 45A ) with somewhat broad pygofer, bilaterally symmetrical; medioventral process (in ventral view) triangular, less than half length of lateral lobes; lateral lobes long and narrow with obtuse apices. Gonostyli not attaining or barely exceeding caudal margin of pygofer in ventral view, thickened at base, slightly narrowing distally before curving outwards and slightly expanding subapically. Aedeagal complex ( Fig. 42B ) with narrow periandrium bearing five processes; venter with sinistral process very short and pointed, directed caudad; left subbasal area bearing long, slender process directed left (directly behind sinistral process); median area bearing long, slender process directed caudad then left at apex before helical curve (sinistrally); dextral process very short and pointed, directed right; dorsum of periandrium (dorsal view) with short pointed process curved left at right apical area of periandrium. Endosoma moderately broad, curved to left (ventral view) with the apex recurved right with two long, straight, and slender dorsoapical processes directed right-cephalad. Anal tube slightly asymmetrical, ovate, longer than wide, apex rounded.

Distribution. Gulf states of the U.S.; with specimen records from Belize and Dominica.

Etymology. The genus name is from the modern Latin word lapsus (“slip” or “error”) + -ium (noun suffix). Named for the atypical location and shape of the processes on the periandrium and endosoma. The name is masculine in gender.

Remarks. Lapsium gen. nov. is externally most similar to Oliaridus gen. nov. and Antilliarus gen. nov. in its general habitus, but differs greatly from those genera in the form of the male terminalia and its lack of red spotting of the eyes. Lapsium gen. nov. is reddish-brown in coloration, similar to Oliaridus gen. nov. and has the center of the mesonotum paler than the lateral margins, plus spot-like markings on the wings at vein branches. However (in males), the narrower vertex, the unusual length of the lateral processes of the periandrium (comparable only to Oliaridus kieferi comb. nov.), the shape and arrangement of the processes of the periandrium, and the shape of the endosoma are distinct. Antilliarus gen. nov. is also reddish-brown in coloration but is larger, usually with a narrower vertex, and distinctive terminalia. The three genera can also be differentiated by the shape of the medioventral process of the male pygofer. Of the taxa addressed in this work, Lapsium gen. nov. has the most elongated lateral lobes of the male pygofer in ventral view ( Figs. 18E , 45A ).

Mead & Kramer (1982: 441) state that the type species is “an uncommon species of Neotropical affinities as yet found only in the southern half of Florida and the Lower Rio Grande Valley in the United States” and “the species has no close relationship to the other members of the Nearctic fauna.” We have seen at least one specimen belonging to this genus from Belize and another from Dominica, supporting Mead & Kramer’s assertion of its Neotropical affinities .

Species Composition

Lapsium difficilis ( Van Duzee, 1912) , new combination.

Genus Oliaridellus new genus

Figs. 20–21 , 42C , 45A

Type species. Oliarus apache Ball, 1934 .

Diagnosis. Medium-sized planthoppers (~ 4–5 mm), dark brown to black. Head subequal or slightly narrower pronotum. Maculae of vertex present, maculae of the frons reduced or absent. Forewings clear, veins bearing conspicuous dark tubercles, RP generally 2-branched apically, some vein nodes and crossveins embrowned. Male pygofer with medioventral process (in ventral view) elongate, triangular or acuminate. Periandrium with sinistral process reduced to hump-like lobe, bearing a process on the left side of aedeagus just below the endosoma. Endosoma narrow and arcuate, forming nearly complete loop; with a short process on outer margin and two to four apical processes. Gonostyli slender, and asymmetrical; only slightly surpassing length of pygofer in ventral view (more so in O. retentus comb. nov.).

Description. Males 3.6–5.1 mm, body dark brown to black.

Head. Head slightly narrower or same width as pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex narrow, longer than wide or long as wide at midline; disc dark brown to black, carinae yellow to brown, lateral maculae present or absent; lateral margins subparallel (or weakly converging anteriorly) or nearly parallel; median carina obscure (or absent); subapical transverse carinae angled (convexly rounded at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black (margins sometimes with pale wash, interpretable as maculae of frons), carinae yellow to orange, maculae of frons reduced or absent; median carina prominent, frontal fovea weakly defined, distinct from dorsoapical fovea; frontoclypeal suture quadrangulate (i.e., frons quadrangularly excised at ventral margin). Rostrum short, not reaching caudal margin of posterior trochanters.

Thorax. Disc of pronotum dark brown to black, dorsal carinae yellow to brown, lateral carinae (between eye and tegula) yellow to concolorous to disc. Disc of mesonotum uniformly dark brown to black, carinae weakly orange or concolorous to disc (often median carina orange with remaining concolorous to mesonotum); intermediate carinae reduced. Forewings ( Fig. 21 ) clear, unmarked with veins pale except some vein forks and crossveins embrowned; tubercles darkly pigmented and conspicuous, present on CuP; vein branching pattern RA 2-branched, RP 2-branched, MP 5-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia ( Fig. 45A ) with broad pygofer; medioventral process (in ventral view) relatively large, elongate triangular or acuminate, broadest at base, half or more length of lateral lobes; lateral lobes bilaterally symmetrical or asymmetrical. Gonostyli only slightly exceeding caudal margin of pygofer in ventral view, very slender at base, narrowing distally before curving outwards (perpendicular to body) and expanding subapically. Aedeagal complex ( Fig. 42C ) with narrow periandrium bearing few processes; venter with sinistral process greatly reduced to hump-like lobe (ventral view); elongated, slender process at left lateral apex; dextral process long and slender, directed caudad; dorsum with retrorse process either at extreme left apex ( Oliaridellus apache comb. nov.) or extreme right apex ( O. retentus comb. nov.), sometimes bifurcate. Endosoma narrow and arcuate, with few processes, forming helical loop with apex pointing caudad; endosoma bearing short process on outer margin (“dorsal process” of Mead & Kramer 1982); process at inner apex either short and broad ( O. apache comb. nov.) or long and slender ( O. retentus comb. nov.); processes at median apex absent ( O. apache comb. nov.) or present ( O. retentus comb. nov.). Anal tube approximately bilaterally symmetrical (dorsal view), ovate, longer than wide, broadly rounded with apex slightly concave.

Distribution. Southwestern U.S. and adjacent Mexico.

Etymology. The genus name is derived from the genus name ‘ Oliaridus’ with the Latin diminutive suffix ‘- ellus ’. The name is masculine in gender.

Remarks. Oliaridellus gen. nov. is very similar to Oliaridus gen. nov., especially in terms of the male terminalia (both of which compose the aridus -group of Mead & Kramer 1982 in addition to Melanoliarus canyonensis and M. papagonus ). The most significant differences between the two groups are the morphology of the head, the structure of the periandrium, and body length. In the former, the vertex is much narrower, but the head itself is quite broad relative to the pronotum; in the latter, the vertex is wide, but the head is notably narrower than the width of the pronotum. In Oliaridellus gen. nov., the rostrum also fails to reach the caudal margin of the posterior trochanters, while in Oliaridus gen. nov., the rostrum attains or surpasses that margin. In terms of the male terminalia, the medioventral process of the male pygofer is larger relative to the lateral lobes in Oliaridellus gen. nov. In Oliaridellus gen. nov., the medioventral process of the pygofer is half or greater than half the length of the lateral lobes, while in Oliaridus gen. nov., the medioventral process of the pygofer is generally much less than half the length of the lateral lobes (excluding Oliaridus hesperius comb. nov.). The gonostyli of Oliaridellus gen. nov. are much smaller and quite slender compared to the long, broad gonostyli of Oliaridus gen. nov. The number of processes on the ventral periandrium is greatly reduced in Oliaridellus gen. nov. in comparison to Oliaridus gen. nov. and their position and shape differ. Oliaridellus gen. nov. has the sinistral process reduced to a hump. Mead & Kramer (1982) recognized this hump for Oliaridellus apache comb. nov., but overlooked or excluded its description in Oliaridellus retentus comb. nov., instead diagnosing the long left apical process of the periandrium as the sinistral process in that species. We interpret the long processes at the left of the periandrium (in ventral view) as homologous between the two species. The form of the endosoma, while looped in both taxa (the defining character of the “ aridus -group” of Mead & Kramer 1982, excluding Cyclopoliarus and Oliaronus ), is also very different in width. The endosoma of Oliaridellus gen. nov. is notably narrow and arcuate, with very few processes at the apex, while the endosoma of Oliaridus gen. nov. is quite broad, with several long processes originating from the apex.

In some ways, Oliaridellus retentus comb. nov. is somewhat intermediate between Oliaridellus apache comb. nov. and Oliaridus gen. nov. Unlike Oliaridellus apache comb. nov., which has a small retrose process on the extreme left apex of the dorsal periandrium (in dorsal view), Oliaridellus retentus comb. nov. has a small retrorse process on the extreme right apex of the dorsal periandrium, much like several species in Oliaridus gen. nov. (e.g., Oliaridus kieferi comb. nov.). Additionally, Oliaridellus retentus comb. nov. has the processes of the endosoma somewhat more elaborated, with the outer process and inner apical processes longer and more slender as well as two additional processes at the apex of the endosoma (similar to Oliaridus gen. nov., but still quite reduced in comparison). However, Oliaridellus retentus comb. nov. is still much closer to Oliaridellus apache comb. nov. in general habitus and form of the male terminalia, especially the gonostyli and ventral periandrium.

Species Composition

Oliaridellus apache ( Ball, 1934) , new combination.

Oliaridellus retentus (Caldwell, 1947) , new combination.

Genus Oliaridus new genus

Figs. 22–23 , 42D , 45C

Type species. Oliarus aridus Ball, 1902 .

Diagnosis. Medium-to-large-sized planthoppers (~ 4–8 mm); brownish-red to brown in coloration. Vertex often wider at base than length at midline, lateral maculae present. Maculae of frons large and conspicuous; eyes mottled with red spots. Mesonotal carinae concolorous to orange; wings with dark tubercles on veins; wing veins intermixed pale and dark; wings marked with spotting at vein forks. Pygofer with small, bluntly sagittate medioventral process (in ventral view), length reaching less than half length of lateral lobes of pygofer. Periandrium narrow, usually with narrow sinistral process, dextral process (when present) longer than sinistral. Endosoma broad and arcuate, helically looped to surpass aedeagus, possessing long process on outer margin, shorter process on inner margin, and numerous long apical processes.

Description. Males 4.0– 8.4 mm, body reddish-brown to dark brown.

Head. Head narrower than pronotum; in lateral view rounded at fastigium, somewhat projecting anteriorly. Vertex narrow to broad, longer than wide or as long as wide at midline; disc reddish-brown to dark brown, carinae light brown to yellow, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina weakly present to obscure (or absent); subapical transverse carinae broadly angled (truncate or convexly rounded at juncture), dorsoapical fovea weakly to well-defined. Eyes mottled with many red spots. Disc of face (frons+clypeus) pale reddish-brown to dark brown (often lustrous), carinae yellow to orange, maculae of frons large and prominent; median carina prominent, frontal fovea well-defined to obsolete, either merged in some species with dorsoapical fovea (apical transverse carina of vertex completely to partially obsolete at merger) or distinct; frontoclypeal suture quadrangulate (i.e., frons quadrangularly excised at ventral margin). Rostrum moderately long, reaching or surpassing caudal margin of posterior trochanters (excepting the variable Oliaridus californicus comb. nov. where the rostrum may not attain the caudal margin of the posterior trochanters in some individuals).

Thorax. Disc of pronotum reddish-brown to dark brown, dorsal carinae light brown to yellow, lateral carinae (between eye and tegula) yellow or concolorous to disc. Disc of mesonotum reddish-brown to dark brown with median portion (within lateral carinae) often paler than lateral portions, carinae orange to concolorous to disc; intermediate carinae often weaker or obscure. Forewings ( Fig. 23 ) transparent or variably patterned, veins often conspicuous, intermixed pale and dark, often embrowned at vein forks and crossveins; tubercles often pigmented darkly, present on CuP vein; vein branching pattern RA 2-branched, RP 3-branched, MP 5-branched, CuA 2 branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia ( Fig. 45C ) with broad pygofer; medioventral process (in ventral view) bluntly sagittate, narrowest at base then sharply expanded before narrowing to apex, reaching less than half the length of lateral lobes; lateral lobes bilaterally symmetrical or asymmetrical, broad in ventral aspect, usually narrowing to point at apex (excepting Oliarodus hesperius comb. nov.). Gonostyli exceeding caudal margin of pygofer in ventral view (excluding Oliaridus kieferi comb. nov. with gonostyli shorter than lateral lobes of pygofer), slender at base before spatulately expanding medially or subapically. Aedeagal complex ( Fig. 42D ) with narrow periandrium bearing numerous processes; venter usually with narrow sinistral process of varied length and position; median process often present at apex of periandrium (variable in length and direction, sometimes two processes present); dextral process variable, but always present, sometimes bifurcate, longer than sinistral process; dorsum but not always, with singular slender process at apex. Endosoma broad and arcuate, narrowest basally, curved left and directed medially, looping behind (and often surpassing) periandrium (excepting Oliaridus truncatus comb. nov.); endosoma with long process on outer margin (“dorsal process” of Mead & Kramer 1982); shorter process present at inner margin; numerous long processes at apex. Anal tube bilaterally symmetrical or asymmetrical, elongate, broadly rounded with apex sometimes concave in dorsal view.

Distribution. Widely distributed in the U.S. and southern Canada, reaching as far south as Belize.

Etymology. The genus name is an amalgam of the genus name ‘ Oliarus’ truncated and combined with ‘ aridus’, the specific epithet of the type species. The name is masculine in gender.

Remarks. This is a genus consisting mostly of larger species where the endosoma of the aedeagus is helically curved to (in ventral view) surpass the aedeagus. Mead & Kramer (1982) tentatively placed the species Oliaridellus apache comb. nov., Melanoliarus canyonensis , Melanoliarus papagonus , and Oliaridellus retentus comb. nov. in their ‘ aridus -group’ (here Oliaridus gen. nov., in part); we have elected to define Oliaridus gen. nov. more narrowly than the ‘ aridus -group’ of Mead & Kramer since each of these listed species is unique in certain crucial characters. We have placed apache and retentus in Oliaridellus gen. nov., which is compared to Oliaridus gen. nov. in the remarks of that genus.

This genus can often be partly diagnosed by the red spots on the eyes, shared by few other Pentastirini . Antilliarus gen. nov. also has red mottling on the eyes but has distinctive male terminalia and inconspicuous tubercles on the wing veins (which are dark in Oliaridus gen. nov.). Oliaronus also shares the red mottling of the eyes and has similar male terminalia, but is distinctive in that the costal and adjacent radial cells of the forewing are highly setaceous and sclerotized and bear approximately 20 apical cells (formed by the RA and RP each 4-branched and the MP 1 3-branched, compared with RA 2-branched, RP 3-branched and MP 1 2-branched in Oliaridus gen. nov.).

The Mexican endemic Melanoliarus abacus bears similarities to this genus (notably in having a sinistrally looping endosoma) but appears reduced in several features of the male terminalia and bears a very prominent process on the ventral face of the endosoma. It also differs in external features (e.g., lacks the pale maculae of the frons). This species is allied to Oliaridus gen. nov. but is not sufficiently similar to include in the genus. For this reason, we have elected to leave M. abacus in Melanoliarus incertae sedis ( Table 2) until additional Mexican and Central American material can be reviewed.

Species Composition

Oliaridus aridus ( Ball, 1902) , new combination.

Oliaridus caldwelli ( Mead & Kramer, 1982) , new combination.

Oliaridus californicus ( Van Duzee, 1914) , new combination.

Oliaridus hesperius ( Van Duzee, 1917) , new combination.

Oliaridus kieferi ( Mead & Kramer, 1982) , new combination.

Oliaridus knullorum ( Mead & Kramer, 1982) , new combination.

Oliaridus lobatus ( Caldwell, 1938) , new combination.

Oliaridus pima ( Kirkaldy, 1907) , new combination.

Oliaridus sementinus ( Ball, 1902) , new combination.

Oliaridus sonoitus ( Ball, 1937) , new combination.

Oliaridus truncatus ( Van Duzee, 1929) , new combination.

Genus Remotiarus new genus

Figs. 24– 25 , 42E , 45D

Type species. Oliarus yavapanus Ball, 1934 .

Diagnosis. Small planthoppers (mostly 4–5 mm); dark brown to black; mesonotal carinae orange; wings unmarked or fuscous, veins dark bearing tubercles, RP 2-branched. Male terminalia with pygofer bearing pentagonal medioventral process (in ventral view), much shorter than lateral lobes of pygofer. Periandrium broad, bearing welldeveloped sinistral and dextral processes (ventral view), dorsum with single slender process (dorsal view). Endosoma curved left with indentation at joint with aedeagus (ventral view); apex bearing single spine-like process.

Description. Males 3.9–4.8 mm, body dark brown to black.

Head. Head narrower than pronotum; in lateral view rounded or weakly angulate at fastigium, somewhat projecting anteriorly. Vertex narrow, longer than wide at midline; disc dark brown to black, carinae light brown to orange, lateral maculae present (though often weakly so); lateral margins subparallel (or weakly converging anteriorly); median carina weakly present to obscure (or absent); subapical transverse carinae narrowly angled (acutely angled or truncate at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black, carinae yellow to orange, maculae of frons absent; median carina prominent, frontal fovea weakly defined, distinct from dorsoapical fovea; frontoclypeal suture arched or quadrangular (i.e., frons semicircularly or quadrangularly excised at ventral margin). Rostrum short, not surpassing caudal margin of posterior trochanters.

Thorax. Disc of pronotum dark brown to black, dorsal carinae light brown to orange, lateral carinae (between eye and tegula) yellow to concolorous to disc. Disc of mesonotum uniformly brown to black, carinae strongly pigmented orange; intermediate carinae prominent. Forewings ( Fig. 25 ) immaculate, sometimes fuscous (especially apically) with veins pale; tubercles dark or concolorous to veins, absent (or sparsely present) on CuP; vein branching pattern RA 2 branched, RP 2-branched, MP 5-branched, CuA 2-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia ( Fig. 45D ) with broad pygofer, bilaterally symmetrical, narrowest at base (in ventral view); medioventral process small and pentagonal, sharply or moderately narrowed to apex; lateral lobes moderately broad in ventral aspect, diverging, pointed or rounded towards apex. Gonostyli slender, broadened and highly curved distad at apex, slightly exceeding caudal margin of pygofer in ventral view. Aedeagal complex ( Fig. 42E ) with broad periandrium bearing several processes; in ventral view periandrium with distinct sinistral, medioventral, and dextral processes; sinistral process broad or slender basally (sometimes bifurcate, split into two opposing processes); medioventral process broad at base and curved left in ventral view; dextral process shorter than other processes and curved right (except in R. bazarus comb. nov., in which the process is longer and curved left); in dorsal view, dorsum of periandrium bearing single slender process medially. Endosoma broad, from ventral view angled left, somewhat curved with slight depression at caudal margin near aedeagus, curved left apex; bearing narrow process at apex and sometimes with subapical processes, varying in length. Anal tube nearly bilaterally symmetrical, longer than wide, uniformly rounded.

Distribution. Southwestern U.S. to central Mexico.

Etymology. The genus name is derived from the modern Latin word “ remotus” (isolated) truncated and combined with a truncation of the genus name Oliarus . The name is a reference to the isolated nature of the described species in this genus. The name is masculine in gender.

Remarks. This is the yavapanus -group of Mead & Kramer (1982). Remotiarus gen. nov. is very similar to Sonorium gen. nov. in its small size, dark brown body color, immaculate to fuscous forewings, and possession of prominent orange carinae on the mesonotum. Sonorium gen. nov. generally differs in its obscure or absent pigmentation of the mesonotal carinae, except the Floridian endemic Sonorium littoralis comb. nov. (although it can be excluded by distribution). It is easiest to differentiate Remotiarus gen. nov. from similar taxa through examination of the male terminalia: the shape and placement of the sinistral and dextral processes of the periandrium, the spine-like dorsal process, and the form of the endosoma (which is broad and indented caudally at the aedeagal joint) are enough to distinguish the genus from externally similar groups.

The central Mexican species Remotiarus bazarus comb. nov. is placed in this genus based on similar external morphology and male terminalia to the U.S. species, although it is rather unusual for this genus. The species is most similar in the form of the endosoma. It also shares the placement of the elaborated sinistral, medioventral, and dextral processes of the ventral periandrium. The shape of these processes differs from the three U.S. species in that they are much more slender. The discovery of related Mexican taxa may warrant a reevaluation of the concept of this genus.

This genus is somewhat similar to the inconspicuous western U.S. species Melanoliarus catus and Melanoliarus coconinus (both treated here in Melanoliarus incertae sedis) in their external morphology. Both species are dark in body color with orange mesonotal carinae, are of similar size, and are primarily southwestern in distribution. However, both have broader vertices than any member of Remotiarus gen. nov. and they differ significantly in the male terminalia.

Species Composition

Remotiarus bazarus (Caldwell, 1951) , new combination.

Remotiarus forcipatus (Caldwell, 1947) , new combination.

Remotiarus uncatus (Caldwell, 1947) , new combination.

Remotiarus yavapanus ( Ball, 1934) , new combination.

Genus Sonorium new genus

Fig. 26–27 , 42F , 45E

Type species. Oliarus corvinus Ball, 1934 . Diagnosis. Small planthoppers (mostly 4–5 mm); dark brown to black. Mesonotal carinae concolorous to bright orange (sometimes intermediate carinae concolorous with median and lateral carinae orange). Forewing clear (sometimes embrowned), veins pale, tubercles evident. Pygofer (ventral view) with triangular to weakly pentagonal medioventral process. Periandrium (ventral view) with prominent sinistral process ‘caliper-like’ in shape (bearing a ventral process that appears opposed to angled apex); dextral process variable in length; dorsum bearing single slender elongated process (dorsal view). Endosoma strongly arcuate, curved left in ventral view, bearing numerous processes apically.

Description. Males 3.4–5.5 mm, body brown to black.

Head. Head narrower than pronotum; in lateral view weakly angulate at fastigium, somewhat projecting anteriorly. Vertex narrow, longer than wide at midline; disc dark brown to black, carinae light brown to orange, lateral maculae weakly present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae narrowly angled (truncate at juncture), dorsoapical fovea well-defined. Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black, carinae yellow to orange, maculae of frons absent; median carina prominent, frontal fovea weakly defined, distinct from dorsoapical fovea; frontoclypeal suture angulate ( S. concinnulus comb. nov., S. corvinus comb. nov.), quadrangulate (e.g., Sonorium dondonius comb. nov., S. littoralis comb. nov.), or arched ( S. altanatus comb. nov., S. sylvaticus comb. nov.). Rostrum short, not surpassing caudal margin of posterior trochanters.

Thorax. Disc of pronotum dark brown to black, dorsal carinae light brown to yellow, lateral carinae (between eye and tegula) light brown to concolorous to disc. Disc of mesonotum dark brown to black, carinae orange to concolorous to disc, intermediate carinae prominent. Forewings clear or embrowned (especially apically), veins pale; tubercles pigmented darkly or concolorous to veins, present on CuP; vein branching pattern RA (1–2)-branched, RP 2-branched, MP 5-branched, CuA 2-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia ( Fig. 45E ) with broad pygofer (often weakly asymmetrical); medioventral process (in ventral view) small and triangular or weakly pentagonal, broadest at base and narrowing towards apex, narrowest at base, reaching up to half length of lateral lobes; lateral lobes moderately broad in ventral aspect, with apices either pointed or rounded (in ventral view). Gonostyli exceeding caudal margin of pygofer in ventral view, slender at base, narrowing distally before thickening and curving cephalad subapically. Aedeagal complex ( Fig. 42F ) with somewhat narrow periandrium bearing numerous processes; venter of periandrium with highly developed sinistral process, usually caliper-like; dextral process always present, variable from short and spine-like to long and apically curved right (in ventral view), surpassing length of sinistral process. Dorsum of periandrium with singular slender process originating from median basal periandrium. Endosoma broad and arcuate, curved helically left (endosomal apex sometimes reaches, but does not exceed, aedeagal shaft), bearing several elongated processes near apex. Anal tube roughly symmetrical, longer than wide, broadly rounded with apex often concave in dorsal view (except S. altanatus comb. nov. with apex rounded).

Distribution. Western U.S. and Canada to Mexico; Florida.

Etymology. “ Sonora ” + - ium (noun suffix). Referring to the primary geographic distribution of many species of this group. The name is masculine in gender.

Remarks. This is the concinnulus -group of Mead & Kramer (1982). Sonorium gen. nov. is similar to several smaller pentastirine groups with unmarked wings but is differentiated through examination of the aedeagal complex. The ‘caliper-like’ sinistral process is distinctive among the American Pentastirini . Melanoliarus catus is externally similar, but differs in features of the terminalia and notably lacks the caliper-like sinistral process. The sinistral process of M. catus is nearly obsolete (a large falcate process originating in the left-median basiventral area may superficially appear to be the sinistral process; Mead & Kramer 1982, fig. 49). Some members of this genus are highly variable in the form of the terminalia, as noted by Mead & Kramer (1982). We regard it highly likely that some of these forms represent additional species present in the western United States, as initially suggested by Mead & Kramer (1982). As with Bifoliarus gen. nov., molecular analysis is likely necessary to delineate these species complexes.

Species Composition

Sonorium altanatus (Caldwell, 1951) , new combination.

Sonorium concinnulus ( Fowler, 1904) , new combination.

Sonorium corvinus ( Ball, 1934) , new combination.

Sonorium dondonius ( Ball, 1934) , new combination.

Sonorium littoralis ( Ball, 1934) , new combination.

Sonorium sylvaticus (Caldwell, 1947) , new combination.

Sonorium zyxus (Caldwell, 1947) , new combination.

Genus Vitroliarus new genus

Figs. 28–29 , 43A , 45F

Type species. Oliarus ecologus Caldwell, 1947 .

Diagnosis. Small planthoppers (about 4 mm), dark brown to black. Mesonotal carinae usually concolorous to mesonotum (sometimes median and lateral carinae orange). Forewings weakly fumose (especially apically), unmarked; veins pale, tubercles concolorous to veins; venation somewhat reduced (RP 2-branched, MP (4–5) branched). Medioventral process of male pygofer small and triangular. Periandrium with six total processes; ventral periandrium with two hyaline left apical processes and two acute and short median subapical processes directed caudad; dorsal periandrium with two long processes.

Description. Males 3.7–4.5 mm, body dark brown to black.

Head. Head narrower than pronotum; in lateral view angulate at fastigium, somewhat projecting anteriorly. Vertex narrow, longer than wide at midline; disc dark brown to black, carinae brown to orange, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae narrowly angled (acutely angled at juncture), dorsoapical fovea weakly defined. Eyes without significant markings. Disc of face (frons+clypeus) dark brown to black, carinae yellow to orange, maculae of frons absent; median carina distinct, frontal fovea somewhat weakly defined, distinct from dorsoapical fovea; frontoclypeal suture angulate (i.e., frons triangularly excised at ventral margin). Rostrum short, not reaching caudal margin of posterior trochanters.

Thorax. Disc of pronotum dark brown to black, dorsal carinae yellow to dark brown, lateral carinae (between eye and tegula) yellow. Disc of mesonotum dark brown to black, carinae concolorous to disc (median and lateral carinae sometimes orange); intermediate carinae somewhat weaker. Forewings ( Fig. 29 ) fumose (often darkening near apex), unmarked, veins pale; tubercles concolorous to wing veins, absent (or sparsely present) on CuP; vein branching pattern RA 2-branched, RP 2-branched, MP (4–5)-branched, CuA 2-branched. Legs yellow to dark brown, paler distally, without banding.

Abdomen. Male terminalia ( Fig. 45F ) with broad pygofer; medioventral process (in ventral view) triangular; usually less than half length of lateral lobes; lateral lobes bilaterally symmetrical, bluntly pointed at apex in ventral view. Gonostyli exceeding caudal margin of pygofer in ventral view, slender at base, narrowing distally before thickening and curving cephalad subapically. Aedeagal complex ( Fig. 43A ) with broad periandrium bearing few processes; venter without sinistral or dextral processes; bearing two hyaline left apical processes curving left, then ventrocephalad apically, the top process small and slender and the bottom process long and broad; two acute and short median subapical processes directed caudad; periandrium in dorsal view with two long medial processes, surpassing endosoma; left process originating basad (left of midline), sinuate, leaning left; right process originating subapically on right, strongly arched to direct dorsad, apex j-hooked. Endosoma broad, strongly angled left in ventral view (with apex curved to direct retrorsely), bearing two subapical processes, one dorsal and one ventral; ventral process long and slender, directed cephalad, dorsal process shorter with broader base (tapering distally) directed left-cephalad (in dorsal view). Anal tube weakly bilaterally asymmetrical in dorsal aspect, ovate, longer than wide, caudal margin slightly concave.

Distribution. Eastern and central U.S.

Etymology. The genus name is derived from the modern Latin word vitrum (glass) + Oliarus , a reference to the vitreous apical processes of the periandrium. The name is masculine in gender.

Remarks. Vitroliarus gen. nov. is easily confused with Bifoliarus gen. nov. due to their similar coloration and tendency to occur in similar habitats. Externally, the two can be separated by the size and shape of the vertex ( Vitroliarus gen. nov. somewhat broader than Bifoliarus gen. nov.), the color of the mesonotal carinae ( Vitroliarus gen. nov. tends to have the carinae orange—especially the median carina—while in Bifoliarus gen. nov. the carinae are usually concolorous to the mesonotum), and the color of the wings (fumose, darkest apically, in Vitroliarus gen. nov., transparent or uniformly slightly fumose in Bifoliarus gen. nov.). This last feature can cause Vitroliarus gen. nov. to be confused with Amiarus (Umbrarus) gen. et. subgen. nov., but the latter has much darker wings and pigmentation on the thorax. Additionally, Vitroliarus gen. nov. is separated from similar eastern U.S. taxa (e.g., Bifoliarus gen. nov., Melanoliarus , Amiarus (Umbrarus) gen. et. subgen. nov.) through examination of the small, triangular medioventral process of the male pygofer, which can be seen without dissection.

As noted by Mead & Kramer (1982), the sole representative of this genus is notable in having a hyaline apical region of the ventral periandrium, which bears two processes. These processes can be very difficult to discern in the cleared terminalia.

Species Composition

Vitroliarus ecologus (Caldwell, 1947) , new combination.

Genus Xenoliarus new genus

Figs. 1A , 30–31 , 43B , 46A

Type species. Oliarus placitus Van Duzee, 1912 .

Diagnosis. Relatively large (5–9 mm), brown to black, maculae of frons conspicuous. Forewings usually clear (markings sometimes present), veins intermixed pale and dark with dark tubercles. Male terminalia with a greatly expanded, spatulate medioventral pygofer process. Gonostyli (ventral view) asymmetrical and arcuate with lobed apices. Periandrium broad bearing several large, elongate processes; ventral periandrium with singular sinistral process. Endosoma highly reduced.

Description. Males 6.1–8.8 mm (eastern U.S. species) or 4.7–6.4 mm (western U.S. species), body dark brown to black.

Head. Head narrower than pronotum; in lateral view rounded at fastigium, projecting anteriorly. Vertex somewhat broad, longer than wide at midline; disc dark brown to black, carinae brown to yellow, lateral maculae present; lateral margins subparallel (or weakly converging anteriorly); median carina obscure (or absent); subapical transverse carinae somewhat broadly angled (convexly rounded at juncture), dorsoapical fovea weakly to well-defined. Eyes often with dark transverse median marking. Disc of face (frons+clypeus) dark brown to black, carinae yellow to concolorous to disc, maculae of frons present (usually reduced to thin lines); median carina distinct, frontal fovea well-defined, distinct from dorsoapical fovea; frontoclypeal suture quadrangulate (i.e., frons rectangularly excised at ventral margin). Rostrum long, surpassing caudal margin of posterior trochanters.

Thorax. Disc of pronotum dark reddish-brown to dark brown, dorsal carinae brown to yellow, lateral carinae (between eye and tegula) yellow. Disc of mesonotum dark reddish-brown to dark brown with median portion (within lateral carinae) often paler than lateral portions, carinae concolorous to disc; intermediate carinae prominent. Forewings ( Fig. 31 ) clear, usually with few dark markings (especially near margin, but sometimes forewings more extensively marked or entirely embrowned, especially females); veins intermittent pale and dark; tubercles darkly pigmented, present on CuP; vein branching pattern RA 2-branched, RP 3-branched, MP 5-branched, CuA 2-branched. Legs with alternating light and dark banding (many in eastern U.S. taxa, few in western U.S. taxa).

Abdomen. Male terminalia ( Fig. 46A ) with broad pygofer; medioventral process greatly expanded and broadly spatulate, slightly asymmetrical, narrowest at base, apex slightly concave; lateral lobes narrow in ventral aspect, diverging, broadly triangular in lateral aspect. Gonostyli exceeding caudal margin of pygofer in ventral view, very slender at base, curving in arch outwards then inwards, in ventral view appearing curved following margin of spatulate medioventral process of pygofer, expanded subapically. Aedeagal complex ( Fig. 43B ) with broad periandrium bearing several large processes; ventral periandrium with large elongate sinistral process (often bifurcate); some species with medioventral process; dextral process absent; periandrium from left lateral aspect with broad and bifurcate (eastern U.S. taxa) or somewhat narrow and simple (western U.S. taxa) process (this process might be understood as originating from right dorsal periandrium); dorsal periandrium with long, large process originating sinistrally or mediobasally. Endosoma reduced, broad at base and either tapering to point (eastern U.S. taxa) or broad and rounded distally (western U.S. taxa); extending primarily caudad and very slightly curved left (ventral view); with (eastern U.S. taxa) or without (western U.S. taxa) elongate apical process; with (eastern U.S. taxa) or without (western U.S. taxa) at least one small subapical process. Anal tube roughly symmetrical, longer than wide, broad and hood-like, moderately concave apically (in dorsal view).

Distribution. Transcontinental U.S. and southern Canada (most abundant in eastern U.S.); Belize.

Etymology. The genus name is derived from the Greek word ξένος (xenos; alien, stranger) + Oliarus , a reference to the unusual saucer-shape of the medioventral process of the pygofer. The name is masculine in gender.

Remarks. This is the placitus -group of Mead & Kramer (1982). The male terminalia of this genus are highly distinctive. Males are readily separated (without dissection) from other Pentastirini by the enlarged, spatulate medioventral process of the pygofer. The aedeagal complex is quite different from any other group addressed in this work. The gonostyli of Xenoliarus gen. nov. are also highly distinctive in being asymmetrical and arcuate with the apices lobed. The endosoma and periandrium appear fused and are indistinct. Mead & Kramer (1982) note the distinctiveness of the endosoma, which is directed primarily caudad, compared to the strong sinistrally-angled form of the endosoma in most members of Melanoliarus s.l. They suggest the group may bear a close affinity to what we have here described as Cilidius gen. nov. (the quinquelineatus -group of Mead & Kramer 1982) which also shares the feature of the endosoma being directed caudad as opposed to curving left in ventral view; the group is also morphologically similar externally. However, Xenoliarus gen. nov. and Cilidius gen. nov. differ significantly in most other features of the male terminalia. Cilidius gen. nov. also differ in that they have conspicuously dark tubercles on the wing veins.

Xenoliarus gen. nov. differ from Melanoliarus s.s. (along with Bifoliarus gen. nov. and Vitroliarus gen. nov.) with their larger and broader form, also bearing a proportionately narrower head relative to the prothorax. Also helpful is that the maculae of the frons are present, although reduced in some species (usually well developed in Xenoliarus placitus comb. nov.). Some individuals (especially females of X. placitus comb. nov.) possess heavilymarked wings with veins intermixed pale and dark with dark tubercles.

The western U.S. taxa are notably smaller than the eastern species. In the eastern U.S., males of Xenoliarus montanus comb. nov. and X. placitus comb. nov. are 6.1–8.8 mm, while in the western U.S., males of X. eximus comb. nov. and X. teximus comb. nov. range from 4.7–6.4 mm. The western U.S. taxa also have different and distinctive forms of the aedeagal complex.

Mead & Kramer (1982) mention that members of this group also possess a “short, stout connective having unusually long ventral arms.” We are not treating the connective in the descriptions or diagnoses for these groups, but recognize that there is diagnostic value to its structure.

Xenoliarus placitus comb. nov. is one of the most common large taxa in the eastern US, and the species comes readily to lights. A light-trapping event by the second author in Frenchtown Woods, Glasgow, Delaware USA during the first week of July (during a new moon) yielded over 200 specimens of the species .

Xenoliarus teximus comb. nov. from different localities have highly variable male terminalia, including within the paratype series, as addressed in Mead & Kramer (1982). The geographic variation seen in this species deserves further exploration.

Species Composition

Xenoliarus eximus (Caldwell, 1947) , new combination.

Xenoliarus montanus ( Metcalf, 1923) , new combination.

Xenoliarus placitus ( Van Duzee, 1912) , new combination.

Xenoliarus teximus (Caldwell, 1947) , new combination.

Figures

FIGURE 1. American Pentastirini spp. in vivo. A) Xenoliarus placitus comb. nov. in Newark, Delaware, USA; B) Antilliarus slossoni comb. nov. in Monroe County, Florida, USA; C) Cilidius quinquelineatus comb. nov. in Groton, Massachusetts, USA; D) teneral Pentastirini sp. and nymph in Marin County, California, USA. Photographs taken by the first author (A), Judy Gallagher (B; CC-BY-NC-ND 1.0, used by permission), Tom Murray (C; CC-BY-NC-ND 1.0, used by permission), and Scott Loarie (D; CC0). B–C were initially sourced from the community science websites BugGuide.net and iNaturalist.org.

FIGURE 2. Morphological terminology of aedeagal complex, illustrations adapted from line art of Mead & Kramer 1982; A) ventral view of aedeagal complex (Amiarus (Umbrarus) humilis comb. nov.); B) dorsal view of aedeagal complex (Melanoliarus coconinus); annotations: Es = endosoma, EsA = apical process(es) of endosoma, EsD= dorsal process of endosoma,Aj = aedeagal joint, Per = periandrium, Sp = sinistral process of periandrium, Dp = dextral process of periandrium, Mvp = medioventral process of periandrium, Mdp = mediodorsal process of periandrium.

FIGURE 5. Melanoliarus, A–E Melanoliarus (Melanoliarus) complectus (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia.

FIGURE 6. Annotated left forewing of Melanoliarus, flipped vertically; from male M. (Amplectiarus) acicus comb. nov. Bold/ black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 7. Amiarus gen. nov., A–D Amiarus sp. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax; E) ventral view of cleared male terminalia.

FIGURE 8. Annotated left forewing of Amiarus gen. nov., flipped vertically; from male A. dimidiatus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 9. Amiarus (Umbrarus) subgen. nov., A–E Amiarus (Umbrarus) humilis comb. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia.

FIGURE 10. Annotated left forewing of Amiarus (Umbrarus) subgen. nov., flipped vertically; from A. (U.) humilis comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 11. Antilliarus gen. nov., A–D Oliarus borinquensis (holotype, male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax; E) ventral view of cleared male terminalia (Antilliarus nr. slossoni).

FIGURE 12. Annotated left forewing of Antilliarus gen. nov. flipped vertically; from male A. nr. slossoni. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 13. Bifoliarus gen. nov., A–D. Oliarus gladensis (holotype, male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax; E) ventral view of cleared male terminalia (Bifoliarus chuliotus comb. nov.).

FIGURE 14. Annotated left forewing of Bifoliarus gen. nov., flipped vertically; from male B. chuliotus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 15. Cilidius gen. nov., A–D Cilidius vicarius comb. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia (Cilidius quinquelineatus comb. nov.); annotations: ff = frontal fovea, lcr = lateral pronotal carina.

FIGURE 16. Annotated left forewing of Cilidius gen. nov., flipped vertically; from C. quinquelineatus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 17. Galapagoliarus gen. nov., A–D Oliarus galapagensis (holotype, female); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax.

FIGURE 18. Lapsium gen. nov., A–E Lapsium difficilis comb. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia.

FIGURE 19. Annotated left forewing of Lapsium gen. nov., flipped vertically; from L. difficilis comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 20. Oliaridellus gen. nov., A–E Oliaridellus apache comb. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia.

FIGURE 21. Annotated left forewing of Oliaridellus gen. nov., flipped vertically; from female O. apache comb. nov. Bold/ black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 22. Oliaridus gen. nov., A–D Oliaridus sonoitus comb. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia.

FIGURE 23. Annotated left forewing of Oliaridus gen. nov., flipped vertically; from male O. aridus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 24. Remotiarus gen. nov., A–D Oliarus forcipatus (holotype, male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia (Remotiarus yavapanus comb. nov.).

FIGURE 25. Annotated left forewing of Remotiarus gen. nov., flipped vertically; from male R. uncatus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins.

FIGURE 26. Sonorium gen. nov., A–D Oliarus dondonius (holotype, male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax; E) ventral view of cleared male terminalia (Sonorium altanatus comb. nov.).

FIGURE 27. Annotated left forewing of Sonorium gen. nov., flipped horizontally; from male S. zyxus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 28. Vitroliarus gen. nov., A–D Oliarus ecologus (holotype, male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax; E) ventral view of cleared male terminalia (Vitroliarus ecologus comb. nov.).

FIGURE 29. Annotated left forewing of Vitroliarus gen. nov., flipped vertically; from male V. ecologus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 30. Xenoliarus gen. nov., A–E Xenoliarus placitus comb. nov. (male); A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) ventral view of cleared male terminalia.

FIGURE 31. Annotated left forewing of Xenoliarus gen. nov., flipped vertically; from male X. placitus comb. nov. Bold/black = primary veins, bold/green = cells, italics/black = crossveins; wing nomenclature following Bourgoin et al. (2015).

FIGURE 32. Flata lunata syntype (undesignated Mead lectotype) (female), A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) labels.

FIGURE 33. Cixius testaceus holotype (female), A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) labels.

FIGURE 34. Oliarus cinereus lectotype, here designated (male), A) dorsal habitus view, B) lateral view, C) frontal view, D) head and thorax, E) labels.

FIGURE 37. Cixius fulvus holotype (female). A) dorsal habitus view, B) lateral habitus view, C) frontal view, D) head and thorax, E) labels.

FIGURE 41. Ventral aedeagal complexes of New World Pentastirini genera. A) Melanoliarus (Melanoliarus) complectus, B) Melanoliarus (Amplectiarus) acicus comb. nov., C) Amiarus (Amiarus) kindli comb. nov., D) Amiarus (Umbrarus) humilis comb. nov., E) Antilliarus slossoni comb. nov., F) Bifoliarus chuliotus comb. nov.; illustrations from Mead and Kramer 1982 (A–B, D–F) and Bourgoin et al. 1998 (C).

FIGURE 42. Ventral aedeagal complexes of New World Pentastirini genera. A) Cilidius vicarius comb. nov., B) Lapsium difficilis comb. nov., C) Oliaridellus apache comb. nov., D) Oliaridus aridus comb. nov., E) Remotiarus yavapanus comb. nov., F) Sonorium corvinus comb. nov.; illustrations from Mead and Kramer 1982.

FIGURE 43. Ventral aedeagal complex of New World Pentastirini genera A) Vitroliarus ecologus comb. nov., B) Xenoliarus placitus comb. nov., C) Nivcentia interrupta, D) Oliarissa armiger, E) Pentastiridius habeckorum, F) Reptalus (Pererepa); illustrations from Mead and Kramer 1982 (A–B, E–F), Fennah 1945a (C), and Fennah 1944 (D).

FIGURE 44. Male terminalia in situ, ventral view; A) Melanoliarus (Melanoliarus), B) Amiarus (Amiarus) gen. nov., C) Amiarus (Umbrarus) subgen. nov., D) Antilliarus gen. nov., E) Bifoliarus gen. nov., F) Cilidius gen. nov.

FIGURE 45. Male terminalia in situ, ventral view, A) Lapsium gen. nov., B) Oliaridellus gen. nov., C) Oliaridus gen. nov., D) Remotiarus gen. nov., E) Sonorium gen. nov., F) Vitroliarus gen. nov.

FIGURE 46. Male terminalia in situ, ventral view, A) Xenoliarus gen. nov., B) Oliaronus, C) Reptalus (Pererepa).