Systematic studies on Pseudomyrmex acacia-ants (Hymenoptera: Formicidae: Pseudomyrmecinae) Author Ward, Philip S. text Journal of Hymenoptera Research 1993 2 117 168 journal article 10.5281/zenodo.10150 2958 80669360-1DB2-4B74-9F4B-99DD3E7EBBFA Pseudomyrmex ferrugineus (F. Smith 1877 ) (Figs. 18, 27,65,66, 70) Pseudomyrma ferruginea F. Smith 1877:64 . Lectotype worker , Mexico ( BMNH ) [ Examined ] , Pseudomyrma belti racefulvescens Emery 1890:64 . Lectotype worker , Guatemala ( Beccari ) ( MCSN ) [Examined] [Synonymy by Ward 1989:437 ; see also Janzen 1967b:391 ] . Pseudomyrma canescens Wasmann 1915: 321 . Syntype workers , T ampico, Mexico ( Brakhoven ) ( MCSN , MCZC ) [Examined] [Synonymy by Ward 1989: 437 ] , Pseudomyrma wasmanni Wheeler 1921:92 . Replacement name, now unnecessary, for P. canescens Wasmann 1915 (nec F. Smith 1877 ). Pseudomyrma belti subsp. bequaerti Wheeler 1942:164 . Lectotype worker , Puerto Castilla , Honduras ( J. Bequaert ) ( MCZC ) [Examined] [Synonymy by Ward 1989:437 ] . Pseudomyrma belti subsp. saffordi Wheeler 1942:162 . Lectotype worker , Chicoasen , Chiapas , Mexico ( G. N. Collins ) ( MCZC ) [Examined] [Synonymy by Ward 1989:437 ] , Pseudomyrma belti subsp. vesana Wheeler 1942:163 . Holotype (unique syntype) worker , Cordoba , Mexico ( F. Knab ) ( MCZC ) [Examined] [Synonymy by Ward 1989:437 ] , Pseudomyrma belti subsp. bequaerti Enzmann 1945:80 . Syntype workers , Puerto Castilla , Honduras ( J. Bequaert ) ( MCZC ) [Examined] [Objective synonym of P. belti bequaerti Wheeler ; Brown 1949:42 ] , Pseudomyrma kuenckeli var. hondurana Enzmann 1945:87 . Lectotype worker , Honduras ( Bates ) ( MCZC ) [Examined] [Synonymy by Ward 1989:437 ] , Pseudomyrmex ferruginea [sic] (F. Smith); Janzen 1966:252 . Pseudomyrmex ferrugineus (F. Smith) ; Kempf Table 1. Summary of nomenclature used in the biological literature on acacia-ants, Pseudomyrmex ferrugineus group.* For all except some of the most recent ( 1979-1990 ) papers, the relationship between the names used in the paper and the species recognized in the current study was established by direct examination of museum specimens collected, or referred to, by the author(s). Only specific and infraspecfic epithets are listed since all species were placed in the same genus, PseudorrryrmaGuerin, until 1952 when M. R. Smith showed that a different generic name, Pseudomyrmex Lund , had priority. By the rules of zoological nomenclature this necessitates a change in the ending of adjectival species names (e.g. ferruginea to ferrugineus ) to agree in gender with Pseudomyrmex (masc.) but this convention was not followed by all authors. Species
Author (s) ferrugineus flavicornis nigrocinctus peperi spinicola veneficus
Norton (1868a) flavidula (part) _ _ flavidula (part) _ _
Belt (1874) - bicolor - - - -
Emery (1890,1891) belti fulvescens belti nigrocincta - spinicola -
Wheeler (1913) belti fulvescens (part) spinicola (part) belti (part) nigrocincta - spinicola (part) belti fulvescens (part) belti (part)
Wasmann (1915,1916) canescens belti nigrocincta - spinicola -
Safford (1922) belti fulvescens (part) belti nigrocincta belti "varieties" (part) spinicola belti fulvescens (part) -
Menozzi (1927a ,b) - belti obnubila peltata - - -
Skwarra (1934a ,b) belti uiasmanni (part) - - belti uiasmanni (part) - -
Wheeler (1942) belti fulvescens belti saffordi (part) belti vesana belti bequaerti belti uiasmanni belti belti fellosa belti obnubila nigrocincta peltata spinicola convarians belti saffordi (part) spinicola spinicola atrox spinicola gaigei spinicola infernalis spinicola scelerosa belti venefica (part)
Janzen (1966,1967a) ferruginea spinicola (part) belti (part) nigrocincta (part) nigrocincta (part) spinicola (part) belti (part)
Janzen (1967b , 1969 , 1973 , 1974 , 1975 , 1983 ) ferruginea (part) belti (part) nigrocincta (part) nigrocincta (part) and "undescribed sp." ferruginea (part) venefica
Hoelldobler & Engel (1979) , Hoelldobler & Engel-Siegel (1985) - - - - ferruginea -
Mintzer (1982) , Mintzer & Vinson (1985) , Mintzer et al. (1987) ferruginea - - - - -
Errard (1984) ferruginea - - - - -
Espelie & Hermann (1988) ferrugineus - - - - -
Ward (1989) ferrugineus flavicornis nigrocinctus peperi spinicola veneficus
Young et al. (1990) - flavicornis nigrocinctus - spinicola -
*Of the four other species in the P. ferrugineus group, P. satanicus has always been correctly identified ( Wheeler, 1942 ; Janzen, 1969 , 1973 , 1974 ); P. mixtecus was mistaken for P. veneficus by Wheeler (1942) and for P. flavicornis (= belti ) by Janzen (1966,1973); P. janzeni was referred to by Janzen (1973:729) as an undescribed species derived from P. ferrugineus-, and P. particeps has received no mention in the literature. Worker measurements (n = 69). - HL 0.99-1.33, HW 0.92-1.15, MFC 0.054-0.108, Cl 0.81-0.94, REL 0.42-0.48, REL2 0.48-0.54, 001 1.39-3.16, VI 0.60-0.78, FCI 0.054-0.101, SI 0.41-0.46, SI2 0.79-0.91, NI 0.61-0.72, PLI 0.54-0.69, PWI 0.56- 0.73, PPWI 1.34-1.70.
Worker diagnosis. - Medium-sized species (HW>0.91; LHT 0.75-1.06) with broad head (Cl> 0.80); anterior margin of median clypeal lobe straight or weakly concave, rounded laterally; palp formula 5,3; frontal carinae well separated (FCI> 0.05) and median lobe of antennal sclerite not strongly exposed (FCI 2 0.45-0.69); mesosomal profile typically as in Fig. 27, with mesonotum notably inclined and with basal face of propodeum rounding gradually into declivitous face, but deviations from this pattern occur; petiole relatively short, high and wide (see relevant metrics: PLI, PWI), with a distinct anterior peduncle (PWI 3 0.36-0.50); posterolateral angles of petiole moderately developed but not as pronounced as in P. peperi (compare Figs. 24 and 27); postpetiole broad. Head densely punctulate, predominantly opaque or subopaque but at least weakly shining on upper third of head between ocelli and compound eye; mesosoma punctulate to punctulate-coriarious, subopaque to sublucid; posterior portions of propodeum opaque to subopaque and usually overlain by larger but weak, irregular punctures or rugulae. Petiole, postpetiole and gaster with fine piligerous punctures, sublucid. Standing pilosity common; pubescence dense but fine and appressed on most surfaces. Color variable, from light reddish- or yellowish-brown to very dark brown, gaster (and usually head) somewhat darker than the mesosoma; mandibles, scapes, fronto-clypeal complex, and apices of legs usually lighter. Comments. - Workers and queens of P. ferrugineus can be recognized by features of head morphology (laterally rounded median clypeal lobe, well separated frontal carinae and correspondingly limited exposure of the median lobes of the antennal sclerites, and moderately broad worker head; see Fig. 18), head sculpture (densely punctulate and (sub)opaque, but weakly shining on upper third of head between the ocelli and compound eye), and coloration (variably brown, not black or orange-brown). This species is most likely to be confused with the allopatric P. janzeni and the partly sympatric P.flavicornis . See under those species for more specific discussion. Distribution and biology. - P. ferrugineus is distributed from eastern and southern Mexico to El Salvador and Honduras (Fig. 70). It is a common species and colonies have been recorded from all swollen-thorn acacia species growing within its range, i.e. Acacia chiapensis , A. collinsii , A. cookii , A. cornigera , A. gentlei , A. globulifera , A. hindsii , A. janzenii , A. mayana and A. sphaerocephala . P. ferrugineus is usually monogynous ( Janzen 1967b , 1973 ) but a few alcohol nest series from Guatemala, belonging to apparently mature colonies (as judged by the presence of alates), contained more than one dealate queen. Janzen (1966 , 1967b ) conducted a massive experimental study of the interaction between P. ferrugineus and Acacia cornigera in Mexico, and the results of these experiments, together with copious additional observations by Janzen, showed conclusively that the ants protect the acacia from herbivores and competing plants (especially vines). P. ferrugineus was also the subject of a study on kinship and nestmate recognition among workers ( Mintzer 1982 ; Mintzer et al. 1985 ) which demonstrated a worker-based and probably inherited component to colony odor. Statements about the biology of " P. ferruginea " from regions south of El Salvador and Honduras (e.g. Janzen 1983 ) refer to a different species, P. spinicola . Material examined ( AMNH , BMNH , CASC , CHAH , CISC , CUIC , INHS , LACM , MCSN , MCZC , MHNG , MNHN , MZSP , NHMV , PSWC , SEMC , UCDC , USNM , WPMC ) . - BELIZE Belize : 2.2mi W Belize, rd.to Chetumal ( D.H.Janzen ) ; 3.8mi NW Belize, rd.to Chetumal ( D.H.Janzen ) ; 4.9mi SW Belize, rd.to Cayo ( D.H.Janzen ) ; Belize ( Baker ; c.u.) ; Manatee [River] ( J.D.Johnson ; c.u.) ; nr. Belize ( N.L.H.Krauss ) ; Cayo : 36.1mi SW Belize ( D.H.Janzen ) ; 36.1mi [S]W Belize ( U.Kansas Mex.Exped .) ; El Cayo [=San Ignacio] ( N.L.H.Krauss ) ; Pine Mtn. Ridge, Mecal R. , 415m ( G.D.Alpert ) ; San Ignacio ( S.E.Schoenig ) ; Corozal : 13.5mi S Sta.Elena (Louisville) ( D.H.Janzen ) ; 15mi S Sta.Elena (Louisville) ( D.H.Janzen ) ; Orange Walk'. 5mi SE Orange Walk , rd.to Belize ( D.HJanzen ) ; Toledo : Punta Gorda ( H.Broomfield ) . EL SALVADOR Chalatenango : 4.7mi NW La Palma , 880m ( D.H.Janzen ) ; 5.5mi SE La Palma , 1130m ( D.HJanzen ) . GUATEMALA Alta Verapaz : San Joaquin, nr. San Cristobal Verapaz , 1080m ( D.H.Janzen ) ; Trece Aguas ( Schwarz & Barber ) ; Chimaltenango : Yepocapa ( H.T.Dalmat ) ; Coatepeque : 2mi W Coatepeque ( D.HJanzen ) ; El Progreso : 4.9mi SW Sanarate , 780m ( D.H.Janzen ) ; Escuintla : 1.7mi S Escuintla , 370m [on CA-2] ( D.H.Janzen ) ; 12.6mi NE Escuintla, rd. to Cd. Guatemala , 1120m ( D.HJanzen ) ; 15km E Escuintla [on CA-9] ( D.H.Janzen ) ; 3mi N Escuintla [on CA-9] ( D.H.Janzen ) ; 3mi S Escuintla [on CA-9] ( D.H.Janzen ) ; 43km S Cd. Guatemala [= 15km E Escuintla] ( D.H.Janzen ) ; 6.8mi S Escuintla , 280m [on CA-2] ( D.H.Janzen ) ; 8.3mi N Escuintla [on CA-9] ( D.HJanzen ) ; 9.2mi N Escuintla , 900m [on CA-9] ( D.H.Janzen ) ; Escuintla ( W.M.Wheeler ) ; Pantaleon ( Champion ) ; Guatemala : 16.2mi NE Cd.Guatemala ( D.H.Janzen ) ; 19kmS Cd.Guatemala [on CA-9] ( D.H.Janzen ) ; 20mi SE Cd.Guatemala , 1060m [on CA-1] ( D.H.Janzen ) ; 6.2mi NE Cd.Guatemala , 700m ( D.H.Janzen ) ; 7.9mi S Cd. Guatemala , 1360m [on CA-9] ( D.H.Janzen ) ; 8.1mi NE Cd. Guatemala , 850m ( D.H.Janzen ) ; Cd. Guatemala ( Champion ) ; Cd. Guatemala , 980m ( D.H.Janzen ) ; Escuintla-Cd.Guatemala [= 19km S Cd. Guatemala] ( D.H.Janzen ) ; Huehuetenango : 12.4mi SE Mex.border at Cd. Cuauhtemoc ( D.H.Janzen ) ; 12.5mi SE Mex.border at Cd. Cuauhtemoc ( D.H.Janzen ) ; 37.6mi NW Huehuetenango , 900m ( D.HJanzen ) ; 7.1mi SE Mex.border at Cd. Cuauhtemoc ( D.H.Janzen ) ; 9.3mi SE Mex.border at Cd. Cuauhtemoc ( D.H.Janzen ) ; Izabal : 1.1mi NE Quirigua , 120m ( D.HJanzen ) ; 2. lmi SW Morales , 50m ( D.HJanzen ) ; Lago Izabal, lkm NE El Estor [= 1.5km NE El Estor] ( D.H.Janzen ) ; Los Amates ( Kellerman ) ; Murciellago ( D.H.Janzen ) ; Quirigua ( W.M.Wheeler ) ; Jutiapa : 6.7mi N San Cristobal , 280m ( D.H.Janzen ) ; 6.9mi N San Cristobal , 290m ( D.H.Janzen ) ; 9mi SW Jutiapa , 950m ( D.H.Janzen ) ; Peten : 70km NW Tikal ( W.R.Tschinkel ) ; Tikal ( T.H.Hubbell ; N.L.H.Krauss ; W.R.Tschinkel ) ; Quezaltenango : 7.2mi NE San Felipe, on Retalhuleu-QuezaltenangoRd. ( D.H.Janzen ) ; 7.5mi NE San Felipe, on Retalhuleu-Quezaltenango Rd. ( D.H.Janzen ) ; Retalhuleu : 1.3mi E Champerico ( D.H.Janzen ) ; 2mi NE Champerico ( D.H.Janzen ) ; 5mi W Retalhuleu ( D.HJanzen ) ; 5mi W Retalhuleu, Hwy. CA-2, at Rio Nil ( D.HJanzen ) ; Champerico ( Baker ; F.Knab ) ; Puenta Samala, 3.8mi NE San Felipe ( D.HJanzen ) ; Retalhuleu ( Stoll ) ; SantaRosa : 25mi SE Escuintla , 200m [on CA-2] ( D.H.Janzen ) ; Solola : " Pacific slope ", 3000ft . ( c.u. ) ; Suchitepequez : Patulul ( W.M.Wheeler ) ; Zacapa : lOmi SW El Lobo , 170m [on CA-9] [= 9.2mi NE Piedras de Afilar] ( D.HJanzen ) ; 2.6mi SW El Lobo , 100m [on CA-9] [= 16.6mi NE Piedras de Afilar] ( D.H.Janzen ) ; 8.1mi SWLos Amates , 160m [on CA-9] [=8.0mi NE El Lobo] ( D.HJanzen ) ; 8.2mi NE Piedras de Afilar , 160m [on CA-9] [= 12.2mi NE Rio Hondo] ( D.H.Janzen ) ; dept, unknown-. " Guatemala " ( Beccari ) ; Concepcion , 1400ft . ( C.N.Ainslie ) ; Mimosa , "Cocepcion" [Concepcion] ( C.N.Ainslie ) . HONDURAS Atldntida : Tela (quarantine New Orleans, U.S.A.) ( TJ.Baker ) ; Colon : Puerto Castilla ( J.Bequaert ) ; Comayagua : " Comayaena " ( S. Passoa ) ; 11.7miS San Antonio , 830m ( D.H.Janzen ) ; 15.7mi N Siguatepeque , 530m ( D.H.Janzen ) ; Minas de Oro ( J.B.Edwards ) ; Copan : 10.4mi S Sta. Rosa de Copan , 980m ( D.H.Janzen ) ; 11.9mi S Sta. Rosa de Copan , 1130m ( D.HJanzen ) ; 17.3mi N Sta. Rosa de Copan , 780m ( D.H.Janzen ) ; Cortes'. 24.6mi SW San Pedro Sula , 240m ( D.H.Janzen ) ; 6.8mi S San Pedro Sula , 480m [= 20.9mi SW Quimistan ] ( D.HJanzen ) ; San Pedro Sula ( S.C.Bruner ; W.M.Mann ) ; Ocotepeque : 4.4mi E [Nueva] Ocotepeque , 1310m ( D.H.Janzen ) ; Santa Bdrbara : 7.3mi E Quimistan ( D.H.Janzen ) ; 13.7mi SW Quimistan , 320m ( D.H.Janzen ) ; Yoro : Coyoles, W of Olanchito ( Echtemacht ) ; dept.unknown : " Honduras " ( Bates ) . MEXICO Camp. : 0.1 mi S Tenabo ( D.HJanzen ) ; 0.8mi E Campeche ( D.H.Janzen ) ; 10.6mi E Campeche ( D.HJanzen ) ; 20mi E Campeche, Hwy. 180 ( D.H.Janzen ) ; 29mi E & 12mi S Campeche (Ruinas Edzna) ( D.H. Janzen ) ; 48mi NE Puerto Real (Isla Aguada), Hwy. 180 ( D.H.Janzen ) ; Ruinas Edzna ( U.Kansas Mex.Exped .) ; Chis. : 10 mi NE [NW?] Tapachula [on Hwy. 200?] ( D.H.Janzen ) ; 10.8 mi S Pichucalco ( D.H.Janzen ) ; 11km SE Pichucalco , 400m ( P.S.Ward ) ; llmi E Arriaga ( D.H.Janzen ) ; 1km WSW Palenque , 80m ( P.S.Ward ) ; 26km E Cintalpa ( W.MacKay ) ; 2mi S Pichucalco ( D.H.Janzen ) ; 32mi W [San] Cristobal de las Casas, Hwy.190 ( D.H.Janzen ) ; 3km ENE Chiapa de Corzo , 500m ( P.S.Ward ) ; 3mi N Soyalo [on Hwy.195] ( D.H.Janzen ) ; 4mi NW Ocosingo ( R.C.Bechtel & E.I.Schlinger ; R.F.Smith ) ; 4mi S Simojovel ( R.C. Bechtel & E.I.Schlinger ; E.I.Schlinger ) ; 56.9mi NE [NW?] Tapachula [on Hwy.200?] ( D.H.Janzen ) ; 5mi SE Tapilula [on Hwy.195] ( D.H.Janzen ) ; 6.9mi N Tapilula ( D.H.Janzen ) ; 7mi SW Teapa [on Hwy.195] ( D.H.Janzen ) ; 8.5mi S La Trinitaria , 1160m ( U.Kansas Mex.Exped .) ; 8.5mi S La Trinitaria, Hwy.190 ( D.H.Janzen ) ; 8mi W Las Cruces, Hwy . 190, 660m ( D.H.Janzen ) ; 96km S Tuxtla Gutierrez , 732m ( D.E. & J.A.Breedlove ) ; Acapetahua Las Bugamvillas ( F.Diaz ) ; Arriaga ( D.H.Janzen ) ; Cd. Cuauhtemoc ( D.H.Janzen ) ; Chicoasen ( G.N. Collins ) ; El Real ( Goodnight & Stannard ) ; Ixtacomitan ( R.Andrews ) ; Llano Grande ( G.N. Collins ) ; Palenque ( c.u. ) ; Pichucalco ( G.N. Collins ) ; Rio Huixtla, Huixtla ( J.G.Pereira ) ; Ruinas Palenque ( E.M.Fisher ) ; San Sebastian [near Tuxtla Gutierrez ; see Safford 1922:393 ] ( G.N. Collins ) ; Santo Domingo, 15mi SE [SW?] Simojovel ( R.F.Smith ) ; Tapilula ( D.H.Janzen ) ; Tonala , 40m ( D.H.Janzen ) ; Tonola [prob.=Tonala] ( A.Petrunkewitch ) ; Tuxtla Gutierrez ( N.L.H.Krauss ; W.P.Stoutamize ) ; Yaxoquintela ( J.E.Rawlins ) ; Yerba Santa ( G.N. Collins ) ; Gro. : 10mi NE Acapulco ( D.H.Janzen ) ; 15.8mi S Chilpancingo ( D.H.Janzen ) ; 18mi S Chilpancingo ( F.D.Parker ; F.D.Parker & L.A.Stange ) ; 25.4mi S Chilpancingo ( D.H.Janzen ) ; 55mi N Acapulco, Hwy.95 ( Cornell Univ.Mex.Field Party ; c.u. ) ; 57.8mi S Chilpancingo ( D.H.Janzen ) ; 59.9mi N Acapulco ( D.H.Janzen ) ; 62.4mi N Acapulco ( D.H.Janzen ) ; 74km N Acapulco ( W.MacKay ) ; Acapulco ( L.J.Lipovsky ) ; Revolcadero ( N.L.H.Krauss ) ; San Geronimo de Juarez ( W.von Hagen ) ; Hgo. : 2km W Orizatlan , 245m ( W.MacKay ) ; Mor. : 15miS Cuernavaca ( E.S.Ross ; W.S.Ross ) ; Cuernavaca ( W.M.Wheeler ) ; Oax. : 11.4-17.0mi W Tehuantepec ( D.H.Janzen ) ; 1mi W Temascal ( D.H.Janzen ) ; 22.2mi N Puerto Escondido , 700m ( D.H.Janzen ) ; 25mi W Tehuantepec ( D.H.Janzen ) ; 3.2mi NE Tehuantepec ( D.H.Janzen ) ; 42km N San Pedro Pochutla , 850m ( W.MacKay ) ; 44mi W Tehuantepec ( E.E.Gilbert & C.D.MacNeil ) ; 5mi E Temascal ( D.H.Janzen ) ; 9.6km E Santiago Astata , 10m ( W.MacKay ) ; Bahias de Huatulco ( W.MacKay ) ; Pinotepa Nacional ( S.W.T.Batra ) ; Salina Cruz ( D.H.Janzen ) ; Tehuantepec ( F.Knab ; W.P.Stoutamize ) ; Temascal ( H.V.Daly ; D.H.Janzen ) ; Temascal , 25m ( D.H.Janzen ) ; Tuxtepec ( J.CamelaG. ; W.M.Mann ) ; Valle Nacional ( G.V.Halffter ) ; Q.Roo : 12.2mi S Peto, Q.Roo-Yucatan border ( D.H.Janzen ) ; 16.9mi W Cd. Chetumal, rd. to Peto ( D.H.Janzen ) ; 22.5mi S Felipe Carillo Puerto ( D.H.Janzen ) ; 6.4mi E Polyuc ( D.H.Janzen ) ; 7mi W Felipe Carillo Puerto ( D.H.Janzen ) ; 8.1mi S Felipe Carillo Puerto ( D.H.Janzen ) ; 8mi S Felipe Carillo Puerto ( D.H.Janzen ) ; Sian Ka'an ( A.Dejean ) ; Sian Ka'an Reserve, nr. Felipe Carillo Puerto ( A.Dejean ) ; Vallarta ( A.Dejean ) ; S.L.P. : 27mi N Tamazunchale ( D.H.Janzen ) ; 40-50mi NW Cd. del Maiz ( W.S.Ross ) ; 4miN Valles , 300ft . ( W.S.Creighton ) ; Cd. Valles ( D.H.Janzen ) ; El Bonito, 7mi S Cd. Valles , 300ft . ( P.H. & M.Arnaud ) ; El Salto ( A.Mintzer ; c.u. ) ; Huichihuayan ( L.J.Lipovsky ) ; Rio Amahac, Tamazunchale , 300ft . ( W. S. Creighton ) ; Tamazunchale ( D. H. Janzen ; W. S. Ross ) ; locality not specified, prob. Tanquian [see Safford 1923:390] (Safford) ; Tab. : 0.6mi S Paraiso onrd. to Cardenas ( D.H.Janzen ) ; 0.9mi S Chontalpa ( D.H.Janzen ) ; 12.8mi S Chontalpa ( D.H.Janzen ) ; 2.1mi E Frontera ( D.H.Janzen ) ; 3 mi W Cardenas ( D.H.Janzen ) ; 30.2mi W Cd. El Carmen ( D.H.Janzen ) ; 37.8mi E Coatzacoalcos, Hwy.180 ( D.H.Janzen ) ; 6.6mi S Chontalpa ( D.H.Janzen ) ; Chontalpa, 26mi S Cardenas ( D.H.Janzen ) ; Cardenas ( D.H.Janzen ) ; Teapa ( J.C. & D.Pallister ; H.H.Smith ; W.P.Stoutamize ) ; Tamps. -. 22.7mi S Cd. Victoria ( D.H.Janzen ) ; 41mi S Cd. Victoria ( C.W.Obrien ) ; 50mi N Valles ( G.E.Bohart ) ; 5mi N Cd. Mante ( A.Mintzer ) ; 7km WSW El Encino , 140m ( P.S.Ward ) ; Antiguo Morelos ( c.u. ) ; Cd. Madero ( F.Infante M. ) ; Ciudad Mante ( N.E. & M. A.Evans ; N.L.H.Krauss ) ; El Limon ( H.E.Evans ) ; Forlon ( L.J.Lipovsky ) ; Gonzales ( c.u. ) ; Llera ( W.E.LaBerge ) ; Mesa de Llera ( A.Mintzer ) ; N of Antiguo Morelos ( A.Mintzer ) ; Rio Guayalejo at Hwy .85 ( A.S.Menke & L.A.Stange ) ; S of Cd. Mante ( A.Mintzer ) ; Tampico ( Brakhoven ; D.L.Crawford ; H.Jourdan ; N.L.H.Krauss ; Locke ; E.Palmer ; E.A.Schwarz ; W.P.Stephen ; c.u. ) ; Tampico, dunes at Cd. Madero ( D.H.Janzen ) ; Xicotencatl ( H.C.Millender ) ; Ver. : " N.M.,Vera Cruz " ( Townsend ) ; " Vera Cruz " ( G.Seurat ; H.H.Smith ) ; 10mi W Veracruz ( G.E.Bohart ) ; 14km ENE La Tinaja , 50m ( P.S.Ward ) ; 15mi W Veracruz ( U.Kansas Mex.Exped .) ; 16km E Cuilahuac [= Cuitlahuac ] ( W.MacKay ) ; 24.9mi NW Acayucan ( D.H.Janzen ) ; 28km N Cardel, Morro de la Mancha ( V.Rico-Gray ) ; 30mi S Acayucan ( F.D.Parker ) ; 3mi N Sayula ( R.C.Bechtel ; R.C.Bechtel & E.I.Schlinger ) ; 4mi NE Minatitlan ( R.C.Bechtel & E.I.Schlinger ) ; 4miNWRinconada Antigua , 1350ft . ( U.KansasMex.Exped. ) ; 4mi W Puente Nacional , 900ft . ( U.Kansas Mex.Exped. ) ; 6.5km N Tierra Blanca , 50m ( W.MacKay ) ; 7mi S Cardel , 600ft . ( c.u. ) ; 9km NNW Sontecomapan , 20m ( P.S. Ward ) ; Boca del Rio ( U.Kansas Mex.Exped. ) ; Buen Pais ( R.C.Bechtel & E.I.Schlinger ) ; Camaron ( E.Skwarra ) ; Cordoba ( F.Knab ; R.R.Snelling ; c.u. ) ; Cotaxtla Exp. Sta., Cotaxtla ( D.H.Janzen ) ; Est. Biol. Los Tuxtlas ( H.A.Hespenheide ) ; Est. Biol. "Los Tuxtlas", nr. San Andres Tuxtla ( G.Ibarra M. ) ; Fortin ( c.u. ) ; Jalapa ( Rangel ; W.M.Wheeler ; c.u. ) ; Jalapilla, mun. Jilotepec ( G. Aleman ) ; Los Cocos ( A.Petrunkewitch ) ; Los Tuxtlas, 10km NNW Sontecomapan , 200m ( P.S.Ward ) ; Mirador ( E.Skwarra ) ; Mocambo ( D.H.Janzen ) ; Orizaba ( c.u. ) ; Palma Sola ( Halffter & Reyes ) ; Panuco ( F.Parker & D.Miller ) ; Playa Azul, Catemaco ( W.P.Stoutamize ) ; Pueblo Nuevo, nr. Tezonapa ( Cornell Univ. Mex. Field Party ) ; Remutadero ( E.Skwarra ) ; Santa Lucrecia [=Jesus Carranza] ( F.Knab ; P.Knab ) ; Tamarindo ( E.Skwarra ) ; Tamos ( F.C.Bishop ) ; Tinajas [presumably La Tinaja, on Hwy.150] ( F.D.Parker & L.A.Stange ) ; Tlacocintla ( E.Skwarra ) ; Veracruz ( G.E.Bohart ; N.L.H.Krauss ; E.Skwarra ; L.A.Stange ) ; Yuc. : 14.8mi S Ticul, "Hwy.180" [prob.Hwy.261] ( D.H.Janzen ) ; 8mi E Merida (rd. to Pto. Juarez) ( D.H.Janzen ) ; Merida ( D.H.Janzen ; N.L.H.Krauss ) ; Merida (S margin of town) ( D.H.Janzen ) ; Tekax, 33mi W Peto ( D.H.Janzen ) ; Temax ( Gaumer ) ; state unknown : " Mex "(" Norton " ) .