Superfamily Coenagrionoidea

Chapter 14.6

Superfamily Coenagrionoidea Laboratorio de Biodiversidad y Gene´tica Ambiental (BioGeA), Universidad Nacional de Avellaneda (UNDAV), Buenos Aires, Buenos Aires, Argentina

Javier Muzo´n Laboratorio de Biodiversidad y Gene´tica Ambiental (BioGeA), Universidad Nacional de Avellaneda (UNDAV), Buenos Aires, Buenos Aires, Argentina

Danielle Anjos-Santos Centro de Investigaciones Esquel de Montan˜a y Estepa Patago´nicas (CIEMEP), Esquel, Chubut, Argentina

Pablo Pessacq Centro de Investigaciones Esquel de Montan˜a y Estepa Patago´nicas (CIEMEP), Esquel, Chubut, Argentina

Chapter Outline Introduction Limitations Terminology and Morphology Material Preparation and Preservation

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INTRODUCTION Coenagrionoidea, with approximately 170 genera and more than 1,700 species, is the largest superfamily within Zygoptera (Dijkstra et al., 2014). Several hypotheses have been proposed to explain its internal arrangement (O’Grady & May, 2003; Rehn, 2003; Carle et al., 2008), and the most recent molecular studies and classificatory scheme (Dijkstra et al., 2013, 2014) suggested that it is composed of three families: Isostictidae, Platycnemididae, and Coenagrionidae. Only the Coenagrionidae are represented within the Neotropical region. The Isostictidae, with 12 genera and 46 species is restricted to Australia, New Guinea, New Caledonia, and neighboring islands; and the Platycnemididae, with 40 genera and more than 400 species, is confined to the Old World. Coenagrionidae is composed of small to large-sized, and usually colorful damselflies commonly known in Spanish as “caballitos del diablo” or “alguaciles.” This is the most diverse family of Zygoptera; with 114 genera and 1,267 species, it represents 73% of all Coenagrionoidea (Dijkstra et al., 2014). Knowledge of the larvae is still very poor; to date, only 200 larvae have been described, which represents about 30% of the total known diversity. This

Key to Coenagrionidae Genera Acknowledgments References

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number includes representatives of just 34 genera, approximately 50% of the generic diversity. The internal topology of Coenagrionidae has long been discussed (see O’Grady & May, 2003). Recent studies suggest that two major groups are recovered, and according to Dijkstra et al. (2013) each group could be treated as a subfamily or even family. The Coenagrionidae sensu Dijkstra et al. (2014) implies the nonmonophyly of longrecognized families within Zygoptera. This has changed the macrosystematics of the order, especially within the Neotropical region where members of the Protoneuridae and Pseudostigmatidae are now included within Coenagrionidae (see Table 14.1). In this chapter, we follow the proposal of Dijkstra et al. (2014). Members of this family are common inhabitants of all kinds of freshwater environments, such as shallow vegetated ponds and shaded streams (von Ellenrieder & Garrison, 2007; Muzo´n et al., 2010). Corbet (1999) included among the habitats of Coenagrionidae fast streams with boulders, slow moving streams with detritus, rivers, and permanent lotic environments. Additionally, larvae of all members of the former Pseudostigmatidae and a few other Coenagrionidae (e.g., Bromeliagrion, Leptagrion, Roppaneura)

Thorp and Covich’s Freshwater Invertebrates, Fourth Edition. https://doi.org/10.1016/B978-0-12-804223-6.00020-2 Copyright Ó 2018 Elsevier Inc. All rights reserved.

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inhabit phytotelmata (tree holes, bamboo internodes, and bromeliads, see Corbet, 1983; Fincke, 1984, 2005; Neiss et al., 2008) in Neotropical forests, and many other Neotropical species are found in temporary ponds. Most Coenagrionoidea larvae live among plants, e.g., submerged weeds, reeds, or grass-like vegetation, and some others may attach themselves to the roots of trees or live among coarse detritus (Suhling & Martens, 2007). The Coenagrionidae in the Neotropics is represented by more than 650 species within 70 genera, of which 60 are endemic or mainly distributed in the Neotropical region (Garrison et al., 2010; Dijkstra et al., 2013). This number is continuously increasing; for example, in the last decade 121 new species and 11 new genera have been described. There are eight genera with more than 20 species each in the region: Acanthagrion, Argia, Epipleoneura, Neoneura, Metaleptobasis, Oxyagrion, Protoneura, and Telebasis. There are 17 monotypic genera (with only one species) in the region: Anomisma, Apanisagrion, Dolonagrion, Enacantha, Fluminagrion, Hesperagrion, Hylaeonympha, Junix, Lamproneura, Leucobasis, Megaloprepus, Proneura, Roppaneura, Schistolobos, Telagrion, Tigriagrion, and Tukanobasis. The remainder 45 genera have intermediary number of species. In the Neotropics, five countries have more than 100 species recorded: Brazil (268), Venezuela (132), Peru (129), Mexico (111), and Colombia (106) (Paulson, 2014, 2016a, 2016b). This coincides with global diversity patterns. Diversity decreases southwards, with Cyanallagma interruptum (Selys) being the southernmost species of Coenagrionidae; it was recorded from Santa Cruz province, Argentina, in Patagonia (Muzo´n et al., 2014).

LIMITATIONS Knowledge of the Coenagrionidae is still very incomplete and many larval descriptions need to be updated. There are several keys for the adults that include Coenagrionoidea (e.g., Lencioni, 2005, 2006; Garrison et al., 2010), but there are a few and only local keys for the larvae (von Ellenrieder & Garrison, 2007; Muzo´n et al., 2014; Neiss & Hamada, 2014) which makes larval identification very difficult.

Despite being the most diverse group within Zygoptera, larval morphology is strikingly uniform, which makes identification difficult, especially at the species level. Larval identification in most cases requires knowledge of either the local fauna from a thorough sampling of adults, or the collection and rearing of larvae to the adult. Odonata larvae are described based on the morphology of the final instar (F0). It has been documented that morphological changes in structures used for identification occur during development (Corbet, 1995). This makes identification of younger instars almost impossible, especially within Coenagrionidae where the general morphology is pretty much uniform within the family.

TERMINOLOGY AND MORPHOLOGY The most important terminology for identifying these damselflies are described in Chapter 14.0 in this book and Chapter 35 (Suhling et al., 2015) in Volume 1 of this series (Thorp & Rogers, 2015). Some of the main morphological features of the family include: head without tubercles; antennae with first article short; prementum either triangular or subrectangular with a variable number of premental setae (often the n þ n notation is used to indicate number of setae on each side of the midline) and with ligula entire, with a variable number of small denticles; labial palps stout with a variable number of palpal setae, a strong and recurved movable hook, and a complex series of distal teeth; the mandibles have generally 4 or 5 incisors with the presence of an accessory tooth y and 2 or 3 molars (the notation of mandibular formula used to describe it follows Watson, 1956). The caudal appendages are extremely variable in morphology, and are used for both locomotion and to aid respiration, especially when there is low concentration of oxygen (Sesterhenn et al., 2013).

MATERIAL PREPARATION AND PRESERVATION In Volume I (Thorp & Rogers, 2015) of this series specimen preparation and preservation are presented.

KEY TO COENAGRIONIDAE GENERA Our knowledge on Neotropical coenagrionid larvae is currently restricted to 34 genera (48.57%) and approximately 210 species (31.81%). Larvae of this family share a trapezoidal labium which generally reaches coxae 2, a slender body with three lamellated caudal appendages (epiproct þ paraprocts), held more or less in the vertical plane. Caudal appendages shape, presence/absence of nodus and spines, and number of premental and palpal setae, are the most commonly used characters to distinguish among genera and species, since they are extremely variable, and are widely used in the keys. Genera previously assigned to Protoneuridae and Pseudostigmatidae are now included within Coenagrionidae (see Dijkstra et al., 2014). Unfortunately, this could be confusing for nonspecialists when tracking references. Therefore, the major changes, which include the inclusion of genera previously assigned to Protoneuridae and Pseudostigmatidae within Coenagrionidae, are summarized in Table 13.6.1. Distributional data for each genus in Neotropical region is indicated between brackets (based on Garrison et al., 2010). 1 1’ 2(1) 2’

Premental setae absent (Fig. 14.6.1) .............................................................................................................................................................. 2 Premental setae present (Fig. 14.6.2)............................................................................................................................................................. 7 Caudal appendages stalked and foliate, widening abruptly (Fig. 14.6.3); larvae inhabit phytotelmata ...................................................... 3 Caudal appendages generally not as above, usually laminar and ending in a filamentous tip (Fig. 14.6.4); if stalked and foliate then larvae not in phytotelmata, usually in lotic environments ................................................................................................................................ Argia [Neotropics. Nearctic]

Superfamily Coenagrionoidea

FIGURE 14.6.1 Argia serva Hagen in Selys, 1865, labium, dorsal view.

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FIGURE 14.6.2 Idioneura ancilla Selys, 1860, labium, dorsal view. Arrows indicate premental setae.

FIGURE 14.6.3 Mecistogaster sp., caudal appendages, lateral view. Original pictures by Neiss & Hamada (2014).

3(2) 3’ 4(3) 4’ 5(4) 5’ 6(5) 6’

Labial palp with five or more setae (Fig. 14.6.5) .......................................................................................................................................... 4 Labial palp with two setae (Fig. 14.6.6) ....................................................................................................................................... Platystigma [Neotropics] Antennomere three length <2 second (Fig. 14.6.7) ................................................................................................................................... 5 Antennomere three length >2 second (Fig. 14.6.8) ................................................................................................................. Microstigma [Neotropics. Nearctic] Caudal appendages apically with a pale area (Figs. 14.6.9, 10) ................................................................................................................... 6 Caudal appendages apically without pale area (Fig. 14.6.3).................................................................................................... Mecistogaster [Neotropics] Caudal appendages strongly petiolate (Fig. 14.6.9) ................................................................................................................. Megaloprepus [South central and Northeast South America] Caudal appendages weakly petiolate (Fig. 14.6.10) ................................................................................................................. Pseudostigma [Central America, Colombia]

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FIGURE 14.6.4 Argia sp., caudal appendages, lateral view.

SUPERFAMILY COENAGRIONOIDEA FIGURE 14.6.5 Mecistogaster amalia (Burmeister, 1839), labium, dorsal view. Illustration adapted from Muzo´n et al. (2010).

FIGURE 14.6.6 Platystigma astictum Machado & Lacerda, 2017, labial palp, dorsal view. Illustration adapted from Lencioni (2006).

FIGURE 14.6.7 Mecistogaster amalia, antenna, dorsal view. Numbers indicates antennomeres. Illustration adapted from Muzo´n et al. (2010).

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FIGURE 14.6.10 Pseudostigma aberrans Selys, 1860, caudal appendages, lateral view. Bar indicates petiole length; arrows indicate tips of caudal appendages with a pale area. Illustration adapted from NoveloGutie´rrez (1993).

FIGURE 14.6.8 Microstigma rotundatum Selys, 1860, antenna, dorsal view. Numbers indicates antennomeres. Illustration adapted from De Marmels (2007).

FIGURE 14.6.11 Bromeliagrion rehni Garrison in De Marmels & Garrison, 2005, labial palp, dorsal view. Arrows indicate palpal teeth. Original pictures by Neiss & Hamada (2014).

FIGURE 14.6.12 Oxyagrion sp., labial palp, dorsal view. Arrows indicate palpal teeth. Original pictures by Neiss & Hamada (2014). FIGURE 14.6.9 Megaloprepus caerulatus (Drury, 1782), caudal appendages, lateral view. Bar indicates petiole length; arrow indicate tip of caudal appendage with a pale area. Illustration adapted from Hedstro¨m & Sahle´n (2003).

7(1) 7’ 8(7) 8’ 9(8) 9’

Labial palp distal margin with two teeth (Fig. 14.6.11)................................................................................................................................ 8 Labial palp distal margin smooth or with numerous small teeth and usually one larger tooth (Fig. 14.6.12).......................................... 13 Caudal appendages uniform in color, if color pattern is visible then without numerous dark spots that are larger near the margins (Fig. 14.6.13); premental setae variable......................................................................................................................................................... 9 Caudal appendages with numerous dark spots, larger near the margins (Fig. 14.6.14); 3 þ 3 premental setae..................... Diceratobasis [Dominican Republic, Jamaica] Labial palp with 10 or more setae (Fig. 14.6.11) ........................................................................................................................................ 10 Labial palp with eight or nine setae (Fig. 14.6.15) ......................................................................................................... Leptagrion (in part) [Brazil, Ecuador, Guianas, Peru, Surinam, Venezuela]

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FIGURE 14.6.13 Leptagrion capixabae Santos, 1965, caudal appendages, lateral view. Illustration adapted from Costa et al. (2009).

SUPERFAMILY COENAGRIONOIDEA

FIGURE 14.6.14 Diceratobasis macrogaster (Selys in Sagra, 1857), caudal appendages, lateral view. Illustration adapted from Westfall (1976).

FIGURE 14.6.15 Leptagrion dardanoi Santos, 1968, labial palp, dorsal view. Illustration adapted from Costa et al. (2009).

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FIGURE 14.6.16 Leptagrion andromache Hagen in Selys, 1876, labium, dorsal view. Illustration adapted from Muzon et al. (2009).

FIGURE 14.6.17 Bromeliagrion rehni, labium, dorsal view. Original pictures by Neiss & Hamada (2014).

FIGURE 14.6.18 Bromeliagrion fernandezianum (Ra´cenis, 1958), labial palp, dorsal view. Illustration adapted from De Marmels (1985).

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FIGURE 14.6.19 Leptagrion andromache, labial palp, dorsal view. Illustration adapted from Muzo´n et al. (2009).

FIGURE 14.6.20 Bromeliagrion fernandezianum, caudal appendages, lateral view. Illustration adapted from De Marmels (1985).

FIGURE 14.6.21 Oxyagron ablutum (Calvert, 1909), caudal appendages, lateral view in relation to abdomen length. Illustration adapted from von Ellenrieder & Lozano (2008).

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FIGURE 14.6.22 Acanthagrion aepiolum Tennessen, 2004, caudal appendages, lateral view in relation to abdomen length. Illustration adapted from von Ellenrieder & Lozano (2008).

FIGURE 14.6.23 Amphiagrion abbreviatum (Selys, 1876), head, dorsal view. Illustration adapted from von Cook & Antonelli (1969).

FIGURE 14.6.25 Roppaneura beckeri Santos, 1966, labial palp, dorsal view, dorsal view.

FIGURE 14.6.24 Idioneura sp., head, dorsal view.

FIGURE 14.6.26 Telebasis obsoleta (Selys, 1876), caudal appendages, lateral view.

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FIGURE 14.6.27 Psaironeura sp.,caudal appendages, lateral view.

SUPERFAMILY COENAGRIONOIDEA FIGURE 14.6.28 Peristicta forceps Hagen in Selys, 1860, labial palp, dorsal view. Illustration adapted from Pessacq (2007).

10(9) 10’ 11(10) 11’ 12(11) 12’ 13(7) 13’ 14(13) 14’ 15(14) 15’ 16(15) 16’

FIGURE 14.6.29 Peristicta forceps, caudal appendages, lateral view.

Prementum with 1 þ 1 primary setae (Fig. 14.6.16) .................................................................................................................................. 11 Prementum with 3 þ 3 primary setae (Fig. 14.6.17) ................................................................................................ Bromeliagrion (in part) Teeth proximal to movable hook truncate (Fig. 14.6.18)............................................................................................................................ 12 Teeth proximal to movable hook pointed (Fig. 14.6.19)................................................................................................. Leptagrion (in part) [Brazil] Caudal appendages without transverse suture ................................................................................................................. Leptagrion (in part) [Brazil] Caudal appendages with transverse suture (Fig. 14.6.20) ......................................................................................... Bromeliagrion (in part) Caudal appendages shorter than abdomen (Fig. 14.6.21) ........................................................................................................................... 14 Caudal appendages as long as or longer than abdomen (Fig. 14.6.22) ............................................ Acanthagrion (except for A. fluviatile) [Neotropics] Postocular lobes rounded (Fig. 14.6.24) ...................................................................................................................................................... 15 Postocular lobes angulate (Fig. 14.6.23)..................................................................................................................................... Amphiagrion [Mexico. Nearctic] Labial palp distal margin with end hook but without defined smaller external teeth (Figs. 14.6.25, 30, 31)........................................... 16 Labial palp distal margin with end hook and defined smaller external teeth (Fig. 14.6.12)...................................................................... 26 Caudal appendages length  3 width (Figs. 14.6.26, 29) ........................................................................................................................ 17 Caudal appendages length  4 width (Fig. 14.6.27)................................................................................................................................ 19

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FIGURE 14.6.30 Psaironeura sp., labial palp, dorsal view. Original pictures by Neiss & Hamada (2014).

FIGURE 14.6.31 Nehalennia minuta (Selys in Sagra 1857), labial palp, dorsal view. Original pictures by Neiss & Hamada (2014).

FIGURE 14.6.32 Psaironeura tenuissima (Selys 1886), femur, dorsal view. Original pictures by Neiss & Hamada (2014).

FIGURE 14.6.33 Forcepsioneura sancta (Hagen in Selys 1860), labial palp, dorsal view.

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FIGURE 14.6.34 Idioneura ancilla Selys, 1860, caudal appendages, lateral view.

SUPERFAMILY COENAGRIONOIDEA FIGURE 14.6.35 Neoneura sp., labium, dorsal view.

FIGURE 14.6.36 Epipleoneura manauensis Santos, 1964, caudal appendages, lateral view.

17(16) 17’ 18(17) 18’ 19(16) 19’

Labial palp with five to seven setae (Fig. 14.6.28) ..................................................................................................................................... 18 Labial palp with eight or nine setae (Fig. 14.6.25), larva lives in axils of Umbelliferae .......................................................... Roppaneura [Brazil] Caudal appendages with defined nodus (Fig. 14.6.26)..................................................................................................................... Telebasis [Neotropics. Nearctic] Caudal appendages with obscured nodus (Fig. 14.6.29) .................................................................................................................. Peristicta [Argentina, Brazil, Paraguay, Uruguay] Labial palp with three to six setae (Fig. 14.6.30)........................................................................................................................................ 20 Labial palp with seven setae (Fig. 14.6.31) .................................................................................................................................. Nehalennia [Neotropics. Nearctic]

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FIGURE 14.6.37 Neoneura sp., caudal appendages, lateral view.

FIGURE 14.6.38 Forcepsioneura sancta, caudal appendages, lateral view.

20(19) 20’

21(20) 21’

22(21) 22’

FIGURE 14.6.39 Epipleoneura manauensis, labial palp, dorsal view. Original pictures by Neiss & Hamada (2014).

Occiput, head mid-dorsal region, femora, and abdomen with filiform or spiniform setae; labial palp usually with three to five setae (Fig. 14.6.33), five to six in Idioneura (Fig. 14.6.2) ................................................................................................................................... 21 Occiput, head mid-dorsal region, femora, and distal margin of S6e8 with concave spiniform setae (Fig. 14.6.32); labial palp with six setae (Fig. 14.6.30) ................................................................................................................................................................................ Psaironeura [Central America, Northern South America] Prementum dorsally with one (Fig. 14.6.35) or two pairs of setae; labial palp with three to five (Fig. 14.6.33) setae; caudal appendages not as below ........................................................................................................................................................................................................ 22 Prementum dorsal surface with three pairs of setae, although third pair may be reduced (Fig. 14.6.2); labial palp with five to six setae (Fig. 14.6.2); caudal appendages distinctly narrower basally, widened distally (Fig. 14.6.34) ..................................................... Idioneura [Brazil] Gills not divided (Fig. 14.6.36).................................................................................................................................................................... 23 Gills divided as a rigid sclerotized basal half and a membranous, paler distal half (Fig. 14.6.37)........................................................... 24

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FIGURE 14.6.40 Protoneura caligata (Hagen in Selys, 1886), caudal appendages, lateral view. Arrow indicates nodus position. Illustration adapted from Westfall (1964).

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FIGURE 14.6.41 Neoneura ethela Williamson, 1917, caudal appendages, lateral view. Arrow indicates nodus position. Illustration adapted from De Souza et al. (2012).

FIGURE 14.6.42 Mesamphiagrion tamaense (De Marmels, 1988), caudal appendages, lateral view. Illustration adapted from von Ellenrieder & Garrison (2008).

FIGURE 14.6.43 Mesamphiagrion laterale (Selys, 1876), S10 dorsal, view. Illustration adapted from De Marmels (2007).

FIGURE 14.6.44 Oreiallagma quadricolor (Ris, 1918), S10 dorsal, view. Illustration adapted from von Ellenrieder & Garrison (2008).

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FIGURE 14.6.48 Hesperagrion heterodoxum (Selys, 1868), caudal appendages, view. illustration adapted from Westfall & May (2006).

FIGURE 14.6.45 Mesamphiagrion laterale, labium, dorsal view. Illustration adapted from De Marmels (2007).

FIGURE 14.6.49 Leptobasis sp., labial palp, dorsal view. Illustration adapted from Trapero Quintana & Naranjo Lo´pez (2009).

FIGURE 14.6.46 Protallagma titicacae (Calvert, 1909), labium, dorsal view. Illustration adapted from Bulla (1972).

FIGURE 14.6.47 Homeoura chelifera (Selys, 1876), caudal appendages, lateral view.

490

23(22) 23’ 24(22) 24’

25(24)

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25’

26(15) 26’ 27(26) 27’ 28(27) 28’ 29(26) 29’ 30(29) 30’ 31(30) 31’ 32(31) 32’ 33(31) 33’

34(33) 34’ 35(34)

35’ 36(34) 36’ 37(36) 37’ 38(37) 38’

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Labial palp with five (Fig. 14.6.33) or six setae; lateral caudal appendage length > 7 width (Fig. 14.6.38) .................. Forcepsioneura [Brazil] Labial palp with three or four setae (Fig. 14.6.39); lateral caudal appendage length < 6 width (Fig. 14.6.36) ................. Epipleoneura [Neotropics] Labial palp with three or four (rarely five) setae; tibiae with one or no dark, transverse band; total length (excluding gills) more than 12 mm ....................................................................................................................................................................................................................... 25 Labial palp with five setae (except P. romanae, with six); tibiae with two or three dark, transverse bands; total length (excluding gills) less than 10 mm (except P. calligata, below) ....................................................................................................................................... Protoneura [Central America, Northern South America] Lateral caudal appendages nodus located at about distal 3/5 (Fig. 14.6.40); lateral caudal appendages rigid portion dorsal margin with about 20 spiniform setae; median caudal appendage ventral margin with about 30 spiniform setae.................................... Protoneura calligata [Cuba] Lateral caudal appendages nodus located at about 1/2, 1/3 or 2/3 of appendages (Fig. 14.6.41); lateral caudal appendages rigid portion dorsal margin with less than 15 spiniform setae (Fig. 14.6.41); median caudal appendage ventral margin with less than 20 spiniform setae ........................................................................................................................................................................................................... Neoneura [Neotropics] Caudal appendages ovoid, with at least one margin convex (Fig. 14.6.42) ............................................................................................... 27 Caudal appendages lanceolate, with margins subparallel at least along proximal third (Figs. 14.6.21, 47, 48)....................................... 29 S10 posterodorsal margin with an arcuate cleft; occipital lobes prominent (Fig. 14.6.43) ....................................................................... 28 S10 posterodorsal margin entire; occipital lobes not prominent (Fig. 14.6.44) ......................................................................... Oreiallagma [Venezuela to Bolivia] Palpal setae five or six; generally with 3 þ 3 or 4 þ 4 long premental setae (Fig. 14.6.45)............................................. Mesamphiagrion [Colombia, Ecuador, Venezuela] Palpal setae three or four; generally with 2 þ 2 long premental setae (Fig. 14.6.46) ................................................................ Protallagma [Argentina, Peru] Caudal appendage apex usually acute (Figs. 14.6.21, 47), if rounded then setae on basal half stouter and more closely spaced than those more distal .................................................................................................................................................................................................... 30 Caudal appendage apex rounded (Fig. 14.6.48), marginal setae subequal in size and spacing along at least 9/10 of the gills Hesperagrion [Me´xico. Nearctic] Labial palp distal margin without bifid teeth (Fig. 14.6.12) ....................................................................................................................... 31 Labial palp distal margin with bifid teeth (Fig. 14.6.49) ............................................................................................................... Leptobasis [Central America, Northern South America, Greater Antilles. Nearctic] Lateral caudal appendage apical sixth with a terminal angle of 70o or more (Fig. 14.6.50) .................................................................... 32 Lateral caudal appendage apical sixth with a terminal angle of 60o or less (Figs. 14.6.21, 47)............................................................... 33 Labium with four premental setae (if five, one or two reduced) and five palpal setae (Fig. 14.6.51) ........................................ Anisagrion [Central America, Ecuador, Venezuela] Labium with five to eight premental setae and six to eight palpal setae (Fig. 14.6.52) .......................................................... Apanisagrion [Me´xico to Honduras. Nearctic] Prementum with two or more setae on each side of midline (Fig. 14.6.53) .............................................................................................. 34 Prementum with one long seta on each side of midline (Fig. 14.6.54), may be accompanied by several short setae .................................. ........................................................................................................................................................................................... Enallagma (in part) [Neotropics. Nearctic] S2 through S8 with lateral carinae slightly concave in dorsal view and with prominent apices bearing two or more stout, curved setae .. ....................................................................................................................................................................................................................... 35 Abdominal segments with lateral carinae straight or slightly convex in dorsal view, without prominent apices and with setae, if present, no larger than the adjacent ones on the same segment .................................................................................................................................... 36 S9 with lateral carinae less prominent than those of preceding segments, and without stout setae; usually four palpal setae..................... ........................................................................................................................................................................................... Enallagma (in part) [Neotropics. Nearctic] S9 with lateral carinae nearly as prominent as those of S8, and carinae of S8 and 9 both bearing stout setae; usually five palpal setae ... ................................................................................................................................................................................................. Neoerythromma Total length of larvae excluding caudal appendages >9 mm ..................................................................................................................... 37 Total length of larvae excluding caudal appendages <6 mm ................................................................................................... Argentagrion [Argentina, Bolivia, Brazil, Paraguay, Uruguay] Eyes lacking pattern of lateral bands, but sometimes with dark spots ....................................................................................................... 38 Eyes usually with a pattern of alternating light and dark lateral bands............................................................................................ Ischnura [Neotropics. Nearctic] Species distributed in Central America......................................................................................................................................................... 39 Species distributed in South America .......................................................................................................................................................... 40

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FIGURE 14.6.50 Apanisagrion lais (Brauer in Selys, 1876), caudal appendages, lateral view. Illustration adapted from Novelo Gutie´rrez (2010).

FIGURE 14.6.51 Anisagrion inornatum Selys, 1876, labium, dorsal view. Illustration adapted from Tennessen (2012).

FIGURE 14.6.52 Apanisagrion lais, larval labium, dorsal view. Illustration adapted from Novelo Gutie´rrez (2010).

39(38)

39’ 40(38)

40’

Lateral carinae and general surface of abdomen (except dorsum of S9e10) without stout setae; antennal article III length 3x article I ... .......................................................................................................................................................................................................... Enacantha [Belize, Cuba, Guatemala, Mexico] Lateral carinae and general surface of abdomen with stout setae; antennal article III length < 3 article I .............. Enallagma (in part) [Argentina, Bolivia, Brazil, Ecuador, Peru, Venezuela. Nearctic] Labial palp medial margin crenulate (Fig. 14.6.55) (there are no generic characters which can be used to separate these genera; knowledge of local fauna is needed in order to be able to separate them) .............................................................. Oxyagrion/Andinagrion/Homeoura [South America]/[Argentina and Uruguay]/[South America] Labial palp medial margin not crenulate ................................................................................................................................... Cyanallagma [Argentina, Brazil, Chile, Uruguay]

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FIGURE 14.6.53 Ischnura fluviatilis Selys, 1876, labium, dorsal view.

FIGURE 14.6.54 Enallagma novahispaniae Calvert, 1907, labium, dorsal view. Illustration adapted from Novelo Gutierrez (2005). FIGURE 14.6.55 Oxyagrion rubidum (Rambur, 1842), labial palp, dorsal view.

Superfamily Coenagrionoidea

ACKNOWLEDGMENTS Figs. 14.6.3, 4, 14.6.12, 14.6.25e27, 14.6.30, 14.6.32, 14.6.36, 14.6.39 were originally presented by Neiss & Hamada (2014) and were generously given by Editora do Instituto Nacional de Pesquisas da Amazoˆnia, Manaus. We express our most sincere gratitude to Dr. Natalia von Ellenrieder for the loan of material.

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