Descriptions of four new species of the Ambrysus signoreti Stål species complex (Heteroptera: Nepomorpha: Naucoridae) Reynoso-Velasco, Daniel Sites, Robert W. Zootaxa 2019 2019-06-13 4615 2 285 302 Reynoso Reynoso-Velasco & Sites 2019 [151,453,153,180] Insecta Naucoridae Ambrysus Animalia Hemiptera 13 298 Arthropoda species sitesi sp. nov.  ( Figs. 3, 8–9)  Description. Macropterous female. HOLOTYPE, length 11.87; maximum width 7.25. Paratypes(n = 10), length 10.70–11.60 (mean = 11.31); maximum width 6.80–7.50 (mean = 7.17). General shape ovate; widest across embolia ( Fig. 8A). Overall dorsal coloration of hemelytra brown to dark brown with yellowish brown marks; head and pronotum light brown with dark brown marks; legs yellowish brown. Dorsal surface coarsely punctate. Ventral coloration of head and thorax yellowish, abdomen with yellowish pubescence.  Head. Head length 1.60; maximum width 3.20. Mostly light brown with dark brown marks along midline and posteriorly; coarsely punctate. Eyes convergent anteriorly, synthlipsis 1.48; thin band of cuticle along posterolateral margin of eye; eyes not raised above level of vertex or pronotum. Anterior margin between eyes slightly convex, extending anteriorly in front of eyes 7.8% of head length; posterior margin between eyes strongly convex, extending posteriorly 29.7% of head length. Labrum width 1.7× length, evenly rounded. Labium with three visible yellowish brown segments, darkening distally, extending 0.49 beyond labrum not including extruded stylets. Antennal proportions 3:14:22:9, length 0.72, extending to lateral margin of eye, elongate hairs on segment 4 and distal half of 3.  Thorax. Pronotum coarsely punctate, ground color medium brown, profuse dark brown markings, dark brown longitudinal line behind eye; transverse sulcus marking anterior border of transverse band in posterior 1/3; transverse band light brown anteriorly, brown posteriorly; lateral margins yellowish brown, convergent, evenly convex, explanate; posterior margin almost straight; anterior margin deeply concave between eyes to embrace convex posterior margin of head; posterolateral corners rounded; width 2.7× length; length at midline 2.28; maximum width at posterolateral corners 6.26. Prothorax ventrally pruinose throughout, except laterally; apices of propleura meeting at midline, not appressed to prosternellum; propleuron yellowish, light brown posteriorly, medial 2/3 of posterior margin with golden setae. Probasisternum with sharp median carina and row of setae lateral to carina. Prosternellum not covered by apices of propleura. Scutellum coarsely punctate, triangular, dark brown, light brown coloration at anterior corners and along midline, width 2.1× length, width 4.32, length 2.00. Hemelytra densely punctate, brown to dark brown, length 8.12 (chord measurement); corium dark brown, yellowish brown ovate and yellowish commashaped marks at lateral and posterior margins, respectively. Clavus dark brown in lateral half, brown in mesal half with dark brown close to base and apex, length 3.8× width, length 4.56, width 1.20; claval commissure dark brown, length 1.30. Embolium length 3.32, greatest width 1.05; lateral margin convex, yellowish in anterior 2/3, brown posteriorly. Membrane dark brown ( Fig. 8A). Hindwings extending to middle of tergum VII. Mesobasisternum tumescence negligible, with sulcus on midline continuing through triangular mesosternellum. Metasternellum (= metaxyphus) transverse, subtriangular, with median carina.  Legs. All leg segments yellowish brown. Profemur yellowish brown; posterior margin with row of tightly arranged setae in basal 2/3, row of short brown spines along middle third; anterior margin with dense pad of setae without associated spines. Protibia and -tarsus with occlusal inner surface flattened and with spatulate setae; tarsus immovable, one-segmented; pretarsal claw single, minute, triangular. Procoxa with cluster of stout, brown anteromedial spines. Meso- and metacoxae partially recessed into thorax. Meso- and metafemora with row of short, light brown spines on anterior margin; spines restricted to basal half on mesofemur, nearly full length of metafemur. Mesotibia with ventrolateral and dorsolateral rows of stout reddish-brown spines; mesolateral rows intermixed with combs of two to six spines. Metatibia with ventrolateral and dorsolateral rows of stout reddish-brown spines. Meso- and metatibiae with semi-circlet of spines at apices on dorsal and mesal margins; two comb rows of stout spines near apex of ventral margin. Meso- and metatibiae and metatarsus with long, yellow swimming hairs; hairs profuse on metatibia and -tarsus. Meso- and metapretarsi with paired claws slender, gently curved, with small basal tooth. Leg measurements as follows: foreleg, femur 2.60, tibia 2.02, tarsus 0.47; middle leg, femur 2.55, tibia 2.30, tarsomeres 1–3, 0.22, 0.49, 0.52; hind leg, femur 3.34, tibia 3.72, tarsomeres 1–3, 0.28, 0.88, 0.67.  Abdomen. Dorsally with lateral margins of III–VI exposed, each brown in anterior 1/3 and light brown posteriorly; lateral margins of II to basal half of III smooth and distal half of III to VI finely serrate, marginal row of profuse and short yellow setae, group of trichobothria near posterolateral corners. Posterolateral corner of II (visible ventrally) narrowly rounded, right angled, III–VII acute. Ventrally yellowish, with dense pile of fine hairs. Lateral margin with thin, glabrous band. Glabrous rounded to oval patches near spiracles on laterosternites II–VI. Mediosternite VII (subgenital plate) asymmetrical, width 1.1× length; length at midline 1.24; maximum width 1.38; lateral lobe at mid-length of left margin broadly rounded; posterolateral corners pointed; wide central lobe produced posteriorly slightly further than posterolateral corners ( Fig. 8B).   FIGURE 8.Structures of  Ambrysus sitesi  n. sp.(A) dorsal habitus of holotype, (B) sternum VI and subgenital plate of female (terminal abdominal segments removed), (C) 6th and 7th abdominal terga of male, (D) 8th abdominal tergum of male, (E) ventral surface of male phallosoma, (F) genital capsule of male (phallosoma removed). ll = lateral lobe, plc = posterolateral corner.  Macropterous male. Paratypes(n = 10), length 9.80–11.00 (mean = 10.52); maximum width 6.20–7.00 (mean = 6.62). Coloration, setation, pronotum proportions, posterolateral corners of pronotum, scutellum proportions, embolium length generally same as for holotype. Otherwise hindwings extending to posterior margin of tergum V. Accessory genitalic process of tergum VI slender, gently curved to right close to the base, widening distally, apex pointed ( Fig. 8C). Medial lobes of tergum VIII (pseudoparameres) symmetrical, posterolaterally produced, posterolateral corner narrowly rounded, extending posteriorly further than posteromesal corner ( Fig. 8D). Phallosoma elongate, linear, slender, slightly constricted basally; left endosomal sclerite reduced, with small and inconspicuous denticles; right endosomal sclerite absent ( Fig. 8E). Parameres symmetrical, almost as long as maximum width, mesal margin rounded, setae emanating from distal 1/3 of dorsal surface. Proctiger slightly shorter than width at base. Pygophore with elongate setae sparsely distributed over most of surface, with thick brush of elongate setae on posterior margin ( Fig. 8F).   Discussion.The conspicuous dorsal pigmentation of specimens of  A. sitesiis similar to that of  A. lunatusUsingerand  A. signoretiStål. Themorphology of the endosomal sclerites of the males suggests that this species could be closely related to  A. lundbladiUsinger. Inhis work on the  Ambrysusof Mexico, La Rivers (1953)described the subspecies  A. lunatus menoidesfrom the central state of Puebla. He cited paratypespecimens from the state of Chiapas(Ixtapa) that were deposited in the UMMZ. We examined a female paratypethat was deposited in the CAS and found that the morphology of the SGP was different to that reported for the subspecies. Instead, the morphology corresponded to the herein described  A. sitesi. Based on this and our extensive collecting in the area, we consider the subspecies A. l. menoidesnot to be distributed in the state of Chiapas.  Diagnostic features and comparative notes.Females of this species exhibit a characteristic SGP with a broadly rounded lateral lobe at mid-length of the left margin. Other species in the  A. signoreticomplex present a lateral lobe on the left margin of the SGP (sometimes also on the right margin), but in general the lobe is low and pointed (e.g.,  A. colimanus,  A. lunatus). The species  A. scalenuspresents a similar left-lobed margin of the SGP (examined on the allotypein the SEMC); however, the lobe extends further laterally and appears truncate. Males of  A. sitesipresent the phallosoma with no right endosomal sclerite and a reduced left endosomal sclerite with inconspicuous rounded denticles along the mesal margin. To our knowledge, the rare  A. lundbladiis the only other species from this species complex that exhibits the phallosoma with an absent right endosomal sclerite and the left sclerite reduced.   Habitat description.The typelocality at Río Rana (L-1906), close to the town of San Gabriel Mixtepec, is in the south central area of the Mexican state of Oaxaca, on the Sierra Madre del Sur mountain range, at an elevation of 712 meters. At the typelocality the stream is 5–10 meters wide, with fast current, and boulders and cobble ( Fig. 9). Specimens were collected by disturbing the substratum with the net along the stream margin.   Distribution.  Ambrysus sitesiis distributed in the highlands of the Mexican states of Chiapas and Oaxaca and the western area of Guatemala, in the Huehuetenango Department( Fig. 3). The records from Oaxaca are in the area of the Sierra Madre del Sur biogeographic province, while those from Chiapas and Huehuetenangoare in the Chiapas biogeographic province. One record from Chiapas (L-1899) is in the Mexican Pacific Coast biogeographic province. This species was collected syntopically with various congeners, including  A. circumcinctus(L-1339, L- 1341, L-1350, L1884, L1897),  A. circumcinctusand  A. pudicus(L-1881, L-1898),  A. circumcinctusand  A. spicatus(L-1349),  A. circumcinctus,  A. parviceps, and  A. pudicus(L-1899);  A. chiapanecusReynoso, Sites & Novelo(L- 1340, L-1886);  A. chiapanecusand  A. spicatus(L-1885);  A. mexicanusMontandon,  A. pygmaeusLa Rivers, and  A. spicatus(L-1397). At its typelocality (L-1906) this species was found syntopically with  A. baeus,  A. henryi  n. sp.,  A. schuhi  n. sp., and  A. spicatus. Additionally,  A. sitesiwas collected with  A. brailovskyi  n. sp.at Finca Prusia ( 12-V-1985), Río Matasano (L-1348), Río Prusia (L-1345), and the Tenejapa municipality ( 14-VII-1964).   Etymology.It is my (DRV) pleasure to name this species in honor of Professor Robert. W. Sites, my former doctoral advisor at the University of Missouriand coauthor of this manuscript. I am grateful for his guidance, help, and friendship throughout these years.  Repositories.The holotypeand some paratypeswill be deposited in the Colección Entomológica del Instituto de Ecología A.C. (Xalapa); additional paratypeswill be deposited in the California  Academy of Sciences( SanFrancisco), Colección Nacional de Insectos(Mexico City), Enns Entomology Museum (Columbia), Snow Entomological Museum Collection (Lawrence), and the United States National Museum of Natural History( Washington D.C.)   Type material examined.  HOLOTYPE ♀.  MÉXICO: OAXACA: Mpio. San Gabriel Mixtepec, km 194 carr. Puerto Escondido-Oaxaca, SanGabriel Mixtepec, Río Rana, 16º06’06.3’’N, 97º03’52.9’’W,  712 m,  03-Apr-2015, L-1906, DRV, Sites, Shepard, Barr & PRH colls.  PARATYPES: same data as holotype ( 2♂, 2♀ CNIN; 10♂, 7♀IEXA; 5♂, 4♀ UMC);  Mpio. Jalapa de Díaz, Sto. Domingo, Río Sto. Domingo, 18°02’15.9’’N, 96°32’57.2’’W,  100 m, L-1881,  25-Mar-2015, DRV, Sites, Shepard, Barr& PRH colls. ( 2♂, 6♀ IEXA; 1♂, 1♀ UMC);  Mpio. Putla Villade Guerrero, Plutade Guerrero, 17º02’21.5’’N, 97º54’42.9’’W,  767 m,  30-May-2012, L-1397, DRV & PRH colls. ( 1♂ UMC);  Mpio. Sta. Maria Tecomavaca, carr. Tehuacán-San Francisco Telixtlahuaca,  16 kmS of San Juande los Cués, Río Salado, 17º55’50.5’’N, 97º00’04.2’’W,  544 m,  01-Jun-2012, L-1407, DRV & PRH colls. ( 1♂ UMC).   CHIAPAS: Mpio. Altamirano, Cuauhtemoc, Balneario Tzaconejá, Río Tzaconejá, 16°41’17.5’’N, 92°01’03.0’’W,  1172 m,  28-Mar-2015, L-1886, DRV, Sites, Shepard, Barr& PRH colls. ( 6♂, 3♀ IEXA; 4♂ UMC);  Mpio.Am- atengo de la Frontera, Barrio Sabinalito, Río Agua Caliente, 15°25’0.1’’N, 92°08’33.7’’W,  942 m, L-1898,  31-Mar- 2015, DRV, Sites, Shepard, Barr& PRH colls. ( 4♂ IEXA); [Mpio. Ángel Albino Corzo], Finca Prusia,  10-V-85, M. Vértiz/ Colección del Instituto de Biología, UNAM. México, D.F ( 1♂ CNIN); [Mpio. Ángel Albino Corzo], Finca Prusia,  12-V-85, F Arias/ Colección del Instituto de Biología, UNAM. México, D.F ( 1♂, 2♀ CNIN); [Mpio. Ángel Albino Corzo], Finca Prusia,  12-V-85, H Velasco/ Colección del Instituto de Biología, UNAM. México, D.F ( 3♂ CNIN); [Mpio. Ángel Albino Corzo], Finca Prusia,  12-V-85, M Vértiz/ Colección del Instituto de Biología, UNAM. México, D.F ( 1♀ CNIN);  Mpio. Ángel Albino Corzo,  1 kmS of Finca Prusia, Río Prusia, 15º43’23.3’’N, 92º47’46.5’’W,  1010 m,  18-May-2012, L-1345, DRV & PRH colls. ( 1♂, 4♀ UMC); Mpio. Frontera Comalapa, El Caracol,  8 kmS of Frontera Comalapa, Río Grijalva, 15°36’07.0’’N, 92°09’14.5’’W,  677 m,  31-Mar-2015, L- 1897, DRV, Sites, Shepard, Barr& PRH colls. ( 1♂ IEXA);  Mpio. Huixtla, Huixtla, Río Huixtla, 15°08’59.9’’N, 92°27’04.6’’W,  67 m,  31-Mar-2015, L-1899, DRV, Sites, Shepard, Barr& PRH colls. ( 1♂ IEXA);  Mpio. Ixtapa, Ixtapa, 16º48’43.7’’N, 92º54’30’’W,  1030 m,  17-May-2012, L-1341, DRV & PRH colls. ( 6♂, 3♀ IEXA); [Mpio. Ixtapa], Ixtapa, #132,  IV-27-1941, IJ Cantrall/  Ambrysus lunatus menoidesLa Rivers  PARATYPE/ Ira La Rivers Collection Bequeathedto the California Academyof Sciences-1978 ( 1♀ CAS);  Mpio. Ixtapa, Río Lajasat Pte. Río Lajas, 16º50’42.6’’N, 92º54’01.7’’W,  1020 m,  17-May-2012, L-1340, DRV & PRH colls. ( 2♂, 1♀ UMC); same but  28-Mar-2015, L-1885, DRV, Sites, Shepard, Barr& PRH colls. ( 10♂, 10♀ CNIN; 20♂, 21♀IEXA; 6♂, 15♀ UMC); [Mpio. Ixhuatán],  4 miN of Tapiula[Tapilula], CL 1097, 5 May1964, JT & MS Polhemus( 1♂, 1♀ USNM); [Mpio. Ocosingo], Ocosingo,  June 25 1950, C & M Goodnight& L Stannard, mountain stream / Chiapas  VI-25-57, Mex. Stannard( 1♂ SEMC);  Mpio. Siltepec, km 72 carr. Jaltenango-Siltepec, nr. Nuevo Galeana, 15º35’54.9’’N, 92º32’28.9’’W,  933 m, L-1349,  19-May-2012, DRV & PRH colls. ( 2♂, 1♀ UMC);  Mpio. Siltepec, km 92 carr. Jaltenango-Siltepec, Campo Aéreo, Río Bravo, 15º35’17.9’’N, 92º28’02.3’’W,  1068 m,  19-May-2012, L-1350, DRV & PRH colls. ( 1♂, 1♀ UMC);  Mpio. Siltepec, Matasano, Río Matasano, 15º36’13.7’’N, 92º33’37.1’’W,  950 m,  19-May-2012, L-1348, DRV & PRH colls. ( 6♂, 6♀ IEXA; 16♂, 19♀ UMC);  Mpio. Solosuchiapa, El Beneficio, Río La Sierra, 17°21’01.1’’N, 93°00’21.1’’W,  303 m,  27-Mar-2015, L-1884, DRV, Sites, Shepard, Barr& PRH colls. ( 1♂ UMC);  Mpio. Solosuchiapa, Solosuchiapa, Río La Sierra, 17º25’40.5’’N, 93º01’35.7’’W,  122 m,  17- May-2012, L-1339, DRV & PRH colls. ( 1♂ UMC); [Mpio. Soyaló], Soyalo, Rio Blanco, CL 1086, 4 May1964, JT & MS Polhemus/ C.J. Drake Accession( 16♂, 10♀ USNM);  Mpio. Tenejapa, ponds in stream above Habenal/  14.VII.1964, coll. D Breedlove/ C.J. Drake Accession/ CAS ( 2♂ USNM);  Rio Laja[s] nr Bochil, CL668,  I-6-74, JT Polhemus( 9♂, 5♀ USNM);  Soyalo, Rio Lajas, CL 1087, 4 May1964, JT & MS Polhemus/  A. guttatipennisclose, No! / C.J. Drake Accession( 1♀ USNM).   GUATEMALA: HUEHUETENANGO: Rio Nentón, ca.  8 kmNE of La Laguna,  10-May-2013, 15.79915º N, 91.88951ºW,  613 m, RS Zackcoll. ( 1♀ WSUC). 2252532372 United States of America Academy of Sciences Coleccion Nacional de Insectos San 16 301 1 District of Columbia 2252532386 2015-04-03 Mexico Mpio. San Gabriel Mixtepec 712 16.101751 San 1 -97.0647 Puerto Escondido-Oaxaca 16 301 1 1 Oaxaca holotype 2252532365 2015-04-03 CNIN, UMC Mexico Mpio. San Gabriel Mixtepec 712 16.101751 San 1 -97.0647 Puerto Escondido-Oaxaca 16 301 30 13 17 Oaxaca paratype 2252532393 2015-03-25 IEXA, UMC de Diaz & Sto. Domingo & Sites & Shepard & Barr Mexico 100 18.03775 Rio Sto. Domingo 1 -96.549225 Mpio. 16 301 10 7 3 Jalapa paratype 2252532410 2012-05-30 UMC Mexico 767 17.039305 Pluta 1 -97.91192 Mpio. Putla Villa 16 301 1 1 Guerrero paratype 2252532377 2012-06-01 UMC Mexico Mpio. Sta. Maria Tecomavaca 544 17.930695 Cues 1 -97.00117 Tehuacan-San Francisco Telixtlahuaca 16 301 1 1 Oaxaca paratype 2252532366 2015-03-28 IEXA, UMC Sites & Shepard & Barr Mexico Mpio. Altamirano 1172 16.688194 Balneario Tzaconeja 1 -92.0175 Cuauhtemoc 16 301 13 3 10 Chiapas paratype 2252532403 2015-03-31 IEXA de la Frontera & Barrio Sabinalito & Sites & Shepard & Barr Mexico 942 15.416695 Rio Agua Caliente 1 -92.14269 Mpio. 16 301 4 4 Chiapas paratype 2252532402 1985-05-10 CNIN M. Vertiz Mexico Coleccion del Instituto de Biologia Finca Prusia 16 301 1 1 Mexico paratype 2252532385 1985-05-12 CNIN F Arias Mexico Coleccion del Instituto de Biologia Finca Prusia 16 301 3 2 1 Mexico paratype 2252532356 1985-05-12 CNIN H Velasco Mexico Coleccion del Instituto de Biologia Finca Prusia 16 301 3 3 Mexico paratype 2252532379 1985-05-12 CNIN M Vertiz Mexico Coleccion del Instituto de Biologia Finca Prusia 16 301 1 1 Mexico paratype 2252532369 2012-05-18 UMC Mexico 1010 15.723139 Rio Prusia 1 -92.79625 Mpio. Angel Albino Corzo 16 301 5 4 1 Chiapas paratype 2252532351 2015-03-31 IEXA El Caracol & Sites & Shepard & Barr Mexico 677 15.601945 Rio Grijalva 1 -92.15403 Frontera Comalapa 16 301 1 1 Chiapas paratype 2252532359 2015-03-31 IEXA Sites & Shepard & Barr Mexico Mpio. Huixtla 67 15.149972 Rio Huixtla 1 -92.45127 Huixtla 16 301 1 1 Chiapas paratype 2252532413 2012-05-17 IEXA Mexico 1030 16.812138 Ixtapa 14 -92.90833 Mpio. Ixtapa 16 301 9 3 6 Chiapas paratype 2252532395 1941-04-27 IJ Cantrall Mexico Ixtapa 16 301 1 Chiapas paratype 2252532352 CAS California Academy Mexico Ira La Rivers Collection Bequeathed 16 301 1 1 Chiapas paratype 2252532391 2012-05-17 UMC Mexico Mpio. Ixtapa 1020 16.845167 Rio Lajas 1 -92.900475 Rio Lajas 16 301 3 1 2 Chiapas paratype 2252532367 2015-03-28 CNIN, UMC Sites & Shepard & Barr Mexico 16 301 82 46 36 Chiapas paratype 2252532376 [238,1432,1844,1871] USNM May & MS Polhemus Mexico 4 mi N of Tapiula 16 301 2 1 1 Chiapas paratype 2252532371 1950-06-25 1957-06-25 1950-06-25 SEMC C & M Goodnight & L Stannard & Mex. Stannard Mexico Ocosingo 16 301 1 1 Chiapas paratype 2252532361 2012-05-19 UMC Mexico Mpio. Siltepec 933 15.598583 Nuevo Galeana 1 -92.54136 Jaltenango-Siltepec 16 301 3 1 2 Chiapas paratype 2252532362 2012-05-19 UMC Mexico Mpio. Siltepec 1068 17 302 15.588305 Campo Aereo 1 -92.46731 Jaltenango-Siltepec 16 301 2 1 1 Chiapas paratype 2252532357 2012-05-19 IEXA, UMC Mexico Mpio. Siltepec 950 15.603806 Rio Matasano 1 -92.56031 Matasano 17 302 47 25 22 Chiapas paratype 2252532353 2015-03-27 UMC El Beneficio & Sites & Shepard & Barr Mexico 303 17.350306 Rio La Sierra 1 -93.00586 Mpio. Solosuchiapa 17 302 1 1 Chiapas paratype 2252532364 2012-05-17 UMC Mexico Mpio. Solosuchiapa 122 17.427916 Rio La Sierra 1 -93.02659 Solosuchiapa 17 302 1 1 Chiapas paratype 2252532415 USNM Soyalo & May & MS Polhemus & C. J. Drake Accession Mexico Rio Blanco 17 302 26 10 16 Chiapas paratype 2252532384 1964-07-14 USNM D Breedlove & C. J. Drake Accession Mexico Habenal Mpio. Tenejapa 17 302 2 2 Chiapas paratype 2252532408 1974-01-06 USNM Bochil, CL & JT Polhemus Mexico Rio Laja 17 302 14 5 9 Chiapas paratype 2252532374 USNM Soyalo & May & MS Polhemus & No & C. J. Drake Accession Mexico Rio Lajas 17 302 1 1 Chiapas paratype 2252532368 2013-05-10 WSUC RS Zack Guatemala 613 15.79915 Rio Nenton 1 -91.88951 17 302 1 1 Huehuetenango paratype