Erythemis attala (Selys, 1857)

Rodríguez, Fredy Palacino, Sarmiento, Carlos E. & González-Soriano, Enrique, 2015, Morphological variability and evaluation of taxonomic characters in the genus Erythemis Hagen, 1861 (Odonata: Libellulidae: Sympetrinae), Insecta Mundi 2015 (428), pp. 1-68 : 9-12

Erythemis attala (Selys, 1857)

( Fig. 1a , 2a,c , 17 , 29 )

Libellula attala Selys in Sagra, 1857:445.

Libellula annulata nec Palisot (Rambur, 1842: 78) . Misidentification.

Libellula annulosa Selys in Sagra, 1857: 445.

Libellula mithra Selys in Sagra, 1857: 445.

Type material. Not examined.

Diagnosis. Abdominal dorsum black or black and yellow, and with yellow spots in several segments. In some cases, the yellow color completely covering the dorsal region of the segment (F). Thorax and basal area on HW black; basal area on HW reaching the media posterior vein (MP) or beyond, covering a small basal region or the row of marginal cells to the anal angle (MRA, Fig. 26), and extending to the base of the triangle to the posterior region. Posterior extension of ventral teeth of male cercus about the same length as the apex of epiproct or less ( Fig. 13 ). Posterior lobe of vesica spermalis covered by lateral lobe ( Fig. 19 ), which extends more posteriorly than the medial lobe. Hook bilobed, triangular, and not perpendicular to the longitudinal axis of vesica spermalis ( Fig. 18 ). Cornua parallel to the transversal axis of vesica spermalis ( Fig. 20 ), with lobes separate apically, and covered by lateral lobes in lateral view. Vulvar lamina suboval with round posterior border ( Fig. 10 ). Lab ≤ 30mm. LcaS4/lclS4 ratio>2.10; WmS2/LcaS5 ratio <3.

Morphometric ratios of males. Acd/Lban<4.10; And/Lban<4.10; Lba/Lban<2.00; Lban/Anb>1.90; LHW/Acd<3.60; LHW/LFW>0.90; Las/Ddas<4.00; LclS3/LcaS3<2.10; AnptFW/LoptFW>0.10; LclS3/ Lecv-clS3<4.00; Lecv-clS3/LcaS3<1.50; Lab/LclS3<1.50; Lab/LcaS3<3.00; Ansbt/Antr>2.70; LFW/ Lnpt<3.00; Antr/Lar>8.00; Lnpt/Anb>1.50; Lban/Lasa-ca>3.00; Lar/Lba<0.20.

Morphometric ratios of females. Lnpt/Anb>1.20 AnptFW/LoptFW>0.20; Lban/Anb<1.50; LHW/ Acd<0.50; LHW/Lab>1.20; Lab/Las>2.50; LclS3/LcaS3<3.00; Lecv-clS3/LcaS3<1.60; Lab/LclS3<1.40; Lab/LcaS3<3.00; Ansbt/Antr>2.40; LFW/Lnpt<2.90; Lar/Lba<0.14.

Head. Ll: 0.012 –0.014 mm (M); 0.013 –0.015 mm (F). Lw: 0.0031 –0.0035 mm (M); 0.0033 –0.0035 mm (F). Hwd: 0.0059 –0.0070 mm (M); 0.0063 –0.0065 mm (F). Hl: 0.0027 –0.0040 mm (M); 0.0033 –0.0037 mm (F).

Legs. Nsmf: 8.0–13 (M); 5.0–7.0 (F). Nshf: 20.0–25.0 (M); 6.0–8.0 (F). Hfl: 0.0060 –0.0066 mm (M); 0.0057 –0.0061 mm (F). Fbsd: 0.0024.00– 0.0025mm (M); 0.0022 –0.0030 mm (F). Mfl: 0.0037 –0.0041 mm (M); 0.0038 –0.0046 mm (F). Dshf (the conditions separated by commas are present in different specimens): short spines-1 to 3 median spines-3 long spines (MF, Fig. 17 ), short spines-3 long spines (M).

Wings. LoptFW: 0.0030 –0.0034 mm (M); 0.0029 –0.0033 mm (F). AnptFW: 0.006 –0.008 mm (M); 0.0006 –0.001 mm (F). LoptHW: 0.0030 –0.0035 mm (M); 0.0030 –0.0038 mm (F). AnptHW: 0.006 – 0.009 mm (M); 0.007 –0.009 mm (F). DfwHW: 0.0056 –0.0070 mm (M); 0.0063 –0.0066 mm (F). LbanFW: 15.00–19.00mm (M); 15.00–17.00mm (F). LbanHW: 13.00–17.00mm (M); 13.00–15.00mm (F). AndFW: 7.37–8.31mm (M); 7.23–7.95mm (F). AndHW: 9.90–11.00mm (M); 9.00– 10.16mm (F). LFW: 31.00– 37.40mm (M); 33.90–36.0mm (F). LHW: 35–38mm (M); 32–35mm (F). AnsbtFW: 0.0020 –0.0025 mm (M); 0.0022 –0.0025 mm (F). AntrFW: 0.006 –0.009 mm (M); 0.004 –0.008 mm (F). LarFW: 0.001 –0.003 mm (M); 0.0025 –0.0037 mm (F). LarHW: 0.0009 –0.003 mm (M); 0.002 –0.003 mm (F). LnptFW: 11.68–13.76mm (M); 12.24–12.66mm (F). LnptHW: 13.08–15.07mm (M); 13.73–14.05mm (F). At-MP: 0.002 –0.006 mm (M); 0.002 –0.004 mm (F). LbaFW: 0.0029 –0.0032 mm (M); 0.0028 –0.0030 mm (F). LbaHW: 0.0031 –0.0034 mm (M); 0.0028 –0.0033 mm (F). RP2-loptFW: 0.001 –0.006 mm (M); 0.001 –0.004 mm (F). RP2-loptHW: 0.0006 –0.004 mm (M); 0.0006 –0.005 mm (F). Dat-lsbt: 0.0009 –0.001 mm (M); 0.0012 –0.0018 mm (F). Lasa-ca: 0.003 –0.008 mm (M); 0.002 –0.007 mm (F). Crp: 7–8. Cal: 3–7. PC-RA-RP1: 2–3. CdfFW: only 3 rows, one cell divided into four sections -3 rows of cells. Vt: 1–2. Csub: 3.

Abdomen. Lab: 25.00–30.00mm (M); 23.00–30.00mm (F). LcaS3: 0.00087 –0.0011 mm (M); 0.0010 – 0.0012 mm (F). LclS3: 19.37–21.87mm (M); 0.0019 –0.0025 mm (F). Lecv-clS3: 0.0012 –0.0016 mm (M); 0.0011 –0.0013 mm (F).

Caudal appendages. Las: 0.0015 –0.0019 mm (M); 0.0008 –0.0011 mm (F). Poas: 0.001 –0.003 mm (M); 0.0012 –0.0018 mm (F). Anas: 0.002 –0.006 mm (M); 0.0025 –0.0031 mm (F). Ddas: 0.004 –0.005 mm (M). Epl: 0.0011 –0.0014 mm (M); 0.0037 –0.0043 mm (F).

Larva. Thirteen preimaginal states were described by Rodrigues-Capítulo (1983) from Punta Indio and Los Talas (Buenos Aires, Argentina). Rodrigues-Capítulo (1983) described the last instar as follows: larvae greenish with a yellowish band on mid-dorsal region of the mesothorax surrounded by two brown spots, mask with whitish and brown irregular spots, seven meridional black and violet stripes on the eyes. Average total length 13.70 mm (live specimens) 15.90–16.50 mm (preserved specimens), this measurement differs from that of Klots (1932) who recorded between 14.50 and 15.00 mm. Head width 4.8–5.0 mm, head length 2.75 mm, head width-length ratio 1.78 mm, antenna length 1.92 mm and 4.60 mm respectively. Anterior, medial and posterior tibial length 2.9, 3.6 and 6.1 mm respectively. Mouth parts with thirteen to fourteen mental setae, small teeth on the distal region of the medial lobe of the mentum, eight to nine palpal setae, outer region of maxillary palp with numerous axial and lateral hairs. Mandibles with 4 + 1 incisive and three molar teeth. Lab 5.60 mm, superior caudal appendage 1.17 mm, lateral 0.67 mm and inferior 1.19 mm. According to Costa and Pujol-Luz (1993), the larva of E. attala bears twelve premental setae, and the cercus exceeds by 1/2 the length of the epiproct.

Material examined. ARGENTINA, Prov. Jujuy, Laguna Calilegua, 23°46’60” 64°46’60”, 675m, 21- 22.iii.2006, R. W. Garrison and N. v. Ellenrieder leg., 1 male ( RWG). BOLIVIA, Santa Cruz, Ichilo, Amboró , 30 km South-Southeast of Buena Vista, 16.iii.1960, R. Steinbach leg., 2 females ( FSCA) ; Same, but Buena Vista , 400m, 12.v.1960, R. Steinbach leg., 3 males ( FSCA) ; the same data as before except collected on the following dates: i.1936 ; ii.1937, 1 male, 4 females; same, but Santiesteban, 4 km. north of Mineros , 25.iii.1960, R.B. Cumming leg., 1 male ( FSCA) ; same, but Ñuflo de Chávez, San Ramón, 4 km SW on Hwy 9, Rio San Julián , 16.38’ 42.3” S 62.30’ 31.6” N, B. Mauffray leg., 1 male ( FSCA) ; Velasco, La Estrella, 24 km N on hwy to la Florida, Río San Martín, 15.23.616S 61.29.770 W, B. Mauffray leg., 1 male ( FSCA). BRAZIL, Amapá, Macapá , Fazendinha , IEPA, Parque Zoobotánico , ix.1997, P. Magno, 1 male ( MNRJ) ; same, but ix.1997, P. Magno, 11 males, 3 females ( MNRJ) ; Amazonas, Benjamin Constant , 20.ii.1942, 1 male ( MNRJ) ; same, but Manaus , N. Santos leg., 27.x.1959, 1 male ( MNRJ) ; same, but Rio Negro , 11.vii.1941, Parko leg., 1 male ( MNRJ) ; same, but 23.vii.1941, 1 female; same, but 23.vii.1941, 1 female; same, but 27.x.1959, 1 female; same, but Igarapé, Itacotiarae Rio Branco , N. Santos leg., 21.x.1959, 2 males ( MNRJ) ; same, but Igarapé do Mariano , Elias leg., xi.1959, 3F ( MNRJ) ; same, but Km. 38, Elias leg., xi.1959, 4M ( MNRJ) ; same, but Poraquequara , i.1973, N. Tangerini leg., 5M, 2 females ( MNRJ) ; same, but Parintins , 18.i.1973, N. Tangerini leg., 5 females ( MNRJ) ; Espírito Santo, Baixo Guandu, Estr. B. Guandu á Hituba km. 13, 01-07.xi.1970, Elias, Paulo leg., 1 male ( MNRJ) ; same, but Linhares , v.1973, Elias, Paulo leg., 1 male ( MNRJ) ; Mato Grosso, Salobra , 21-23.i.1955, A. Passarelli leg., 3 females ( MNRJ) ; Pará, Belém, Parque Museu , 03.xi.1963, Arlé leg., 1 male ( MNRJ) ; same, but Utinga , 12.ii.1963, Roppa, Mielke leg., 1 male ( MNRJ) ; Paraná, Curitiba , 01.ii.1966, O. Mielke leg., 2 males ( MNRJ) ; same data as before except collected on: 04.xii.1966, 1 female ; Rio Grande do Sul, Pelotas , 18.v.1943, C. Biezanko leg., 1 male ( MNRJ) ; same data as before except collected on following dates: 01.iv.1945, 1 male ; 15.ii.1949, 1 female ; 05.v.1943, 1 female; 02.ii.1970, 1 male; same data as before except deposited in FSCA and collected on following dates: 29.i.1956, 1 female ; 24.xii.1956, 1 male ; 04.i.1957, 1 male; 09.i.1957, 1 male, 2 females; 10.i.1957, 1 male, 1 female; 23.i.1957, 1 male; 24.i.1957, 1 male, 1 female; 25.i.1957, 2 males; 26.i.1957, 1 male, 1 female; 27.i.1957, 1 female; 29.i.1957, 1 female; 01.ii.1957, 1 female; 24.xi.1957, 1 male, 1 female; 01.i.1958, 1 male; 02.i.1958, 1 male; 05.i.1958, 1 male; 07.i.1958, 2 males; 10.i.1958, 1 female; 13.i.1958, 1 female; 18.i.1958, 1 female; 01.ii.1958, 2 males, 3 females; 02.ii.1958, 1 male; 04.ii.1958, 1 male; 02.iii.1958, 1 female; 28.iii.1958, 1 male; 02.iv.1958, 1 male; 03.iv.1958, 1 female; Rondônia, Fazenda Rancho Grande , 62 km SW of Ariquenes, 10°31’48” 62°48’0”, 165 m, 2-11.xi.1989, R. W. Garrison leg., 1 female ( RWG) ; same, but Porto Velho, Guaporé cachocina do Samuel , Rio Yamary , Pauro leg., x.1944, 1 female ( MNRJ) ; Roraima, Ariquenes , viii.1980, Bento leg., 1 male ( MNRJ) ; São Paulo, Rio Paraná , 27.i.1971, Castilho leg., MNRJ ; 33 males Sem procedência, 1 male ( MNRJ). COLOMBIA, Atlántico, Ciénaga La Larga, Palmar de Varela , 3.xi.2007, M. Lobo leg., 2 males 1 female ( UARC) ; Bolívar, Mompós, Hda La Sebastiana , 27.ii.2000, M. Wolff leg., 1 male ( CEUA) ; Meta, Villavicencio, Corpoica , “Estación la Libertad”, 385m, 1.iii.2008, E. Realpe leg., 1 male ( ANDES) ; same, but Puerto López , 220 m, 01.x.2002, S. Remolino leg., 1 male ( ANDES) ; No data, 2 males, 1 female ( UARC) ; Valle, Cali , 1000 msnm, 15.ii.1999, G. Vargas leg., 1 male (MUSENUV) ; same, but pasto, 19.ii.1999, N. Tigreros, 1 male (MUSENUV); same, but 05.iii.1998, M. Albarracín, 1 male (MUSENUV); same, but, 23.ii.1999, A. Barona leg., 1 male (MUSENUV); same, but Universidad del Valle , 1000 msnm, 01.iii.2000, P. Agudelo leg., 1 male (MUSENUV) ; same, but Tulua , 1100msnm, 17.xi.2001, A. Ramírez leg., 2 males (MUSENUV) ; same, but Jardín Botánico Mateguadua , 4º1’42”N 76º10’0,7”W, 1200msnm, 13.viii.2003, C. Bermúdez leg., 1 male (MUSENUV) ; same, but Jardín Botánico Mateguadua, 4º1’42”N 76º10’0,7”W, 1107msnm, 18.ix.2009, J. Mendivil leg., 1 male (MUSENUV). EC- UADOR, Guayas, Cantón Durán, 02.17S 79.83W, 30 ft, S.N. Rhoads leg., 12.ii.2011, 1 female ( FSCA). GUATEMALA, Quirigua , 16.ix.1915 -18, W. Schaus and J. T. Barnes leg., 1 female ( FSCA). MÉXICO, Morelos, Las Estacas , i.1984, R. Novelo leg., 1 male ( CNIN) ; Veracruz, Catemaco , 15.ix.1958, 1 male ( CNIN) ; same, but Río Otapa, 8 Km. S of La Tinaja, 18°45’0” 96°26’24”, 90m, 13.viii.1976, R.W. and J.A. Garrison leg., 1 male ( RWG). NICARAGUA, Matagalpha, Moyua , 12.36.09N 86.03.18W, 1420 ft., 28.viii.2003, F.C. Sibley leg., 1 male ( FSCA). PARAGUAY, Asunción, iv.1944, Mis. Cient. Brazil leg., 1 male ( MNRJ). PERÚ, Loreto, Iquitos , xii.1977, RWG ; 1 female same, but Explorama Lodge, 50 mi. NE Iquitos, on Amazon River at junction with Yanamono River , 24.viii.1989, S.W. Dunkle leg., 1 male, 2 females ( FSCA) ; same, but Amazon River, 50 miles NE @ Explorama Lodge , 24.vii.1989, H.V. Weems Jr. Leg., 1 male ( FSCA) ; same, but Maynas, Yanamono river at Explorama Lodge , 11.ii.1987, J. Castner leg., 1 male ( FSCA) ; same data as before except collected on: 10.ii.1987, 1 female ; same, but Muenacaño, Río Amazonas cerca Iquitos , viii.1839 ; J. Schunke leg., 1 male ( FSCA) ; same, but Pucallpa , 600 ft., 09.xi.1946, J.C. Pallister leg., 1 male ( FSCA) ; San Martín, Bella Vista, 1500 ft., 09.xii.1946, J.C. Pallister leg., 1 male ( FSCA) .

Remarks. Calvert (1928) provides data for larvae of this species, but he also warns of the potential for misidentification. This concern also is shared by Klots (1932) for the key she proposed for this species. Later Rodrigues-Capítulo (1983) redescribed the immature stages of E. plebeja clarifying the diagnostic characters proposed by the previous authors.

Biology. Larvae have been shown to dwell in a habitat where Azolla filiculoides Lam. and various species of Lemnaceae were present. Rodrigues-Capítulo (1983) described color variations in larvae, which vary depending on the substrate they use, and explains that rotifers, protozoans, oligochaetes, and filamentous algae are present due to the relative inactivity of the larva. There are reports of univoltine or semivoltine lifecycles with a facultative diapause that probably regulates the populations of both larvae and adults (Rodrigues-Capítulo 2000). Adults may be associated to ponds and lakes in forests, agricultural regions and sugar cane fields. E. attala perches on dead branches very close to the ground and it is common to find males of this species attacking those of E. plebeja (F. Palacino, pers. obs.).

Distribution. From U.S.A. to Argentina ( Fig. 29 ), between 4–2000 m.asl. De Abenante and Philippi (1982) report E. attala from Uruguay, however, without indicating any locality.

Figures

Figure 1. a) Plot showing continuous variation for the character distance between lateral and ventral carinae (dlvc), opposite to meeting point of lateral and medial transverse carinae on S3 (lmt). Numbers represent the following species: 1) E. attala, 2) E. carmelita, 3) E. collocata, 4) E. credula, 5) E. haematogastra, 6) E. mithroides, 7) E. peruviana, 8) E. plebeja, 9) E. simplicicollis, 10) E. vesiculosa. b) S1-3 E. carmelita males showing the measures from Williamson (1923): distance between lateral and ventral carinae (dlvc), divided by the measure of distance between meeting point of lateral and medial transverse carinae and ventral carinae (lmt).

Figure 2. Plots of the first two discriminant functions of the discriminant analysis. a) The first two discriminant functions explain 72.5% in males, the cases in where species are overlapped were analyzed by separated in b. b) The first two discriminant functions explain 88.3% in males, and c) The first two discriminant functions explain 83.7% in F. Polygons define the limits of proposed groups.

Figure 29. Erythemis attala distribution map.

Figures 10–17. Discrete characters of vulvar lamina, abdominal appendages, thorax, and femur.Numbers represent the following species: 10) E. peruviana female; 11) E. plebeja female; 12) E. simplicicollis male; 13) E. mithroides male; 14) E. vesiculosa male; 15) E. peruviana female; 16) E. vesiculosa female, 17) E. attala male. Abbreviations indicate the following characters, S 8 -S 9: Abdominal segments 8-9; LV: Vulvar lamina; Pb: Posterior border; Vt: Teeth ventral; Ai: Inferior appendage; As: Antehumeral stripe.

Figures 18–25. Discrete characters of vesica spermalis. Numbers represent the following species:18) E. mithroides, dorsal view; 19) E. carmelita, left side view; 20) E. vesiculosa, left side view; 21) E. peruviana, left side view; 22) E. credula, dorsal wiew; 23) E. haematogastra, right side view; 24) E. credula, finger-shaped structures, dorsal view; 25) E. plebeja, dorsal view. Abbreviations indicate the following characters, C: Cornua; L: Lateral lobe; H: Hook; Pb: Posterior border; M: Medial lobe; P: Posterior lobe; NH: Hook not bilobed.