Argia medullaris Selys, 1865

Garrison, Rosser W. & Ellenrieder, Natalia Von, 2018, Damselflies of the genus Argia (Odonata: Coenagrionidae) from Ecuador with descriptions of five new species, Zootaxa 4470 (1), pp. 1-69 : 20-24

Argia medullaris Selys, 1865

Figs. 18 (head, thorax, S1– 3 ♂), 63, 64 (S7– 10 ♀), 80 (mesostigmal plates ♀), 98 (genital ligula), 115–117 (appendages ♂), 129 (lectotype labels), 130 (holotype labels A. rectangula ), 132 (lectotype labels A. columbiana ), 133 (holotype labels A. augustana ), 135 (S8– 10 ♀ holotype of A. augustana [from Navás 1934]), 138 (map), 147 (photo).

Argia medullaris Hagen in Selys 1865: 412 (40 separate) (description of ♂, " Bogota, Nouvelle-Grenade, par Lindig. (Collect. Hagen, de Selys)"); Calvert in Calvert & Hagen 1902: 111 (catalog, Plate 1, Figs. 2, 2a .); Calvert 1907: 374 ( Guatemala, Costa Rica); Förster 1914: 64 (comparison of A. fraudatricula sp. nov. to A. medullaris ); Ris 1918: 116 ( A. variabilis compared to A. medullaris ); De Marmels 1990: 337 ( Venezuela); Ramírez et al. 2000: 249 ( Costa Rica); Förster 2001:11 ( Guatemala, Costa Rica); Abbott et al. 2002: 126 ( Nicaragua); Esquivel 2005: 286 ( Mexico [in error], Guatemala, Costa Rica–Colombia, Ecuador, Venezuela); von Ellenrieder & Garrison 2007: 19 (mention of syntypes and identity); Heckman 2008: 350, 351, 384 (key, figs. based on Calvert & Hagen 1902); Hoffmann 2009: 27, 39 ( Peru); Garrison et al. 2010: 213 (listed,); Pérez-Gutiérrez & Palacino-Rodriguez 2011: 212 ( Colombia); Garrison & von Ellenrieder 2013: 16, 18, 22 ( Costa Rica, field photo); Pérez-Gutiérrez et al. 2013: 100 ( Colombia, field photo);.

Argia ? medullaris: Calvert 1902: 91–92 , p. 92, Tab. IV. figs. 6, 54, 54s (in part, specimens from Guatemala, Costa Rica, Colombia, determined as possible A. medullaris ).

Argia rectangula Navás 1920: 1–2 (Patria. Colombia: Viotá, I[J]ul. 1915, specimen ♀ in mea collectione a F. Apollinari missum). NEW SYNONYMY.

Argia columbiana Navás 1924: 327–328 [16–17 separate] (Patria. Colombia: La Mesa, Julio de 1925 [sic! Should be 1915]; Tusagasuga, Julio de 1913; Cordillera central, Cordillera de Bogotá, vertiente occidental. (Col. m.).). NEW SYNONYMY.

Argia augustana Navás 1934: 180 [28 separate] (Patria. Colombia: San Antonio de Tena, agosto de 1926. Un ejemplar ♂ que recibí del P. Uribe y que tenía todavía sin estudiar en mi colección, aguardando ver otro sexo.). Garrison et al. 2003: 27 (catalog, synonymy with A. medullaris ).

Argia charalana: Navás 1935: 33–34 [1–2 separate] ( Colombia: Pensilvania, 1913, 1918; Villavicencio, Septiembre de 1912; Pandi, Agosto de 1911; Medina, Enero de 1918).

Types: of medullaris: The description was based on at least two syntype males ("Collect. Hagen, de Selys"). One, the MCZ paralectotype, lacks the last four abdominal segments and the other, a complete male in the Selys collection ( von Ellenrieder & Garrison 2007), is the one that we here designate as lectotype. Paralectotype ♂:"St. Fe de Bogota / Lindig 1863 [written]", " A medullaris Hag. [written]", "Hagen [printed]", "Penis / drawn [written by C.H. Kennedy]". Type [printed] MCZ 12177 [written], in MCZ; lectotype: same labels but "42. A. medullaris / ♂ [written]; "medullaris / ♂ [written]"; "desseiné / Williamson [penciled]"; "Desseiné par/Santos—3.x.64 [written].", in RBINS ( Fig. 129 );—of augustana : Holotype ♀ —small, pale blue hand written label, "S. Antonio de Tena / ( Colombia) / Agosto 1926 "; rectangular green label, " Argia / augustana ♂ Nav. [hand printed] / P. Navás S. J. det. [printed]"; small pink label, "Typus" [printed in Navás's hand] ( Fig. 133 ); on accompanying envelope, " Argia augustana ♂ Nav. / Colombia / tipo Regalo", in UMMZ;—of columbiana : small, white hand written label, " La Mesa / VII-15"; rectangular green label, " Argia / columbiana Nav. [hand printed] / P. Navás S.J. det. [printed]"; small pink label, "Typus" [printed in Navás' hand]; small printed green label, "MUSEUM PARIS"; white label, " Argia columbiana / Navas 1924 / Lectotype ♂ [in ink by Legrand] / J. Legrand dés. [printed] 1996 [in ink], rectangular printed red label, " LECTOTYPE " ( Fig. 132 ). The specimen is the Lectotype [MNHN]; ♀: small, white hand written label, " La Mesa / VII-15"; rectangular green label, " Argia / columbiana & Nav. [hand printed] / P. Navás S.J. det. [printed]"; small pink label, "Typus" [printed in Navás' hand]; small printed green label, "MUSEUM PARIS"; white label, " Argia columbiana / Navas 1924 / Paralectotype & [in ink by Legrand] / J. Legrand dés. [printed] 1996 [in ink], rectangular printed red label, " PARALECTOTYPE ". The specimen is a paralectotype. Lacking S7-10 [MNHN];—of rectangula : small, white hand written label, " Viotá (C.) / VII-15"; rectangular green label, " Argia / rectangula ♀ Nav. [hand printed] / P. Navás S.J. det. [printed]"; small pink label, "Typus" [printed in Navás' hand]; small printed green label, "MUSEUM PARIS"; white label, " Argia rectangula / Navás 1920 / Holotype ♀ [in ink by Legrand] / J. Legrand dés. [printed] 1994 [in ink], rectangular printed red label, " HOLOTYPE " ( Fig. 130 ). The specimen is the Holotype [MNHN].

Specimens examined (only those from Ecuador listed). 521 ♂♂, 198 ♀♀: Imbabura Prov.: 2 ♂♂, Guallupe , {0°47'0'' N, 78°18'0'' W}, 1,600 m, 10 February 1946 [no collector given] [ CSCA] ; same data but 2 ♂♂ [ RWG] ; same data but 1 ♀ [ UMMZ] ; same data but 1 ♂, 11 February 1946 [ CSCA] ; same data but 2 ♂♂ [ UMMZ]; Manabí Prov.: 1 ♂, 1 ♀, Cojimies {0°22'0'' N, 80°2'0'' W, 5 m}, March 1949, William Clarke-Macintyre leg. [ UMMZ]; Pichincha Prov.: 2 ♂♂, Bella Vista {0°0'57'' S, 78°40'50'' W}, August 1948, William Clarke-Macintyre leg. [ CSCA] ; same data but 2 ♂♂ [ RWG] ; same data but 7 ♂♂ [ UMMZ]; 1 ♂, Sachatamia Lodge, 4 km northeast of Mindo, Pachijal River tributary (0°1'24'' S, 78°45'30'' W, 1630 m), 5 May 2009, William Haber & Fred Morrison leg. [ WAH]; Santo Domingo de los Tsáchilas Prov.: 1 ♂, Santo Domingo de los Colorados {0°15'0'' S, 79°9'0'' W, 604 m}, July 1939, William Clarke-Macintyre leg. [ CSCA] ; same data but 1 ♀ [ RWG] ; same data but 1 ♂ [ UMMZ]; Napo Prov.: 1 ♂, 17 km east of Coyuja, small stream {0°27'51'' S, 77°54'30'' W, 1,823 m}, 21 August 1980, Sidney W. Dunkle leg. [ RWG]; 1 ♂, marsh 17 km east of Cuyujúa on road to El Chaco {0°27'0'' S, 77°53'0'' W, 1,796 m}, 17 August 1980, Sidney W. Dunkle leg. [ RWG]; 1 ♀, marsh/ stream 3.5 km north of El Chaco {0°18'59'' S, 77°47'31'' W, 1,513 m}, 7 June 1995, Sidney W. Dunkle leg. [ RWG]; 1 ♂, Verde Yacu River {0°43' S, 77°58' W, 2,280 m}, 4–29 April 1984, Herman G. Real leg. [ RWG]; 1 ♂, seepage marsh 10.6 km south of Baeza {0°30'30'' S, 77°52'44'' W, 1,800 m}, 15 June 1995, Sidney W. Dunkle leg. [ RWG]; 1 ♂, 1 ♀, seepage, route E45, about 10 km south of Baeza (0°22'31'' S, 77°49'9'' W, 1,580 m), 4 November 2013, Kenneth J. Tennessen leg. [ FSCA]; 1 ♂, small stream 5 km south of Baeza {0°27'0'' S, 77°53'0'' W, 1,796 m}, 25 July 1996, Kenneth J. Tennessen leg. [ RWG]; 1 ♂, small stream, 5 km south of Baeza {0°28'6'' S, 77°52'2'' W, 1,900 m}, Sidney W. Dunkle leg. [ RWG]; 1 ♂, 1 ♀, Yanayacu trail, Yanayacu Biological Station & Center for Creative Studies, 5 km west of Consanga {0°35'57'' S, 77°53'24'' W, 2,100 m}, 11 June 2006 [no collector given] [ RWG]; 1 ♂ , same data but Yanayacu trail to falls [ RWG]; 1 ♂, 1 ♀, Mapoto, Pastaza River , 1,300 m, 28 September 1938, William Clarke- Macintyre leg. [ RWG]; Tungurahua Prov.: 1 ♀, Baños {1°24'0'' S, 78°24'0'' W, 2,000 m}, 26 February 1936, William Clarke-Macintyre leg. [ RWG] ; same data but 1 ♀ [ UMMZ] ; same data but 1 ♂, 1 ♀, 2 March 1936 [ CSCA] ; same data but 2 ♂♂, 1 ♀ [ RWG] ; same data but 1 ♂ [ UMMZ] ; same data but 1 ♂, 7 March 1936 [ UMMZ] ; same data but 1 ♂, 9 March 1936 [ CSCA] ; same data but 3 ♂♂, 1 ♀, 11 March 1936 [ CSCA] ; same data but 2 ♂♂, 2 ♀♀ [ RWG] ; same data but 5 ♂♂, 1 ♀ [ UMMZ] ; same data but 1 ♂, February 1936 [ CSCA] ; same data but 3 ♂♂, June 1936 [ RWG] ; same data but 1 ♂, 2 ♀♀, February 1936 [ UMMZ] ; same data but 1 ♂, June 1936 [ UMMZ] ; same data but 4 ♂♂, April 1936 [ UMMZ]; 1 ♂, 1 ♀, Baños, Lamas {1°24'0'' S, 78°24'0'' W, 2,000 m}, 4 January 1938, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♀, 17 October 1938 [ UMMZ]; 1 ♂, 1 ♀, Baños, Punzan {1°24'0'' S, 78°24'0'' W, 2,200 m}, 6 March 1946, William Clarke-Macintyre leg. [ CSCA] ; same data but 2 ♂♂, 1 ♀ [ RWG] ; same data but 10 ♂♂, 1 ♀ [ UMMZ]; 1 ♀, Baños, Pastaza River watershed {1°24'0'' S, 78°24'0'' W, 1,900 m}, 25 May 1937, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♀, 1 January 1938 [ UMMZ] ; same data but 1 ♀, 25 April 1939 [ RWG] ; same data but 1 ♂, December 1937 [ CSCA] ; same data but 1 ♂, 1 ♀, September 1938 [ RWG]; 2 ♂♂, brook into Blanco River, 12 km east of Baños, Pastaza River watershed {1°23'0'' S, 78°18'0'' W, 1,700 m}, 9 April 1936, William Clarke-Macintyre leg. [ RWG]; 1 ♂, Chaupi, on Mount Tungurahua {1°6'0'' S, 78°39'0'' W, 3,000 m}, 24 September 1937, William Clarke-Macintyre leg. [ CSCA]; 1 ♂, 1 ♀, El Tablón , near Baños {1°8'0'' S, 78°37'0'', 2,000 m}, May 1936, William Clarke-Macintyre leg. [ CSCA] ; same data but 2 ♀♀ [ UMMZ]; 3 ♂♂, Guama Yacu (= Guamba River), near Agoyan , Pastaza River watershed {1°24'45'' S, 78°22'16'' W}, May 1936, William Clarke-Macintyre leg. [ UMMZ]; 1 ♂, Guama , Pastaza River watershed {1°24'0'' S, 78°25'0'' W, 2,500 m}, 15 June 1936, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♂, June 1936 [ UMMZ]; 1 ♂, 1 ♀, Hacienda La Merced, Pastaza River watershed , {1°23'0'' S, 78°17'0'' W, 1,900 m}, 5 September 1936, William Clarke-Macintyre leg. [ CSCA] ; same data but 2 ♂♂, April 1936 [ RWG] ; same data but 2 ♂♂ [ UMMZ]; 1 ♀, Hacienda La Palmera , {1°25'0'' S, 78°12'0'' W, 1,300 m}, 7 December 1938, William Clarke- Macintyre leg. [ UMMZ] ; same data but 1 ♀, 8 December 1938 [ UMMZ] ; same data but 1 ♀, 9 December 1938 [ UMMZ] ; same data but 1 ♂, 12 December 1938 [ CSCA] ; same data but 1 ♀, December 1938 [ RWG] ; same data but 4 ♀♀ [ UMMZ]; 1 ♂, La Esperanza, Blanco River , Pastaza watershed {1°23'57'' S, 78°21'3'' W, 1,500 m}, August 1936, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♂, July 1936 [ UMMZ]; 2 ♂♂, Chin-chin River, near Baños {1°23'0'' S, 78°19'0'' W, 1,800 m}, 8 March 1946, William Clarke-Macintyre leg. [ UMMZ]; 1 ♂, 1 ♀, Blanco River Valley , Camp 4, Pastaza watershed {1°23'0'' S, 78°18'0'' W, 1,600 m}, May 1936, William Clarke-Macintyre leg. [ CSCA]; 1 ♂, Blanco River , Pastaza River watershed {1°23'0'' S, 78°18'0'' W, 1,600 m}, 7 August 1936, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♂, April 1937 [ RWG] ; same data but 8 ♂♂, April 1937 [ UMMZ] ; same data but 1 ♂, May 1937 [ UMMZ] ; same data but 1 ♀, 5 September 1936 [ UMMZ] ; same data but 2 ♂♂, 21 August 1937 [ UMMZ] ; same data but 1 ♀, 27 November 1937 [ UMMZ] ; same data but 1 ♀, August 1938 [ UMMZ] ; same data but 4 ♂♂, 1 ♀, 15 October 1938 [ UMMZ] ; same data but 2 ♂♂, 4 ♀♀, November 1938 [ UMMZ] ; same data but 1 ♀, January 1939 [ UMMZ] ; same data but 1 ♀, February 1939 [ UMMZ] ; same data but 3 ♂♂, March 1939 [ UMMZ] ; same data but 2 ♂♂, 12 May 1939 [ UMMZ] ; same data but 1 ♀, 29 June 1939 [ RWG] ; same data but 1 ♂, July 1939 [ UMMZ] ; same data but 2 ♂♂, 2 ♀♀, 2 September 1939 [ UMMZ] ; same data but 2 ♀♀, 6 September 1939 [ UMMZ] ; same data but 2 ♂♂, 1 ♀, 18 September 1939 [ CSCA] ; same data but 10 ♂♂, 15 ♀♀ [ UMMZ] ; same data but 14 ♂♂, 3 ♀♀, 5 October 1939 [ UMMZ] ; same data but 1 ♂, 1 ♀, 18 October 1939 [ UMMZ] ; same data but 2 ♂♂, 22 October 1939 [ UMMZ] ; same data but 2 ♂♂, 23 October 1939 [ RWG] ; same data but 1 ♂, 1 ♀, 25 October 1939 [ CSCA] ; same data but 1 ♂, 1 ♀ [ RWG] ; same data but 2 ♂♂, 3 ♀♀ [ UMMZ] ; same data but 1 ♂, 26 October 1939 [ RWG] ; same data but 1 ♂, 1 ♀ [ UMMZ] ; same data but 1 ♀, 28 October 1939 [ RWG] ; same data but 1 ♂ [ UMMZ] ; same data but 8 ♂♂, 2 ♀♀, 1 November 1939 [ CSCA] ; same data but 5 ♂♂, 3 ♀♀ [ RWG] ; same data but 64 ♂♂, 17 ♀♀ [ UMMZ] ; same data but 1 ♂, 1 ♀, 10 May 1941 [ RWG] ; same data but 1 ♂ May 1949 [ CSCA] ; same data but 4 ♂♂, 4 ♀♀, May 1949 [ RWG] ; same data but 2 ♂♂, May 1949 [ UMMZ]; 1 ♂, Huambo River, Baños, Pastaza River watershed , {1°24'0'' S, 78°24'0'' W, 1,900 m}, 24 July 1936, William Clarke-Macintyre leg. [ CSCA]; 2 ♂♂, Pastaza River between Yunguilla and Abitagua {1°25'0'' S, 78°14'0'' W, 1,600 m}, 1 November 1939, William Clarke-Macintyre leg. [ CSCA] ; same data but 6 ♂♂ [ RWG] ; same data but 27 ♂♂, 3 ♀♀ [ UMMZ]; 1 ♂, 1 ♀, San Pedro River ( Baños ) {1°25'0'' S, 78°14'0'' W, 1,400 m, 8 March 1946, William Clarke-Macintyre leg. [ CSCA] ; same data but 2 ♂♂, 3 ♀♀ [ RWG] ; same data but 11 ♂♂, 2 ♀♀ [ UMMZ]; 7 ♂♂, Topo River, Pastaza River watershed , 1,500 m, 12 March 1936, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♂, 27 March 1936 [ RWG] ; same data but 1 ♀ [ UMMZ] ; same data but 1 ♂, 28 March 1936 [ RWG] ; same data but 1 ♂, 1 ♀ [ UMMZ] ; same data but 1 ♂, 29 March 1936 [ RWG] ; same data but 2 ♂♂, 12 December 1948 [ RWG] ; same data but 1 ♀, 2 February 1949 [ RWG]; 1 ♂, 1 ♀, San Francisco, Pastaza River , {1°18' S, 78°30' W, 1,200 m}, 16 October 1937, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♂, 1 October 1938 [ UMMZ]; 1 ♀, Valencia, Baños {1°24'2'' S, 78°7'30'' W, 1,600 m}, 27 April 1937, William Clarke-Macintyre leg. [ CSCA] ; same data but 1 ♀ [ UMMZ]; 1 ♂, vicinity of Baños {1°24'0'' S, 78°24'0'' W, 1,800 –2,500 m}, 1936, William Clarke-Macintyre leg. [ UMMZ]; 1 ♂, Yunguilla, Pastaza River {1°23'58'' S, 78°20'57'' W, 1,600 m}, June 1936, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♀, 8 September 1936 [ UMMZ] ; same data but 6 ♂♂, 2 ♀♀, 10 September 1936 [ UMMZ] ; same data but 3 ♂♂, May 1938 [ UMMZ] ; same data but 1 ♂, 1 ♀, 21 April 1939 [ UMMZ] ; same data but 1 ♂, 27 April 1939 [ UMMZ] ; same data but 1 ♀, April 1939 [ UMMZ] ; same data but 1 ♀, 23 August 1939 [ RWG] ; same data but 6 ♂♂, 8 ♀♀, 23 August 1939 [ UMMZ] ; same data but 1 ♂, 1 ♀, 18 October 1939 [ RWG] ; same data but 1 ♀ [ UMMZ]; 1 ♂, Yunguilla, where Blanco River enters Pastaza River {1°23'0'' S, 78°18'0'' W, 1,500 m}, 20 August 1936, William Clarke-Macintyre leg. [ UMMZ] ; same data but 1 ♂, 10 September 1936 [ BMNH] ; same data but 1 ♂, 1 ♀ [ UMMZ] ; same data but 1 ♀, 30 October 1937 [ UMMZ] ; same data but 2 ♂♂, 1 ♀, 10 June 1939 [ UMMZ] ; same data but 2 ♂♂, 1 ♀, 25 July 1939 [ UMMZ] ; same data but 1 ♂, 1 ♀, 28 July 1939 [ UMMZ] ; same data but 1 ♀, December 1939 [ UMMZ]; Pastaza Prov.: 1 ♂, Abitagua, Pastaza River watershed , {1°27'0'' S, 78°9'0'' W}, 1,200 m, July 1936, William Clarke-Macintyre leg. [ RWG] ; same data but 3 ♂♂, October 1936 [ UMMZ] ; same data but 1 ♂, 10 November 1936 [ UMMZ] ; same data but 1 ♂, 13 October 1937 [ UMMZ] ; same data but 2 ♂♂, 25 May 1939 [ UMMZ] ; same data but 2 ♂♂, 26 May 1939 [ UMMZ] ; same data but 1 ♂, 23 August 1939 [ RWG] ; same data but 1 ♂, 25 September 1939 [ RWG] ; same data but 1 ♂, 1 October 1939 [ CSCA] ; same data but 1 ♂ [ UMMZ] ; same data but 1 ♂, 1 ♀, 2 October 1939 [ CSCA] ; same data but 1 ♂, 1 ♀ [ RWG] ; same data but 15 ♂♂, 1 ♀ [ UMMZ] ; same data but 1 ♀, 6 October 1939 [ RWG] ; same data but 2 ♂♂, 9 October 1939 [ RWG] ; same data but 1 ♂, 10 October 1939 [ RWG] ; same data but 7 November 1939 [ UMMZ] ; same data but 1 ♂, 1 ♀, 8 November 1939 [ UMMZ] ; same data but 1 ♂, 1 ♀, 13 November 1939 [ RWG] ; same data but 1 ♂ [ UMMZ] ; same data but 1 ♂, 21 November 1939 [ UMMZ] ; same data but 1 ♂, 12 December 1939 [ RWG] ; same data but 2 ♂♂, 20 December 1939 [ UMMZ] ; same data but 1 ♀, 30 December 1939 [ RWG] ; same data but 1 ♂, 18 January 1940 [ RWG] ; same data but 1 ♂, 19 March 1940 [ RWG] ; same data but 1 ♂, 9 April 1940 [ UMMZ] ; same data but 1 ♂, 14 April 1940 [ CSCA] ; same data but 1 ♀, 19 February 1941 [ CSCA] ; same data but 1 ♂, 20 February 1941 [ RWG] ; same data but 3 ♂♂, 1 ♀, December 1948 [ RWG]; 1 ♀, Abitagua, Pastaza River watershed, creek Guillermina {1°27'0'' S, 78°9'0'' W, 1,200 m}, 26 May 1941, William Clarke-Macintyre leg. [ UMMZ]; 1 ♂, Abitagua , Pastaza River watershed , mouth of Fox River , {1°27'0'' S, 78°9'0'' W, 1,200 m}, 25 February 1941, William Clarke-Macintyre leg. [ CSCA]; 2 ♂♂, between Puyo and Mera {1°29' S, 78°3' W, 1,000 m}, February 1939, William Clarke-Macintyre leg. [ RWG]; 2 ♀♀, El Sucre, Verde River , Pastaza River watershed {1°24'12'' S, 78°18'3'' W, 1,500 m}, Aril 1937, William Clarke-Macintyre leg. [ UMMZ]; 1 ♂, Hacienda Guillermina, Abitagua , Pastaza River watershed , {1°27'0'' S, 78°9'0'' W, 1,200 m}, 10 December 1939, William Clarke-Macintyre leg. [ UMMZ]; 2 ♂♂, 1 ♀, Puyo (1°28'0'' S, 77°59'0'' W, 1,000 m), 17 April 2008, T. & K. Miyashita leg. [ HK] ; same data but 1 ♂ [ RWG] ; same data but 2 ♂♂, April 2009 [ HK]; 1 ♂, 1 ♀, Napo River, Pastaza River watershed {1°28'0'' S, 77°59'0'' W, 2,500 m}, 2 April 1936, William Clarke-Macintyre leg. [ CSCA] ; same data but 1 ♂ [ MCZ] ; same data but 2 ♂♂, 1 ♀ [ RWG] ; same data but 3 ♂♂ [ UMMZ] ; same data but 3 ♂♂, 1 ♀, 30 April 1936 [ CSCA] ; same data but 10 ♂♂ [ RWG] ; same data but 6 ♂♂ [ UMMZ] ; same data but 1 ♂, 1 ♀, May 1936 [ RWG] ; same data but 1 ♀, 1 October 1936 [ UMMZ] ; same data but 1 ♂, 1 ♀, 5 October 1936 [ BMNH] ; same data but 1 ♂ [ RWG] ; same data but 13 ♂♂, 4 ♀♀ [ UMMZ] ; same data but 1 ♂, 1 ♀, 12 October 1936 [ RWG] ; same data but 5 ♂♂, 1 ♀ [ UMMZ] ; same data but 1 ♂, 24 October 1936 [ UMMZ]; 2 ♂♂, Verde Chico River, Pastaza River watershed , {1°23'24'' S, 78°23'29'' W, 1,700 m}, 7 February 1939, William Clarke-Macintyre leg. [ RWG] ; same data but 5 ♂♂, 3 ♀♀, May 1939 [ RWG] ; same data but 7 ♂♂, 1 ♀, May 1939 [ UMMZ] ; same data but 2 ♂♂, 1 ♀, May 1949 [ RWG] ; same data but 14 ♂♂, 5 ♀♀, May 1949 [ UMMZ]; 1 ♂, Verde River, Pastaza River watershed {1°24'12'' S, 78°18'3'' W, 1,500 m}, 25 October 1935, William Clarke-Macintyre leg. [ CSCA] ; same data but 1 ♂, 27 October 1938 [ RWG] ; same data but 1 ♂, 29 June 1939 [ RWG]; 2 ♂♂, 1 ♀, Vizcay, Pastaza River watershed , June 1936, William Clarke-Macintyre leg. [ CSCA] ; same data but 7 ♂♂, 2 ♀♀, June 1936 [ UMMZ] ; same data but 2 ♀♀, January 1939 [ CSCA] ; same data but 2 ♀♀, January 1939 [ RWG] ; same data but 2 ♂♂, 2 ♀♀, January 1939 [ UMMZ] ; same data but 1 ♂, 1 ♀, March 1939 [ CSCA] ; same data but 1 ♂, 1 ♀, March 1939 [ MCZ] ; same data but 1 ♂, March 1939 [ RWG] ; same data but 5 ♂♂, 1 ♀ March 1949 [ UMMZ]; Morona Santiago Prov.: 1 ♂, 1 ♀, Guilca, Upano River [received September 28, 1945] {1°57'0'' S, 77°51'46'' W}, 1,400 m, Leopoldo Gomez A. leg. [ UMMZ]; 1 ♂, 2 ♀♀, Macas , Upano River [received September 28, 1945] {1°57'0'' S, 77°51'46'' W, 1,050 m}, Leopoldo Gomez A. leg. [ UMMZ] ; same data but 1 ♂ [ UMMZ]; 3 ♂♂, 1 ♀, Mangosisa, Upano River [received Sept. 28, 1945] {2°31'28'' S, 77°53'41'' W, 850 m}, Leopoldo Gomez leg. [ RWG] ; same data but 2 ♂♂ [ UMMZ]; 3 ♂♂, Salado, Upano River {2°15'0'' S, 78°16'0'' W, 2,550 m}, November 1948, William Clarke-Macintyre [ RWG]; El Oro Prov.: 2 ♂♂, Moro Moro , 1,500 m, 15 July 1941, David B. Laddey leg. [ RWG] ; same data but 1 ♂ [ UMMZ]; 2 ♂♂, Pinas {3°40'0'' S, 79°39'0'' W, 1000 m}, 25 July 1941, David B. Laddey leg. [ RWG] ; same data but 2 ♂♂ [ UMMZ]; Loja Prov.: 1 ♂, San Francisco Scientific Station, San Francisco River [ID 25] (3°58'26'' S, 79°4'45'' W, 1717 m), 28 October 2014, Christoph Willigalla leg. [ CW] .

Remarks: Argia medullaris is a common species in Ecuador but it has been confused in the past with the more northerly ( Mexico) A. variabilis Selys. The latter species is not treated further here but we recognize both as separate species based on examination of type material. Males of Argia medullaris and A. variabilis Selys are unique in possessing a largely pale coloration on the thorax, dorsum of abdomen, and a continuous black inferior stripe along S3–10 ( Fig. 18 ). Argia medullaris was described from at least two males from "Bogota, Nouvelle- Grenade [ Colombia], par Lindig (Collect. Hagen, de Selys)". An incomplete male lacking S7–10 is in the MCZ and the other, the lectotype, is in the RBINS ( von Ellenrieder & Garrison 2007) and is illustrated here ( Fig. 115 ).

Applying the name A. medullaris to various populations has been challenging. We have examined specimens we believe to represent this species from Honduras, El Salvador, Nicaragua, Costa Rica, Panama, Venezuela, Colombia, and Ecuador based on comparison with the complete syntype A. medullaris lodged in the RBINS ( Fig. 115 ). The late L.K. Gloyd recognized at least three (and possibly more) phenotypes (these given various letters including sp. D [= angustana], T [= columbiana ] and Te [from Ecuador]) from material primarily from Colombia and Ecuador and it is obvious from her notes that she believed the three (and possibly other populations from Central America) to represent distinct species. She recognized what she thought were differing ratios between the width of the torus and length of the distal margin of the cercus. Although our early analysis also led us at first to conclude that more than one species was involved, subsequent relaxing and manipulation of the caudal appendages convinced us that A. medullaris comprises a wide-ranging species that varies slightly in cercal morphology in the males and slight differences in the mesostigmal plates of the females. We have been unable to correlate these minor differences to geography. We believe various postmortem positions of the caudal appendages have contributed to differences that are more apparent than real. For example, the elongate cerci can be moved posterodistally so that they appear long and linear posteriorly, or they can be positioned posterolaterally so that the appendages may resemble the cerci of A. variabilis . The external margin of the cercus can also be seen as a decumbent flap when the cercus is directed posteriorly. This flap first led us to believe in a species-specific difference that disappeared when the cercus was relaxed and positioned posterodistally.

The taxonomy of A. medullaris is further complicated by a series of poor descriptions and illustrations of specimens of Argia from Colombia described by Navás. Disposition of these names is as follows:

Argia rectangula Navás, 1920 . This name was applied to a juvenile female in fair condition lacking S5–10 that we have examined. The mesostigmal plates match those of typical A. medullaris (as in Fig. 80 ) and show no significant differences in body pattern. Consequently, we treat this name as a junior synonym of A. medullaris . Navás was apparently impressed with the Hw quadrilateral as this was the only illustration ("Regio quadrilateri in ala post.") accompanying the original description.

Argia columbiana Navás, 1924 . This name was applied to a male and female both of which we have examined. The caudal appendages of the male lectotype ( Fig. 117 ) show no significant differences from the lectotype of A. medullaris ( Fig. 115 ) and neither does body pattern. Consequently, we treat this name as a junior synonym of A. medullaris .

Argia augustana Navás, 1934 . The holotype was relaxed and placed in an envelope and sent to L. K. Gloyd in the 1930's. Examination by Gloyd and us showed it to be a junior synonym of Argia medullaris . Mesostigmal plates of holotype illustrated ( Fig. 80 ). Navás' (1934) figure of the dorsal view of S8–10 ( Fig. 135 ) is misleading as it shows a pair of isolated pale spots dorsolaterally on S9. However, these spots extend ventrally as is typical for the female of this species (Figs. 63, 64).

Argia charalana Navás, 1924 . We have examined two males and three females from Medina (♂, in UMMZ), Pandi (♂ [ Fig. 116 ], ♀, in UMMZ) and Pensilvania (2♀, in UMMZ & MNHN) labeled by Navás as A. charalana Navás, 1924 . The original description of A. charalana includes only a male with the following data: " Colombia: Charalá (Santander), Marzo de 1915". None of the series mentioned above possess data consistent with that given in the original description. Four of the five specimens listed above were later cited by Navás (1935: 33–34 [1–2 separate]) where he described the female and added the following localities: " Colombia: Pensilvania, 1913, 1918; Villavicencio, Septiembre de 1912; Pandi, Agosto de 1911; Medina, Enero de 1918.". The original description and illustration of A. charalana differ from the series later identified above by Navás as A. charalana . Illustrations of the appendages in the original description of A. charalana strongly suggest those of A. fissa Selys (see Garrison & von Ellenrieder 2017 for illustrations). Navás also describes the wings of A. charalana as tinted ("Alae membrana levissime, vix sensibiliter fulvo tincta.....") a character unique to Argia fissa in Colombia. Although the above series of A. charalana examined by Gloyd (unpubl.) and us matches Argia medullaris , we consider Argia charalana , based on the evidence given above, to be a junior synonym of Argia fissa Selys, 1865 .

Figures

FIGURES 14–18. Color pattern of male head, thorax, and S1–4. Head, dorsal view, thorax and S1–4, lateral view.

FIGURES 1–8. Color pattern of head. Head, anterior view (1); head posterior view (2–5); labrum, anterior view (6–8).

FIGURES 128–130. Type labels.

FIGURES 131–135. Type labels (131–133) and reproduced illustrations (134, 135). The penciled "♀" and "T or D?" accompanying the labels for Fig. 133 (A. augustana) were probably written by L. K. Gloyd.

FIGURES 115–118. Male caudal appendages. (a) lateral view; (b) dorsal view; (c) mediodorsal view; (d) posterior view.

FIGURES 77–83. Female mesostigmal plates. (a) dorsal view; (b) laterodorsal view; (c) lateral view; (d) detail of mesostigmal lobe, dorsal view; (e) detail of mesostigmal lobe, medial view; (f) frontal (anterior) view; (g) detail of mesostigmal lobe, posterior view; (h) posterior view.