Erythemis simplicicollis (Say, 1840)

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 : 25-28

Erythemis simplicicollis (Say, 1840)

( Fig.1 , 2a–c , 5, 6, 9 , 12 , 26, 37)

Libellula simplicicollis Say, 1840: 28 .

Libellula imbuta Say, 1840: 32 .

Libellula caerulans Rambur, 1842: 64 .

Libellula maculiventris Rambur, 1842: 87 .

Mesothemis simplicicollis Hagen, 1861: 170

Mesothemis gundlachii Scudder, 1866: 195 .

Mesothemis simplicicollis Calvert, 1893: 265 (Diagnosis) .

Type material. Not examined.

Diagnosis. Thorax green, and dorsum of abdomen green with black dorsal bands (female and sexually immature male), or thorax and abdomen pruinose blue (mature male). McVey (1985) provides information about rates of color maturation in relation to age, diet, and temperature in males of this species. Abdominal appendages are white. Basal area hyaline. Posterior lobe of vesica spermalis absent. Hook is bilobed and not perpendicular to the longitudinal axis of the vesica spermalis ( Fig. 18 ), and the lateral lobe is extended more posteriorly than the medial lobe. Cornua is parallel with respect to the transversal axis of the vesica spermalis, with lobes fused to apex, and not covered by lateral lobes in lateral view ( Fig. 20 ). Lab ≤ 30mm.

Morphometric ratios of males. Acd/Lban<4.10; And/Lban<4.10; Lban/Anb<1.90; LHW/Acd<3.60; LHW/LFW<0.97; Las/Ddas>4.00; Las/Anas<4.00; LclS3/LcaS3<2.10; LHW/Lab>1.20; Lecv-clS3/ LcaS3<1.50; Lab/LclS3<1.50; Lab/LcaS3<3.00; Antr/Lar<8.00; LFW/Lnpt>3.00; Lnpt/Anb<1.50; Lba/ Lban>2.00; Lban/Lasa-ca<3.00.

Morphometric ratios of females. AnptFW/LoptFW<0.22; Lban/Anb<1.55; LHW/Acd<0.54; 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; Antr/Lar<3.00; LFW/Lnpt<2.90; Lar/Lba<0.14; Lnpt/Anb>1.20.

Head. Ll: 0.0132 –0.0137 mm (M); 0.012 –0.013 mm (F). Lw: 0.033 –0.038 mm (M); 0.031 –0.034 mm (F). Hwd: 0.057 –0.065 mm (M); 0.058 –0.063 mm (F). Hl: 0.030 –0.038 mm (M); 0.032 –0.034 mm (F).

Legs. Nsmf: 8–12 (M); 7–9 (F). Nshf: 18–30 (M); 10–18 (F). Hfl: 0.060 –0.070 mm (M); 0.056 –0.067 mm (F). Fbsd: 0.022 –0.024 mm (M); 0.021 –0.027 mm (F). Mfl: 0.038 –0.043 mm (M); 0.034 –0.041 mm (F). Dshf (the conditions separated by commas are present in different specimens): short spines-1 or 2 median spines-3 long spines, short spines-3 long spines.

Wings. LoptFW: 0.030 –0.034 mm (M); 0.034 –0.035 mm (F). AnptFW: 0.005 –0.007 mm (M); 0.006 –0.007 mm (F). LoptHW: 0.031 –0.035 mm (M); 0.036mm (F). AnptHW: 0.005 –0.007 mm (M); 0.0062 –0.0068 mm (F). DfwHW: 0.057 –0.062 mm (M); 0.057 –0.063 (F). LbanFW: 14.99–16.24mm (M); 15.75–16.20mm (F). LbanHW: 12.73–13.75mm (M); 12.06–13.53mm (F). AndFW: 6.86–7.83mm (M); 8.28–8.88mm (F). Wb: 8.52–9.60mm (M); 8.93–9.60mm (F). LFW: 30.00– 34.10mm (M); 33.38–34.54mm (F). LHW: 29.27–33.73mm (M); 31.00– 32.76mm (F). AnsbtFW: 0.018 –0.024 mm (M); 0.020 –0.021 mm (F). AntrFW: 0.005 –0.009 mm (M); 0.006 –0.008 mm (F). LarFW: 0.003 –0.009 mm (M); 0.0043 –0.0056 mm (F). LarHW: 0.0012 –0.0025 mm (M); 0.0006 –0.0018 mm (F). LnptFW: 10.08–12.20mm (M); 11.12–12.28mm (F). LnptHW: 10.91–13.36mm (M); 12.22–13.31mm (F). At-MP: 0.0031 –0.0043 mm (M); 0.003 –0.013 mm (F). LbaFW: 0.030 –0.032 mm (M); 0.028 –0.032 mm (F). LbaHW: 0.033 – 0.033 mm (M); 0.030 –0.034 mm (F). RP2-loptFW: 0.000 – 0.002 mm (M); 0.006 –0.012 mm (F). RP2-loptHW: 0.0000 – 0.0018 mm (M); 0.0018 – 0.0056 mm (F). Dat-lsbt: 0.0012 –0.0018 mm (M); 0.0012 –0.0018 mm (F). Lasa-ca: 0.0068 –0.0112 mm (M); 0.008 –0.012 mm (F). Crp: 0.008 –0.011 mm. Crp: 5–8 (FW), 4–7 (HW). Cal: 3–6. pC-RA-RP1: 2–3. CdfFW: only 3 rows of cells. CdfHW: 1–2 individual cells-2 to 4 double cells. Vt: 1. Csub: 3.

Abdomen. Lab: 25.55–29.80mm (M); 25.16–30.61mm (F). LcaS3: 0.010 –0.011 mm (M); 0.0100 – 0.0106 mm (F). LclS3: 0.020 – 0.025 mm (M); 0.021 –0.028 mm (F). Lecv-clS3: 0.013 –0.016 mm (M); 0.012 –0.014 mm (F).

Caudal appendages. Las: 0.015 –0.018 mm (M); 0.010 –0.011 mm (F). Poas: 0.0018 –0.0037 mm (M); 0.0018mm (F). Anas: 0.004 –0.005 mm (M); 0.003 –0.004 mm (F). Ddas: 0.0025 –0.0043 mm (M). Epl: 0.009 –0.013 mm (M); 0.004 –0.005 mm (F).

Larva. Bick (1941) mentions 13 larval instars, a total larval period of 113 days, and an average hatching time as 11.6 days. The small and robust larvae of Erythemis simplicicollis are habitat generalists. They can inhabit swamps, marshes, ponds, lakes, reservoirs and small pools of creeks where they live under the detritus, feeding on tadpoles ( May and Baird 2002; Relyea 2003). Osburn (1906) listed E. simplicicollis larvae living in “slightly salt water”, Wright (1943) reported E. simplicicollis larvae in Mississippi River delta and central Gulf Coast salt marshes in brackish areas of varying salinities (14–57% seawater), Bick et al. (1953) found E. simplicicollis larvae associated with naturally acid streams in southern Louisiana, and Smith and Smith (1995) exposed larvae to six salinities (0–80% seawater) and found that larvae survived for at least 24 h.However, survival time and eclosion significantly decreased at salinities above 20% with only 50% of the larvae emerging at that those higher concentrations (Smith and Smith 1995). Also, little development occurred at salinities of 60% and 80%, suggesting that increasing salt concentration could produce physiological stress on the larvae. Catling (2005) found E. simplicicollis as indicator of higher water quality. Both temperature and time of day influence the occurrences and frequencies of ventilation in larvae of this species, under constant temperature conditions, the vertical migration has its highest peak during photophase and its lowest during dark phase ( Cofrancesco and Howell 1982). According to Morin (1984), the larvae are eaten by fish, compete for food with larvae of Tramea lacerata Hagen (Libellulidae) (Wissinger and McGrady 1993). Larger larvae of E. simplicicollis attack and predate on Hyalella (Crustacea) amphipods ( Cothran 2008).

Material examined. HAITI, Samana, Frazar , no data, 1 male . MEXICO, Jalisco, La Huerta. Arroyo Chamela , 1.ix.2001, E. González-Soriano leg., 1 male ( CNIN) ; Coahuila, Puente San Rodrigo II bridge, c. 64 km S of Ciudad Acuña , elev. c. 28°44’01.0” 100°54’45.8”, 320 m, 23.vi.2007, R.A. Behrstock leg., 2 females ( CNIN) . U.S.A., Nebraska, Cherry Co., Boardman Creek at Rd. 16C, Merritt Reservoir State Recreation Area , W of Nebr. Hwy 97, about 26 mi SW of Valentine, 42°34’57” 100°54’56”, 18.vii.1998, R. W. Garrison leg., 1 male ( RWG) ; California, Cumberland Co., Texas Pond by Honeycutt Road , Fort Bragg Military Reservation , 35°8’34” 78°55’56”, 53 m, 16.vii.1967, R. W. Garrison leg., 1 male ( RWG) ; Texas, Williamson Co., Mustang Creek, by Carlos G. Parker Blvd. (= Loop 427), Taylor , 30°34’37” 97°27’18”, 25.viii.1976, J. E. Hafernik, Jr. leg., 1 male ( RWG) ; same, but Field near road to Manor, Taylor , 30°34’37” 97°27’18”, 16.viii.1975, J. E. Hafernik, Jr. leg., 1 female ( RWG) .

Remarks. The paratypes of E. simplicicollis were included in the Say’s paratypes collection, but most of his collection is known to have been lost.

Biology. Erythemis simplicicollis adults has been sigth in swarms (Paulson 1966), trapped from light traps (Wright 1944; Frost 1975), and Harrison and Lighton (1998) found that, under laboratory conditions, they are even able to fly with low concentrations of oxygen. E. simplicicollis may to have one, two or several generations per year depending of the climatic conditions ( Montgomery 1980; Harrison and Lighton 1998). This species usually prefer to perch on exposed logs or rocks (Robey 1975) and their sexual recognition is based on visual cues of body coloration ( Andrew 1966). Males are territorial, exhibiting intraspecific flight pattern when two males are in the same area ( Currie 1963), and showing interspecific aggression toward males of Pachydiplax longipennis (Burmeister) ( Baird and May 2003) . McVey (1988) found that lifetime reproductive success was highly variable, suggesting that this variation is important for selection between both sexes, especially when males are territorial ( Koenig and Albano 1987). After mating (which is not preceded by courtship behavior), the male tries to prevent other males from mating with his female or female may mate with different males in one day and store sperm to fertilize their eggs for several days, however, McVey and Smittle (1984) found that the last male in copulate fertilizes the up to 95% of the eggs.

According to collection labels, adults of E. simplicicollis feeds on females of the same species, as well as other Libellulidae (Odonata) such as Celithemis ornata (Rambur) , Perithemis tenera (Say) , and Orthemis ferruginea (Fabricius) , Coenagrionidae (Odonata) such as Enallagma weewa Byers , and Araneae (Arachnida) such as Leucauge argyra (Walckenaer) . E. simplicicollis also hunts Diceroprocta delicata (Osborn) ( Hemiptera : Cicadidae ) (Sanborn 1996), caterpillars of the noctuid moth Helicoverpa zea (Boddie) ( Bell and Whitcomb 1961) and Phidippus pulcherrimus Keyserling ( Araneae : Salticidae ) ( Edwards 1980; 1987). In New York ( USA), Olberg et al. (2000) found that 97% of prey-capture flights by E. simplicicollis males ended in successful captures. On the other hand, some birds ( Calvert 1893), fish and Asilidae (Diptera) such as Promachus hinei Bromley ( Chatfield 2011) may feed on E. simplicicollis . Adults of E. simplicicollis are parasitized by Actinocephalus (Koern.) and Geneiorhynchus Schneider (Eugregarinorida: Actinocephalidae ) ( Locklin and Vodopich 2010), the presence and abundance of these parasites appear to be independent of season and do not affect the host fitness characters ( Locklin and Vodopich 2009; 2010), but in central Texas ( USA), the same authors found that parasite prevalence was biased towards males because of behavioral and environmental aspects that influenced them. Painter et al. (1996) found that repeated applications of Bacillus thuringiensis israelensis on E. simplicicollis , did not affect development, morphology or their flight capability. Recently, a novel adipokinetic hormone (neuropeptide) of corpora cardiaca ( Gäde and Kellner 1999; Gäde et al. 2011), and one Cyclovirus new species, were found in E. simplicicollis adults (Rosario et al. 2012).

Distribution. From U.S.A. to Costa Rica ( Fig. 37 ), from 0 to 1825 m. asl.

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 37. Erythemis simplicicollis distribution map.

Figures 4–9. Morphometric characters. 4) S1-3 of E. carmelita male, 5-6) E. simplicicollis male; 7) E. peruviana female; 8) FW of E. vesiculosa male; 9) E. simplicicollis male. Abbreviations indicate the following characters, LFW, LHW: FW or HW length; Lnpt: Nodus-pterostigma length; Antr: Triangle width; Lar: Arculus-second antenodal length; Lba: wing base-arculus length; Lban: Wing base-nodus length; Anpt: Pterostigma width; Lopt: Pterostigma length; Ansbt: Subtriangle width; Acd: HW discoidal field posterior border width; Anb: HW base width; Lasa-ca: Supplementary anal vein-Cu-A crossvein, Lab: Abdomen length; Las: Cercus length; Ddas: reach of teeth on ventral region of cercus; Anas: Anterolateral width of cercus; S: Abdominal segment 3; LclS3: Lateral carina length; LcaS3: 3 Apical carina length; Lecv: Ventral carina length; Lecv-clS3: Basal area between ventral-lateral carinae on S3; Wm: Width medial region of the abdominal segment; S 8: Abdominal segment 8; S 9: Abdominal segment 9; Lal: Female lamina length from base of basal lobe to apex.

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.