Phylloicus aeneus ( Hagen, 1861:285 ) Macronema Anisocentropus Ulmer 1905a:79 Phylloicus P. nigripennis Ross 1952:34 Flint 1967:17 P. nigripennis Denning et al. 1983:182 P. mexicanus Phylloicus ornatus ( Banks, 1909:342 ) Notiomyia Flint 1967:17 Phylloicus Holzenthal 1988:72 Phylloicus centralus ( Navás, 1924:82 ) Macronema Holzenthal 1988:72 Phylloicus P. aeneus P. nigripennis mexicanus latus sagittosa P. aeneus mexicanus nigripennis latus sagittosa ornatus centralus aeneus Revision of the Neotropical caddisfly genus Phylloicus (Trichoptera: Calamoceratidae) PRATHER, AYSHA L. Zootaxa 2003 2003-08-29 275 1 1 214 4H7JS (Hagen) Hagen [264,593,523,549] Insecta Calamoceratidae Phylloicus Animalia Trichoptera 20 21 Arthropoda species aeneus   Figs. 1, 6, 15, 16, 118      Phylloicus aeneus( Hagen, 1861:285)[ Typelocality: Mexico[ Veracruz]; MCZ; female; in  Macronema]; 1864:804 [to  Anisocentropus]. —  Ulmer 1905a:79[to  Phylloicus, redescription of male now in  P. nigripennis].  Ross 1952:34[male]. —  Flint 1967:17[illustration of male now in  P. nigripennis]. —  Denning et al.1983:182[redescription, illustration is probably of  P. mexicanus]. —Bowles & Flint 1997:58 [variation].     Phylloicus ornatus( Banks, 1909:342)[ Typelocality: United States, Texas, Brownwood; MCZ; female; in  Notiomyia]. —  Flint 1967:17[to  Phylloicus]. —  Holzenthal 1988:72[distribution]. — Bowles & Flint 1997:53 [redescription, male, female, larva, pupa]. NEW SYNONYM.     Phylloicus centralus( Navás, 1924:82)[ Typelocality: Costa Rica; MNHNP; male; in  Macronema]. —  Holzenthal 1988:72[to  Phylloicus]. NEW SYNONYM. The identity of  P. aeneushas been the subject of some confusion, largely due to the holotypebeing a female in very poor condition. Several authors have redescribed this species based on specimens mistakenly associated with the type female, and thus judged  P. nigripennis, mexicanus,  latus, and  sagittosato be conspecific with  P. aeneus. Based on the evidence discussed below,  mexicanusand  nigripennisare valid species — with  latusand  sagittosajunior synonyms of the latter — and  ornatusand  centralusare synonyms of  aeneus.   Hagen (1861)described  Macronema aeneumfrom a female specimen labeled “Sallé, V.C.” [for Veracruz, Mexico, handwritten on a single green label]. Ross (1952)identified this type specimen in a paper on lectotypesof Hagen species in the Museum of Comparative Zoology. He did not refer to the specimen as a lectotypein the paper. A yellow label in Ross’s hand on the specimen names it lectotype, but there is nowhere in Hagen’s original description or Ross’s 1952 paper (except by implication in the paper’s title) any indication that it was selected from among the type series. This specimen is badly damaged; the abdomen, which has been cleared and placed in a microvial of glycerin, is in many pieces; only a portion of one forewing remains, and even this is faded, so that it is not possible to determine what color pattern it might once have exhibited, although the epithet “  aeneum”translates as “coppery.” The holotypefemale of  M. aeneumbears a tuft of dark stout setae at each posterolateral margin of tergite VIII. The female holotypeof  Notiomyia ornatawas collected in central Texas. It is in good condition and the wing pattern is clearly discernable. The pattern is very distinctive, and the species can be easily discriminated on the basis of wing pattern alone. The wings can be described as copper­colored. The tufts of setae on tergite VIII observed in  P. aeneusare present also on the female holotypeof  N. ornata. I have seen these setal tufts only on females of  P. aeneus, ornatus, mexicanus,and  panamensis, new species. Specimens of  P. mexicanushave other distinguishing characters that rule out conspecificity; of the remaining two species, only  P. ornatusis known to be from Mexico. The terminalia of the type specimens of  P. aeneusand  ornatusare indistinguishable. Only one form of  Phylloicusis found in Texas, and thus there is no doubt of the association of male and female. The males bear short coremata on abdominal segment IV, with very short, broad projections of the posterolateral tergal margin. The male holotypeof  M. centralehas terminalia and modifications of abdominal segment IV identical with those observed in  P. ornatusmales. The wing pattern is faded, but what remains is consistent with the distinctive pattern of  P. ornatus. Ulmer redescribed  P. aeneusbased on several specimens from Morelos; these specimens appear to be conspecific with the typeof  P. nigripennis. Flint (1967)compared the female typesof  aeneusand  nigripennisand concluded that they were conspecific; he illustrated a male labeled “ Mexico, Sallé,” stating that it was probably part of the original typeseries. I have examined this and another specimen labeled “ Mexico, Sallé,” and labeled by Flint in 1965, as  Phylloicus aeneusmale. The collection labels on these specimens are printed (possibly cut from a journal article) with “ Mexico” on the upper label, and “Sallé” on the second label; these labels do not resemble those on the typeof  aeneusexcept in the collector’s name. These specimens are males of  P. nigripennis; the wings are dark and without pattern, no coremata are present on the abdomen, and they exhibit other characteristics consistent only with that species. The redescription of  P. aeneusby Denning et al.(1983)includes a mention of the abdominal coremata. Their illustrations were prepared from specimens collected in Sonora, which may have been either  P. aeneusor  mexicanus; if the specimens depicted in the illustration had abdominal coremata, they must have been  P. aeneus. Given this evidence, the logical conclusion is that  P. aeneus, centralus,and  ornatusare conspecific. Pinned specimens of  P. aeneusare easily identified by the distinctive pattern of dark patches near the base of the forewing. As well, in  P. aeneusand  mexicanus,  nigripennis, and  panamensis, there is an oblique fold of the forewing’s apical third, toward the midline ( Fig. 118). The male genitalia of  P. aeneusare most similar to those of  P. mexicanus, nigripennis,and  panamensis.  Phylloicus aeneusmay be distinguished from these three species by the presence of short, single­lobed coremata, covered by a short posterior tergal process, on abdominal segment IV. Adult. Forewing length 9.6­12.3 mm, n = 201. Head orange­brown. Maxillary palps black. Antenna twice forewing length; dark brown, with narrow patches of pale sensilla on anteromesal surface of each flagellomere. Dorsal pterothorax orange­brown, lateral margins black; ventrolateral thorax golden brown. Femora golden brown; foretibiae pale tan; mesotibiae white proximally, dark distally; metatibiae white proximally, dark distally; tarsi black. Metathoracic leg of male without posterior fringe. Tibial spur formula 2,4,3. Forewing apical third folded obliquely toward midline; chestnut brown; with two transverse bands; proximal band golden, not reaching either wing margin, 1/2 width of wing or less; distal band golden, extending from anterior to posterior wing margins, zig­zagging, with golden oval surrounding nygma; with single basal stripe, golden; with large dark patch anterobasally and golden semicircle posterobasally ( Figs. 105, 118). Hind wing basal brush present in female, orange. Male. Preterminalic abdominal terga with anteromesal notch. Corematic structures present. Tergum IV with expanded lateral flanges, lateral coremata; lateral coremata with basal globose lobes. Tergum V without sclerotized modifications ( Fig. 15F). Sternum VII with short, acute anteromesal process. Sternum VIII similar to anterior sterna, sternum IX not elongate. Tergum IX with mesal ridge extending 2/3 length of segment; posterior margin smoothly rounded; posteromesal surface covered with fine pilosity ( Fig. 15B); lateral ridge absent; dorsal pleural setae approximately 15, ventral pleural setae approximately 5 ( Fig. 15A); sternum IX with paired mesolateral ridges; sternum IX ( Fig. 15C). Preanal appendage approximately length of tergum X, of uniform diameter throughout length, setae long, but not filamentous or longer than appendage ( Fig. 15A, B). Tergum X sagittate basally; basodorsal process short and bifid; basolateral processes absent; apex, in lateral view, acute, in dorsal view, notched, notch square; with 2 longitudinal rows of short setae mesally; with paired shallow processes on lateral surfaces ( Fig. 15A, B). Harpago long, curving mesally and tapering apically; peglike setae few, apical ( Fig. 15A, C). Phallic endotheca with paired basolateral lobes, basolateral lobes large and round; phallotremal sclerites average size, longest dimension less than diameter of phallobase; dorsal sclerite narrow, width less than height ( Fig. 15D, E). Female. Abdominal terga I­III black. Preterminal abdominal terga with anteromesal notch. Sternum VII with short pointed anteromesal process. Tergum VIII with dense posterolateral brush of setae ( Fig. 16B); sternum VIII cleft posteromesally, cleft not reaching anterior ridge; sternum VIII ( Fig. 16C). Tergum IX with mesal ridge extending length of segment ( Fig. 16B). Sternum IX anterior and posterior lobes darkly sclerotized and striate, with deep sublateral invaginations ( Fig. 16A). Tergum X appendage longer than mesal lobe, base indistinct, apex triangular; mesal lobe lightly sclerotized; digitate lateral processes absent ( Fig. 16B). Sternum X without setae in membrane ( Fig. 16A). Vaginal apparatus anterior and posterior sclerites equal in length; anterior sclerite rounded anteriorly, posterolateral projections rounded; posterior sclerite ovoid; posterior end of spermatheca a sclerotized ovoid ( Fig. 16A).   Materialexamined.  BELIZE: Caya: Rió Frio, mouth of Rió Frio Cave, San Ignacio,  18.vi.1975, Resh— 2 males, 1 female( UMSP);   COSTA RICA:1920, Serre—  P. centralus holotypemale ( MNHNP);   Alajuela:  20 kmS Upala,  1­20.xi.1990, Parker— 1 female( EMUS);  Río Toro, 3.0 km (road) SW Bajos del Toro, 10°12'14”N, 84°18'58"W,  1530 m,  3­4.ix.1990, Holzenthal, Blahnik, & Huisman— 1 male, 2 females( NMNH);  Reserva Forestal San Ramón, Río San Lorencito& tribs., 10°12'58”N, 84°36'25"W,  980 m,  1­4.x.1986, I & A Chacón— 1 male( UMSP);   30.iii.­1.iv.1987, Holzenthal, Hamilton, & Heyn— 1 male( UMSP);   1­4.v.1990, Holzenthal& Blahnik— 1 female( UMSP);   28­ 30.vii.1990, Holzenthal, Blahnik, & Muñoz— 1 female( UMSP);   6­10.ii.1991, Holzenthal, Muñoz, & Huisman— 2 males, 1 female( UMSP);   Cartago: Reserva Tapantí, Quebrada Palmitos& falls, ca. 9 km(road) NW tunnel, 09°43'12”N, 83°46'48"W,  1400 m,  24­25.iii.1991, Holzenthal, Muñoz, & Huisman— 1 male( UMSP);  Río Grande de Orosí, 09°41'10”N, 83°45'22"W,  1650 m,  23.i.1985, Janzen& Hallwachs— 1 male( NMNH);   Guanacaste:Cerro ElHache, W side,  310 m,  31.v.1986, Janzen& Hallwachs— 1 male, 1 female( UMSP);  Finca Montezuma, 3 kmSE R. Naranjo,  17.xi.1991, Parker— 1 female( EMUS);   20­31.i.1992, Parker— 1 female( EMUS);   v.1992, Parker— 1 female( EMUS);   16­31.v.1993, Parker— 1 female( EMUS);  Maritza, confluence with Río Tempisquito, 10°57'25”N, 85°29'42"W,  580 m,  31­19.v.1994— 1 male& pupal exuviae, 1 female( UMSP);   14.ii.­1.v.1994— 2 males& pupal exuviae, 2 females( UMSP);  Maritza, Rio Orosi, 10°59'28”N, 85°25'42"W,  675 m,  31.i.­27.v.1994— 2 males& pupal exuviae, 1 female( UMSP);  Quebrada Garcia, Quebrada Garcia, 10.6 km ENE Quebrada Grande, 10°51'43”N, 85°25'41"W,  470 m,  8.iii.1986, Holzenthal& Fasth— 1 female( UMSP);  Río Los Ahogados, Río Los Ahogados, 11.3 km ENE Quebrada Grande, 10°51'54”N, 85°25'23"W,  470 m,  7.iii.1986, Holzenthal& Fasth— 1 male( UMSP);  Parque Nacional Guanacaste, Estacion Maritza, Río Tempisquito, 10°57'29”N, 85°29'49"E,  550 m,  29.i.1992, Sweeney— 1 male( NMNH);  Río Orosí, Estación Pitilla, 10°59'28”N, 85°25'41"W,  700 m,  22­25.v.1990, Holzenthal& Blahnik— 1 female( UMSP);  Río San Josecito, 10°55'19”N, 85°28'12"W,  960 m,  3­4.iv.1987, Holzenthal, Hamilton, & Heyn— 1 male( INBIO);  Río Tempisquito, Maritza, 10°57'29”N, 85°29'49"W,  550 m,  19­ 20.vii.1987, Holzenthal, Morse, & Clausen— 1 female( UMSP);  Parque Nacional Rincón de la Vieja, Río Negro, 10°45'54”N, 85°18'47"W,  810 m,  3.iii.1986, Holzenthal& Fasth— 1 male, 3 females( UMSP);   Límon: Limon,  16 kmW Guapiles,  400 m,  ii.­iii.1989, Hanson— 1 male( UMSP);  Reserva Biológica Barbilla, trib. to Río Dantas, 13 (km) S Pacuarito, 09°59'42”N, 83°28'37"W, 0 m,  1.ii.1992, Holzenthal, Muñoz, & Kjer— 1 male, 1 female( UMSP);   Puntarenas: Río Bellavistatrib., Las Alturas, road to quarry, 08°57'07”N, 82°50'53"W,  1480 m,  19.iii.1991, Holzenthal, Muñoz, & Huisman— 1 female( UMSP);  Río Bellavista, ca. 1.5 km NW Las Alturas, 08°57'04”N, 82°50'46"W,  1400 m,  15­17.vi.1986, Holzenthal, Heyn, & Armitage— 1 male( UMSP);   8­9.iv.1987, Holzenthal, Hamilton, & Heyn— 2 males, 2 females( INBIO); —  9 males, 9 females( UMSP);   2­3.viii.1987, Holzenthal, Morse, & Clausen— 2 males, 1 female( UMSP);   10­ 11.viii.1990, Holzenthal, Blahnik, & Muñoz— 1 male, 1 female( UMSP);   16­17.iii.1991, Holzenthal, Muñoz, & Huisman— 7 males( UMSP);  Río Cotón, in Las Alturas, 08°56'17”N, 82°49'34"W,  1360 m,  12.viii.1990, Holzenthal, Blahnik, & Muñoz— 2 females( UMSP);   18.iii.1991, Holzenthal, Muñoz, & Huisman— 1 male( INBIO);  Río Guineal, ca 1 km(air) E Finca Helechales, 09°04'34”N, 83°05'31"W,  840 m,  22.ii.1986, Holzenthal, Morse, & Fasth— 4 males, 2 females( UMSP);   4.viii.1987, Holzenthal, Morse, & Clausen— 1 male( UMSP);  Río Singrí, ca 2 km(air) S Finca Helechales, 09°03'25”N, 83°04'55"W,  720 m,  21.ii.1986, Holzenthal, Morse, & Fasth— 1 male, 1 female( UMSP);  San Vito, 2 kmS., Finca Las Cruces, 08°42'00”N, 83°00'00"W,  4000 m,  20.xi.1988, Solis— 1 male( INBIO);  Jardín BotanicoR & C Wilson, unnamed trib., Sendro del Agua, 08°48'00”N, 82°57'36"W,  1180 m,  8.viii.1990, Holzenthal, Blahnik, & Muñoz— 1 female( UMSP);  Zona Protectora Las Tablas, Río Cotón, Sitio Cotón, 08°56'28”N, 82°47'13"W,  1460 m,  15.iv.1989, Holzenthal& Blahnik— 11 males, 10 females( UMSP);   San José: Motel Prado, San Isidro del General— 1 male( UMSP);  Pedregoso,  640 m, 21.ii., Rounds— 1 male( MCZ);  Río Parrita Chiquito, rt. 12, 6.5 km SW jct. rt. 2, 09°42'11”N, 83°58'12"W,  1990 m,  10.iv.1987, Holzenthal, Hamilton, & Heyn— 1 male( INBIO);   GUATEMALA: Las Mercedes,  914 m, 1879­1881, Champion— 1 male, 1 female( BMNH);  Panima, Vera Paz, 1879­1881, Champion— 1 female( BMNH);   Chiquimula: Padre Miguel,  19.viii.1965, Flint& Ortiz— 2 females( NMNH);   El Progreso: Est. de la Virgen,  11­12.viii.1965, Flint& Ortiz— 1 male, 2 females( NMNH);   Jutiapa: San Jerónimo, 1879­1881, Champion— 2 males, 3 females( BMNH);   Suchitepequez: Cuyotenango,  10­20.vi.1966, Flint& Ortiz— 1 male( NMNH);  MEXICO:  3048 m, 1871, Bilimek— 1 male( NMW);   Chiapas:2.9 km S of Jitotol,  11.viii.1967, Flint— 2 males( NMNH);  58 kmS Palenque,  17.v.1981, C & O Flint— 1 male( NMNH);  Dolores, Rt.190, km 1190,  7.viii.1965, Flint& Ortiz— 1 male( NMNH);  Soconusco, 27 Jul, Purpus— 1 female( ZMHU);   Chihuahua: Arroyo Toro, Toro Basin,  1720 m,  23.vi.1987, Kondratieff& Baumann— 1 male( NMNH);  Riito, Hwy  16, 10 miE of Yepachic,  28.vi.1987, Kondratieff& Baumann— 1 male( NMNH);   Durango:  10 mi.W El Salto,  10.vi.1964, McAlpine— 1 female( NMNH);   Jalisco:11.2 mi. E Cuititian,  4 mi.E Durazno,  1128 m, 29­  30.viii.1987, Bloomfield— 1 female( NMNH); 2.3 mi. E Durazno,  1158 m,  6­8.vi.1989, Bloomfield— 1 male( NMNH);   Mórelos: Cuernavaca, Banks— 1 female( MCZ);   Nayarit:0.6 mi. E Riitos, road to San Andreas,  9­11.iii.1987, Bloomfield— 1 male( NMNH);   Nuevo Leon: Municipio de Santiago, Streamat La Nogaleraabove Cola de Caballo,  27.v.1991, Contreras& Harris— 1 male, 2 females( UMSP);   Oaxaca:  8 kmS Valle Nacional,  25.v.1981, C & O Flint— 1 male, 1 female( NMNH);   San Luis Potosí:  Huichihuayan, Rt.85, km 399,  7.viii.1966, Flint— 1 male, 1 female( NMNH);   Sancti Spíritus: Sótano de Tlamaya,  26.xi.1964, Raines& Bell— 2 males( NMNH);   Sinaloa: Portrerillos,  15 mi.W El Palmito,  1524 m,  8.vii.1964, McAlpine— 1 female( CNC);   20.vii.1964, Mason— 1 female( CNC);   Sonora:small stream near Yecora,  20.viii.1986, Kondratieff— 1 female( NMNH);   Tamaulipas: Rioverge.  25 kmSW Ciudad Victoria,  14­16.iii.1982, Gillaspy— 1 male( NMNH);   Veracruz: Hagen—  P. aeneus holotypefemale ( MCZ); 1.6 mi. N Coscomatepec,  22.vii.1966, Flint& Ortiz— 1 male, 1 female( NMNH);  Cordoba, 4­23.viii.  Aug 1965, Lau— 1 female( NMNH);   10­20.viii.1966, Lau— 1 male( NMNH);   1.ix.1966, Lau— 1 male( NMNH); nr. Huatusco,  25­26.vii.1965, Flint& Ortiz— 1 male( NMNH);  Río Tacolapan,  25­ 26.vii.1966, Flint& Ortiz— 1 male, 1 female( NMNH);  Las Tuxtlas Biological Station, Las Tuxtlas area, nr. Balzapote,  31 kmNE of Catemaco,  3­15.v.1981, C & O Flint— 1 male, 3 females( NMNH);   NICARAGUA: Estelí: Area Protegida Miraflor, Quebrada Grande, 13°13'20”N, 86°15'33"W,  1200 m,  19.vii.2000, Chamorro& Lacayo— 1 female( UMSP);   Jinotega: Peñas Blancas, 13°17'00”N, 85°33'00"W,  1300 m,  25.vii.1997, Maes& Hernández— 5 males( NMNH);   PANAMA: Chiriqui: Boquete,  6.vi.1940, Fairchild— 1 female( MCZ);  Bugaba,  244­457 m,  21.x.1881­  11.iii.1883, Champion— 1 male( BMNH);  Fortuna Dam Sitenr. Hornitos, 08°55'00”N, 82°16'00"W,  1050 m,  24.xi.­ 20.xii.1976, Wolda— 6 males( NMNH);   12­18.x.1977, Wolda— 1 male, 1 female( UMSP);   9­6.xii.1977, Wolda— 1 male, 2 females( NMNH);   16­29.xi.1977, Wolda— 2 males, 1 female( NMNH);   30­6.xii.1977, Wolda— 3 males( NMNH);   UNITED STATES: Texas: Brownwood, Banks—  P. ornatus holotypefemale ( MCZ);  Fort Davis, Davis Mountains,  1524 m, vii­viii.1928, Poling— 1 female( CAS);  Brewster County, Windows Creek,  5­10.iv.1993, Gelhaus— 1 female( NMNH);  Jeff Davis County, Barrilla Draw,  4 mi.E Ft. Davis,  11­12.iv.1993, C & O Flint— 1 male( NMNH);  Kerr County, Kerrville,  1.iv.1959, Wigmore— 1 female( CNC). Distribution. Belize, Costa Rica, Guatemala, Mexico, Nicaragua, Panama, USA( Texas). 3319300662 1975-06-18 UMSP Resh Belize Material Rio Frio Cave Rio Frio 22 23 3 1 2 Caya 3319300446 MNHNP Costa Rica Serre 22 23 1 holotype 3319300850 1990-11-01 1990-11-20 1990-11-01 EMUS Parker Costa Rica 20 km S Upala 22 23 1 1 Alajuela holotype 3319300889 1990-09-03 1990-09-04 1990-09-03 NMNH Holzenthal & Blahnik & Huisman Costa Rica 1530 10.203889 Bajos del Toro 21 -84.31612 Rio Toro 22 23 3 2 1 Alajuela holotype 3319300739 1986-10-01 1986-10-04 1986-10-01 UMSP I & A Chacon Costa Rica 980 10.216111 Rio San Lorencito 21 -84.60694 Reserva Forestal San Ramon 23 24 1 1 Alajuela holotype 3319300997 1987-03-30 1987-04-01 1987-03-30 UMSP Holzenthal & Hamilton & Heyn Costa Rica 23 24 1 1 Alajuela holotype 3319300424 [608,1270,364,390] 1990-05-01 1990-05-04 1990-05-01 UMSP Holzenthal & Blahnik Costa Rica 23 24 1 1 Alajuela holotype 3319300714 1990-07-28 1990-07-30 1990-07-28 UMSP Holzenthal & Blahnik & Munoz Costa Rica 23 24 1 1 Alajuela holotype 3319300565 1991-02-06 1991-02-10 1991-02-06 UMSP Holzenthal & Munoz & Huisman Costa Rica 23 24 3 1 2 Alajuela holotype 3319300477 1991-03-24 1991-03-25 1991-03-24 UMSP Holzenthal & Munoz & Huisman Costa Rica 1400 9.719999 Quebrada Palmitos 21 -83.78001 Reserva Tapanti 23 24 1 1 Cartago holotype 3319300410 1985-01-23 NMNH Janzen & Hallwachs Costa Rica 1650 9.686111 Rio Grande de Orosi 21 -83.75611 23 24 1 1 Cartago holotype 3319300934 1986-05-31 UMSP Janzen & Hallwachs Costa Rica 310 El 23 24 2 1 1 Guanacaste holotype 3319300673 1991-11-17 EMUS R. Naranjo & Parker Costa Rica Finca Montezuma 23 24 1 1 Guanacaste holotype 3319301002 [381,878,684,710] 1992-01-20 1992-01-31 1992-01-20 EMUS Parker Costa Rica 23 24 1 1 Guanacaste holotype 3319301020 [892,1317,684,710] 1992-05 EMUS Parker Costa Rica 23 24 1 1 Guanacaste holotype 3319301083 [264,784,724,750] 1993-05-16 1993-05-31 1993-05-16 EMUS Parker Costa Rica 23 24 1 1 Guanacaste holotype 3319300576 1994-05-19 1994-05-31 1994-05-19 UMSP Costa Rica 580 10.956944 Rio Tempisquito 21 -85.494995 Maritza 23 24 2 1 1 Guanacaste holotype 3319300369 [383,1155,804,830] 1994-02-14 1994-05-01 1994-02-14 UMSP Costa Rica 23 24 4 2 2 Guanacaste holotype 3319300558 1994-01-31 1994-05-27 1994-01-31 UMSP Costa Rica 675 10.991112 Rio Orosi 21 -85.42833 Maritza 23 24 3 1 2 Guanacaste holotype 3319300921 1986-03-08 UMSP Holzenthal & Fasth Costa Rica Quebrada Garcia 470 10.861945 Quebrada Grande 21 -85.428055 Quebrada Garcia 23 24 1 1 Guanacaste holotype 3319300930 1986-03-07 UMSP Holzenthal & Fasth Costa Rica Rio Los Ahogados 470 10.865001 Quebrada Grande 21 -85.42305 Rio Los Ahogados 23 24 1 1 Guanacaste holotype 3319301061 1992-01-29 NMNH Sweeney Costa Rica Parque Nacional Guanacaste 550 10.9580555 Rio Tempisquito 21 85.49694 Estacion Maritza 23 24 1 1 Guanacaste holotype 3319300689 1990-05-22 1990-05-25 1990-05-22 UMSP Holzenthal & Blahnik Costa Rica 700 10.991112 Estacion Pitilla 21 -85.428055 Rio Orosi 23 24 1 1 Guanacaste holotype 3319301053 1987-04-03 1987-04-04 1987-04-03 INBIO Holzenthal & Hamilton & Heyn Costa Rica 960 10.921945 Rio San Josecito 21 -85.47 23 24 1 1 Guanacaste holotype 3319300658 1987-07-19 1987-07-20 1987-07-19 UMSP Holzenthal & Morse & Clausen Costa Rica 550 10.9580555 Maritza 21 -85.49694 Rio Tempisquito 23 24 1 1 Guanacaste holotype 3319300778 1986-03-03 UMSP Holzenthal & Fasth Costa Rica 810 10.765 Rio Negro 21 -85.31306 Parque Nacional Rincon de la Vieja 23 24 4 3 1 Guanacaste holotype 3319300669 1989-02 1989-03-31 1989-02 UMSP Hanson Costa Rica 400 Limon 23 24 1 1 Limon holotype 3319300385 1992-02-01 UMSP Holzenthal & Munoz & Kjer Costa Rica Reserva Biologica Barbilla 9.995 Pacuarito 21 -83.476944 Rio Dantas 23 24 2 1 1 Limon holotype 3319300341 1991-03-19 UMSP Holzenthal & Munoz & Huisman Costa Rica 1480 8.951944 Las Alturas 21 -82.84806 Rio Bellavista 23 24 1 1 Puntarenas holotype 3319300308 1986-06-15 1986-06-17 1986-06-15 UMSP Holzenthal & Heyn & Armitage Costa Rica 1400 8.951111 Las Alturas 21 -82.846115 Rio Bellavista 23 24 1 1 Puntarenas holotype 3319301094 1987-04-08 1987-04-09 1987-04-08 INBIO Holzenthal & Hamilton & Heyn Costa Rica 23 24 4 2 2 Puntarenas holotype 3319301107 1987-04-08 1987-04-09 1987-04-08 UMSP Holzenthal & Hamilton & Heyn Costa Rica true 23 24 18 9 9 Puntarenas holotype 3319300830 [379,1270,1644,1670] 1987-08-02 1987-08-03 1987-08-02 UMSP Holzenthal & Morse & Clausen Costa Rica 23 24 3 1 2 Puntarenas holotype 3319300375 1990-08-10 1990-08-11 1990-08-10 UMSP Holzenthal & Blahnik & Munoz Costa Rica 23 24 2 1 1 Puntarenas holotype 3319301069 1991-03-16 1991-03-17 1991-03-16 UMSP Holzenthal & Munoz & Huisman Costa Rica 23 24 7 7 Puntarenas holotype 3319300927 1990-08-12 UMSP Holzenthal & Blahnik & Munoz Costa Rica 1360 8.938056 Las Alturas 21 -82.82611 Rio Coton 23 24 2 2 Puntarenas holotype 3319300550 [485,1264,1804,1830] 1991-03-18 INBIO Holzenthal & Munoz & Huisman Costa Rica 23 24 1 1 Puntarenas holotype 3319300661 1986-02-22 UMSP Holzenthal & Morse & Fasth Costa Rica 840 24 25 9.076111 Finca Helechales 21 -83.09195 Rio Guineal 23 24 6 2 4 Puntarenas holotype 3319301077 1987-08-04 UMSP Holzenthal & Morse & Clausen Costa Rica 24 25 1 1 Puntarenas holotype 3319301090 1986-02-21 UMSP Holzenthal & Morse & Fasth Costa Rica 720 9.056945 Finca Helechales 21 -83.08194 Rio Singri 24 25 2 1 1 Puntarenas holotype 3319300604 1988-11-20 INBIO Solis Costa Rica 4000 8.7 Finca Las Cruces 21 -83.0 San Vito 24 25 1 1 Puntarenas holotype 3319300620 1990-08-08 UMSP Holzenthal & Blahnik & Munoz Costa Rica Jardin Botanico 1180 8.8 Sendro del Agua 21 -82.96 Wilson 24 25 1 1 Puntarenas holotype 3319300491 1989-04-15 UMSP Holzenthal & Blahnik Costa Rica Zona Protectora Las Tablas 1460 8.941112 Sitio Coton 21 -82.78694 Rio Coton 24 25 21 10 11 Puntarenas holotype 3319300539 [487,1318,603,630] UMSP Costa Rica San Isidro del General Motel Prado 24 25 1 1 San Jose holotype 3319300370 [264,884,644,670] MCZ Rounds Costa Rica 640 Pedregoso 24 25 1 1 San Jose holotype 3319300722 1987-04-10 INBIO Holzenthal & Hamilton & Heyn Costa Rica 1990 9.703055 Rio Parrita Chiquito 21 -83.97 24 25 1 1 San Jose holotype 3319300692 BMNH Champion Guatemala 914 Las Mercedes 24 25 2 1 1 holotype 3319301027 BMNH Champion Guatemala Vera Paz Panima 24 25 1 1 holotype 3319300877 [388,1317,804,830] 1965-08-19 NMNH Flint & Ortiz Guatemala Padre Miguel 24 25 2 2 Chiquimula holotype 3319300723 1965-08-11 1965-08-12 1965-08-11 NMNH Flint & Ortiz Guatemala Est. de la Virgen 24 25 3 2 1 El Progreso holotype 3319300744 [389,1318,884,910] BMNH Champion Guatemala San Jeronimo 24 25 5 3 2 Jutiapa holotype 3319300340 [264,1229,924,950] 1966-06-10 1966-06-20 1966-06-10 NMNH Flint & Ortiz Guatemala Cuyotenango 24 25 1 1 Suchitepequez holotype 3319300578 NMW Bilimek Guatemala 3048 24 25 1 1 Suchitepequez holotype 3319300884 1967-08-11 NMNH Flint Guatemala S of Jitotol 24 25 2 2 Chiapas holotype 3319301013 1981-05-17 NMNH C & O Flint Guatemala Palenque 24 25 1 1 Chiapas holotype 3319300488 [454,1318,1044,1070] 1965-08-07 NMNH Flint & Ortiz Guatemala Rt. Dolores 24 25 1 1 Chiapas holotype 3319300941 [264,845,1084,1110] ZMHU Purpus Guatemala Jul Soconusco 24 25 1 1 Chiapas holotype 3319301035 1987-06-23 NMNH Kondratieff & Baumann Guatemala 1720 Toro Basin Arroyo Toro 24 25 1 1 Chihuahua holotype 3319300650 1987-06-28 NMNH Kondratieff & Baumann Guatemala Hwy Riito 24 25 1 1 Chihuahua holotype 3319300812 1964-06-10 NMNH McAlpine Guatemala 10 mi. W El Salto 24 25 1 1 Durango holotype 3319300753 1987-08-null NMNH E Cuititian & Bloomfield Guatemala 1128 4 mi. E Durazno 24 25 1 1 Jalisco holotype 3319300953 1989-06-06 1989-06-08 1989-06-06 NMNH E Durazno & Bloomfield Guatemala 1158 24 25 1 1 Jalisco holotype 3319300705 MCZ Banks Guatemala Cuernavaca 24 25 1 1 Morelos holotype 3319300916 1987-03-09 1987-03-11 1987-03-09 NMNH Bloomfield Guatemala San Andreas Riitos 24 25 1 1 Nayarit holotype 3319300441 1991-05-27 UMSP Contreras & Harris Guatemala Municipio de Santiago Cola de Caballo Stream 24 25 3 2 1 Nuevo Leon holotype 3319300880 1981-05-25 NMNH C & O Flint Guatemala 8 km S Valle Nacional 24 25 2 1 1 Oaxaca holotype 3319300933 1966-08-07 NMNH Flint Guatemala Huichihuayan, Rt. Rt. Huichihuayan 24 25 2 1 1 San Luis Potosi holotype 3319300469 1964-11-26 NMNH Raines & Bell Guatemala Sotano de Tlamaya 24 25 2 2 Sancti Spiritus holotype 3319300757 1964-07-08 CNC McAlpine Guatemala 1524 Portrerillos 24 25 1 1 Sinaloa holotype 3319301047 [649,1131,1604,1630] 1964-07-20 CNC Mason Guatemala 24 25 1 1 Sinaloa holotype 3319301006 1986-08-20 NMNH Kondratieff Guatemala Yecora 24 25 1 1 Sonora holotype 3319300515 1982-03-14 1982-03-16 1982-03-14 NMNH Gillaspy Guatemala Rio 24 25 1 1 Tamaulipas holotype 3319300637 MCZ Guatemala Hagen 24 25 1 Veracruz holotype 3319300456 1966-07-22 NMNH Flint & Ortiz Guatemala Coscomatepec 24 25 2 1 1 Veracruz holotype 3319300530 1965-08 NMNH Lau Guatemala Cordoba 24 25 1 1 Veracruz holotype 3319300442 [391,882,1804,1830] 1966-08-10 1966-08-20 1966-08-10 NMNH Lau Guatemala 24 25 1 1 Veracruz holotype 3319301081 [897,1318,1804,1830] 1966-09-01 NMNH Lau Guatemala 24 25 1 1 Veracruz holotype 3319300726 [264,1067,1844,1870] 1965-07-25 1965-07-26 1965-07-25 NMNH Flint & Ortiz Guatemala Huatusco 24 25 1 1 Veracruz holotype 3319300766 1966-07-25 1966-07-26 1966-07-25 NMNH Flint & Ortiz Guatemala 25 26 Rio Tacolapan 24 25 2 1 1 Veracruz holotype 3319300795 1981-05-03 1981-05-15 1981-05-03 NMNH C & O Flint Guatemala Las Tuxtlas Biological Station Balzapote Las Tuxtlas area 25 26 4 3 1 Veracruz holotype 3319300596 2000-07-19 UMSP Chamorro & Lacayo Nicaragua 1200 13.222221 Quebrada Grande 21 -86.25916 Area Protegida Miraflor 25 26 1 1 Esteli holotype 3319300668 1997-07-25 NMNH Maes & Hernandez Nicaragua 1300 13.283334 Penas Blancas 21 -85.55 25 26 5 5 Jinotega holotype 3319300986 1940-06-06 MCZ Fairchild Panama Boquete 25 26 1 1 Chiriqui holotype 3319300695 1881-10-21 1883-03-11 1881-10-21 BMNH Champion Panama 351 Bugaba 25 26 1 1 Chiriqui holotype 3319300748 1976-11-24 1976-12-20 1976-11-24 NMNH Wolda Panama 1050 8.916667 Hornitos 21 -82.26667 Fortuna Dam Site 25 26 6 6 Chiriqui holotype 3319300945 1977-10-12 1977-10-18 1977-10-12 UMSP Wolda Panama 25 26 2 1 1 Chiriqui holotype 3319300626 [381,994,644,670] 1977-12-06 1977-12-09 1977-12-06 NMNH Wolda Panama 25 26 3 2 1 Chiriqui holotype 3319300561 1977-11-16 1977-11-29 1977-11-16 NMNH Wolda Panama 25 26 3 1 2 Chiriqui holotype 3319300542 [578,1073,684,710] 1977-12-06 1977-12-30 1977-12-06 NMNH Wolda Panama 25 26 3 3 Chiriqui holotype 3319300791 MCZ United States of America Banks Brownwood 25 26 1 Texas holotype 3319301088 CAS Poling United States of America 1524 Davis Mountains Fort Davis 25 26 1 1 Texas holotype 3319300486 1993-04-05 1993-04-10 1993-04-05 NMNH Gelhaus United States of America Brewster County Windows Creek 25 26 1 1 Texas holotype 3319301021 1993-04-11 1993-04-12 1993-04-11 NMNH C & O Flint United States of America Jeff Davis County Barrilla Draw 25 26 1 1 Texas holotype 3319300448 1959-04-01 CNC Wigmore United States of America Kerr County Kerrville 25 26 1 1 Texas holotype