The levis group of Orthemis revisited: a synopsis including a synonymy and description of six new species of Orthemis from South America (Odonata: Libellulidae)

The levis and ferruginea groups of Orthemis are redefined. Six new species of Orthemis are described: O. aciculata sp. nov. (♂ holotype: Surinam, Para Dist., road near forest, Zanderij I (5°32′ N, 55°10′ W), 17 January 1957, leg. J. Belle [RMNH]), O. celata sp. nov. (♂ holotype: Brazil, Pará State, Rio Gurupí, Canindé (0°30′57′ ′ S, 51°14′00′ ′ W), 27–28 February 1966, leg. B. Malkin [RMNH]), O. faaseni sp. nov. (♂ holotype: Brazil, Rondônia State, Porto Velho (8°46′ S, 63°54′ W), 28 February 1922, leg. J.H. Williamson & J.W. Strohm [UMMZ]), O. garrisoni sp. nov. (♂ holotype: Panama, Panamá Prov., 7 km NW of Gamboa, Pipeline Road, trail near palm forest, 30 July 1979, leg. RWG & J.A. Garrison [FSCA]), O. paulsoni sp. nov. (♂ holotype: Ecuador, Sucumbíos Prov., forest trail ca. 3 km W of Shushufindi (0°09′59′ ′ N, 76°41′30′ ′ W), 14 October 2009, leg. K.J. Tennessen [FSCA]), and O. teres sp. nov. (♂ holotype: Bolivia, Cochabamba Dept., Chapare, Cristal Mayo (17°00′57′ ′ N, 65°38′09′ ′ W), October/November 1994, leg. R. Andreas [FSCA]). Orthemis plaumanni Buchholz is found to be a junior synonym of O. ambinigra Calvert. A lectotype is designated for O. concolor Ris. All species of the levis group and the new species of the ferruginea group described here are diagnosed, illustrated, mapped, and keyed.


Introduction
Orthemis Hagen, 1861 currently comprises 23 New World species, primarily neotropical in distribution, which have been loosely classified into two groups based on abdominal width (von Ellenrieder, 2009): the ferruginea group, whose species are characterized by a robust and depressed abdomen, and the levis group, represented by species with a relatively slender abdomen.
Names assigned to species of the ferruginea group have been particularly difficult to associate with specimens, and a number of species included within this complex are not yet resolved (Donnelly, 1995(Donnelly, , 2001Paulson, 1998Paulson, , 2001. Species diagnoses in the levis group were originally based mostly on color (i.e. Calvert, 1902;Ris, 1910Ris, , 1919. Examination of structural characters of male genitalia showed several specimens with similar color patterns represented different species, leading recently to the description of five new species (von Ellenrieder, 2009).
After examining over 500 additional specimens from several collections, and borrowing type material of several species whose diagnoses based on original descriptions were not completely reliable, namely O. ambinigra, O. flavopicta, and O. plaumanni, I found O. ambinigra to be the senior subjective synonym of O. plaumanni, and discovered another five new species of the levis group and one of the ferruginea group which I describe in this paper. Among the examined material there were several pairs in copula, which allowed the hitherto unknown females of O. anthracina and O. coracina to be associated with their males, as well as those of O. cultriformis and of several of the new species. Based on correlation with male characters I also tentatively describe a female as O. harpago. I found the shape of the female vulvar lamina and associated structures to be diagnostic, allowing for species identification of isolated females which thus far was not always possible.
Here I redefine the ferruginea and levis groups of Orthemis, provide a synopsis of the levis group, including descriptions of five new species, diagnoses, illustrations, updated maps, an updated key to males, and a partial key to females based on the species described to date, and describe one new species in the ferruginea group.

Material and methods
Nomenclature follows Garrison et al. (2006). Measurements are given in millimeters; average dimensions are given as average ± standard deviation and were taken from 10 specimens when available; Pt length was measured along the costal side, cercus and epiproct length in lateral view, and epiproct maximum and subapical width in ventral view (as indicated in Figure 16a; total length includes appendages and abdomen length excludes them. Width/length ratio of S4 cannot be always measured with exactitude due to the variable degrees at which the abdomen can be artificially compressed (with opposite ventral terga touching, pleura folded under them and sterna entirely hidden) or dilated (with sterna, and sometimes pleura, fully exposed) during preservation; here S4 length was measured along the ventrolateral carina, and S4 apical width at level of the transverse posteroventral carina (as indicated in Figure 6a), and the widest possible range including compressed and dilated states was taken. Specimens in which genital diagnostic characters were not visible were softened in a wet chamber in order to allow spreading of male hamules, extrusion of vesica spermalis from the genital fossa, and unfolding of lateroventral flaps of female S8 using insect pins and watch forceps, then these structures were fixed in the desired position on a piece of balsa wood with minuten pins, and dehydrated in acetone again on the wood before removing the pins. Drawings were made with the aid of a camera lucida coupled to Wild M8 and Nikon SMZ1500 stereomicroscopes. Wings were scanned from specimens.
Short descriptive summaries and diagnoses are provided for species previously known and full descriptions for new species and hitherto unknown females. Species are treated alphabetically. Synonymic lists include only species descriptions, major treatments of the genus, and confirmed misidentifications in the literature, excluding isolated mentions and records. Several of the new species were erroneously assigned to some of the older names in the literature, therefore maps represent distribution records from material examined and type material, and do not include literature records unless confirmed by examination of specimens; they were created electronically from the Digital Chart of the World (1:1,000,000) using Arc View 9.3; longitude/latitude coordinates were culled from the Global Gazetteer website (http://www.fallingrain.com/world/) and Google Earth when not provided in the labels.
Acronyms for collectors and collections are as follows: ANSP -Academy of Natural Sciences, Philadelphia, Pennsylvania, USA; BMNH -The Natural History Museum, London, UK; CMNH -Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA; DRP -Dennis R. Paulson,18. Pterothorax dark reddish brown to black with yellow stripes in young adults (Figure 2k), to progressively obscured, with pterothorax entirely dark in mature males; Pt dark reddish brown to black; inner and outer branches of hamule separated by a distance longer than length of inner branch ( De Marmels, 1988;Donnelly, 2001;Meurgey & Daigle, 2007;Paulson, 2001;Ris, 1910) 4. Vulvar lamina consisting of posterior margin of sternum of S8 forming an inverted U-shaped rim (Figure 18d   14 . Abdomen narrowing abruptly at base, with base of S4 less than half as high as base of S3 in lateral view ( Figure 7l); transverse end of vulvar lamina more posterior, separated from posterior margin of S8 by a distance ca. equal to its width ( Figure 18o); Peru and Brazil ( Figure 22)

Etymology
This species is named aciculata (from the Latin noun acicula meaning splinter) in reference to the splinter-shaped sclerotized projection on medioectal lobe of distal segment of vesica spermalis.

Male holotype
Head. Prementum black; labial palps pale yellow with medial black stripe as wide as about 0.40 of palp width and anterior margin narrowly black (as in Figure 1b); labrum black with Abdomen. Sides linear, gradually narrowing from S3 to S4 in ventral view (Figure 6a), S4 about three times as long as wide (ratio apical width/length = 0.33). Dorsal terga ( Figure 4a): S1-2 reddish yellow, with pale yellow mid-dorsal stripe; S3-4 red, except posterodorsal black spot extending ventrally to lateral carina on S4; S5-7 black with red mid-dorsal and ventrolateral stripes, the latter narrowing from S5 to S7; S8 black with ventrolateral stripe and mid-dorsal carina red; S9 black with small diffuse red spot at ventrobasal third; S10 and caudal appendages dark reddish brown. Ventrolateral carina on S2 vestigial, on S3-8 well developed and black, on S9 well developed along basal two thirds, absent at distal third. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga ( Figure 6a): S1 yellow; S2 pale yellow along ventrolateral carinae, ventrally reddish brown; S3-8 dark reddish brown with pale yellow stripe along lateral margin; S9-10 dark reddish brown. Anterior lamina in lateral view shorter than hamule and as high as genital lobe ( Figure 10a); hamule bifid with small inner branch forming short pointed spine, and larger outer branch with blunt tip bent ventrally over inner branch, separated by a distance longer than inner branch length; outer corner of outer branch smoothly rounded ( Figure 11a). Distal segment of vesica spermalis with basal portion trapezoidal in ectal view (Figure 13a), and with long flagella on ental surface; distal lobes represented on each side by elongate outer oval shaped lobe longer than basal portion in lateral view, and medioectal membranous lobe with sclerotized lateral projection margined with denticles and distolateral sclerotized splinter-shaped projection (Figures 12a,13a). Cercus markedly curved ventrally in lateral view, with row of 4-6 ventral tubercles along distal 0.50 and tip upturned ( Figure 15a); in dorsal view converging along basal 0.70 with opposite cercus, then approximately parallel to it along distal 0.30 (Figure 14a). Epiproct

Variation in male paratypes
Head. As for holotype but labial palp medial black stripe as wide as 0.40-0.50 of palp width; in one paratype postclypeal lobes not fringed with dark line along distal margin; in one paratype labrum with pair of rounded laterobasal pale yellow spots.
Abdomen. As for holotype but dorsal terga of S1-2 orange red dorsally and S5 mostly red in three paratypes (Figure 19a)

Variation in female paratypes
Head. As for holotype but medial black stripe of labial palp as wide as 0.33-0.40 of palp width (Figure 1a-b); labrum reddish brown with black along distal margin and pair of yellow laterobasal spots; ventral half of antefrons pale brown medially and pale yellow laterally, dorsal portion of antefrons, postfrons, and vertex pale to dark reddish brown, occipital triangle pale to dark reddish brown, rear of head reddish brown with two yellow spots behind eyes.
Thorax. As for holotype but prothorax reddish brown except anterior lobe, mediodorsal area of middle lobe, and posterior margin of posterior lobe yellow. Pterothorax (Figure 2b-c) as for holotype but sometimes a narrow stripe along medial third of mesepisternal-mesepimeral carina; mesepimeron with triangular spot on anterodorsal corner; metafemur armed with 12-19 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur; arculus between Anx 2 and Anx3, usually closer to Anx 2, sometimes midway; Fw discoidal field with 3 rows of cells at base to 4-6 rows at hind margin, Hw with 2-3 at base, then 2, then increasing to 12-17 at hind margin; anal loop enclosing 22-26 cells. Pt orange to reddish brown, overlying 4-6 cells. Anx: 14-17 in Fw, 12-15 in Hw; Pnx: 14-18 in Fw, 15-18 in Hw.
Abdomen. As in holotype but S4 as long as ca. twice to almost three times its apical width (ratio width/length = 0.38-0.53). Dorsal terga (Figure 7a) with mid-dorsal carina bordered by yellow in S1-4, and by pale orange in S5-7; S1-3 pale reddish brown with ventral third yellow; S4-7 pale orange red with medial third diffusely brown to dark reddish brown or black; S8 dark reddish brown to black with ventrolateral flap pale yellow; S9-10 and caudal appendages dark reddish brown to black. Ventrolateral carina on S2 vestigial, on S3-7 well developed and black. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 8a) reddish brown with yellow stripe along ventrolateral carinae on S1-6, orange in S7; S8 pale yellow; S9-10 The levis group of Orthemis revisited 141   (Figure 16a, d, f, g-h, j, l, o). It differs from all of them by the splinter-shaped lateral projection on medioectal lobe of vesica spermalis distal segment (Figure 13a; versus no sclerotized projections or of different shape, Figure 13d, d, g-h, m), and by hamule shape, with inner and outer branches ca. equally long (Figure 11a; versus inner branch longer than outer branch in O. harpago, Figure 11m),  Figure 18k).

Remarks
Several specimens of this species from Surinam were misidentified as O. flavopicta [RMNH, Belle, 2002], one male from Panama as O. anthracina [RWG, von Ellenrieder, 2009], and one female from Trinidad as O. cultriformis [RWG, von Ellenrieder, 2009]. Correlation of shape of vesica spermalis and hamule in a relatively large series of specimens, and association of males and females based on distribution, allows me to conclude that this is a separate undescribed species.
The levis group of Orthemis revisited 143

Diagnosis
Male of O. ambinigra has metallic blue to purple frons and vertex, labial palp with medial black stripe as wide as 0.66 of each palp or more, pterothorax brown with yellow stripes, and abdomen red except for carinae narrowly black and a black mediodorsal longitudinal spot on S8-9 ( Figure 5a). Abdomen narrows gradually from S2 to S4 (Figure 6b), and S4 is about as wide at level of apical carina as 0.35-0.50 of its length. Hamule inner and outer branches are located in the same plane (Figure 10b), with both ending at about the same level; inner corner of outer branch is angled, and outer corner is smoothly rounded (Figure 11b). Distal segment of vesica spermalis is trapezoidal, with sides slightly convex and maximum width near mid-length of basal sclerotized portion in ectal view (Figure 13b), with paired distal medial longitudinal membranous lobes beset with denticles along distal margin on ectal side, paired outer lateral lobes shorter than basal sclerotized portion and folded medially to the sides, and paired flagella (Figures 12b, 13b). Cercus curves ventrally gradually ( Figure 15c) and is widest at base in dorsal view ( Figure 14b). Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16b). It differs from all of them by the wider extension of black on premental palps (0.60 or more versus 0.40 or less, with the possible exception of O. teres known only from the holotype missing labium), S8-9 with black mediodorsal stripe (versus S8-9 or at least S8 entirely red dorsally), and shape of outer branch of hamule (Figure 11b), with inner corner angled (inner corner rounded in O. ambirufa, Figure 11c) and outer surface not grooved (grooved in O. tambopatae and O. teres, Figure 11t-u).
Female shares with male: labial palp with wide medial black stripe (0.50 or more of each palp), pterothorax brown with yellow stripes (Figure 2d), abdomen narrowing gradually from S2 to S4 (Figures 7b, 8b), and S4 about as wide at level of apical carina as 0.35-0.50 of its length. Color of abdomen is pale orange to reddish brown, with yellow stripes and brown diffuse spots (Figures 7b, 8b), and black mid-dorsal stripes on S7-9 to S8-9. Ventrolateral flap on S8 is pale yellow and relatively wide, ca. 0.28-034 as wide as its length, with a smoothly curved contour ( Figure 17b Buchholz's (1950) description of O. plaumanni was based on three males from Nova Teutonia, Santa Catarina, Brazil. He did not include illustrations except for a lateral view of the genital fossa, his diagnosis from other species in the levis group of Orthemis being based solely on the presence of two bridge crossveins. Even though some specimens from Peru I examined earlier (identified as O. plaumanni in von Ellenrieder, 2009) seemed to fit well the description of O. plaumanni in terms of venation and color, the shape of male hamule differed from that illustrated by Buchholz (1950) Buchholz (1950). alt: antero-lateral triangular projection; de: paired depression; me: medial excision; mlr: medio-longitudinal ridge; lm: lateral margin; sr: semi-circular rib; tr: transverse ridge; trp: posterior projection of lateral end of transverse ridge; ur: inverted U-shaped ridge. and ventral terga, as well as color pattern of the holotype of O. plaumanni Buchholz fully agree with those of the holotype of O. ambinigra Calvert. The holotype of O. plaumanni has two bridge crossveins in both Fw and only one in both Hw. Supernumerary bridge crossveins are common in O. ambinigra, with ca. 55% of the examined specimens having them in at least one wing, although their presence and number are variable and cannot be used as a diagnostic character: ca. 5% of examined specimens have two to three bridge crossveins in three wings, ca. 25% two to three bridge crossveins in two wings, and ca. 25% two bridge crossveins in one wing. Supernumerary bridge crossveins are also found on occasion in other species of Orthemis, i.e. I observed them in O. paulsoni, O. philipi, O. tambopatae, and O. teres.

Specimens examined
Total 10

Diagnosis
Male of O. ambirufa has reddish to orange-brown frons and vertex lacking metallic reflections, labial palp with medial black stripe as wide as 0.33 or less of palp width, pterothorax reddish brown with yellow stripes (Figure 2e), and dorsal terga of S2-10 red except for ventrolateral and The levis group of Orthemis revisited 155 posterior carinae narrowly black. Abdomen narrows gradually from S2 to S4, and S4 is about as wide at level of apical carina as 0.33-0.55 of its length. Hamule inner and outer branches are located in the same plane ( Figure 10c), with both ending at about the same level; inner and outer corners of outer branch are smoothly rounded (Figure 11c). Distal segment of vesica spermalis is ca. rectangular, with sides approximately parallel in ectal view (Figure 13c), with paired distal medial longitudinal membranous lobes beset with denticles along distal margin on ectal side, paired outer lateral lobes about as long as basal sclerotized portion and folded medially to the sides (Figure 12c), and paired flagella. Cercus curves ventrally only slightly and gradually ( Figure 15d) and is almost straight and cylindrical in dorsal view ( Figure 14c). Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16c Female shares with male: labial palp with narrow medial black stripe (0.33 or less of palp width), pterothorax reddish brown with yellow stripes, abdomen narrowing gradually from S2 to S4 (Figures 7c,8c), and S4 about as wide at level of apical carina as 0.34-0.44 of its length. Color of dorsal terga S2-10 is pale orange to reddish brown, with longitudinal yellow stripes along mediodorsal and ventrolateral carinae (Figures 7c, 8c). Ventrolateral flap on S8 is yellow and relatively narrow, ca. 0.21-0.26 as wide as its length, with a smoothly curved contour ( Figure 17c). Sternum S8 is smooth, with vulvar lamina consisting of posterior margin of S8 forming a shallow inverted U-shaped rim, with the medial excision wider than its lateral margins ( Figure 18d). Orthemis attenuata, O. concolor, and O. levis also have a smooth sternum S8 with posterior margin of sternum S8 forming an inverted U-shaped rim, but the medial excision is narrower than its sides (Figure 18e, i, p), and their abdomen is much more slender, abruptly narrowed at base (Figures 7e, g, m, 8e, g), while the abdomen is wider and only slightly and gradually narrowed at base in O. ambirufa (Figures 7c, 8c).

Description of female
Head. Labium black, with palps pale yellow with medial black stripe as wide as 0.33-0.40 of palp width and anterior margin narrowly black ( Figure 1d); labrum black with pair of yellow laterobasal spots; base of mandibles yellow; lateral portion of clypeus along eyes pale yellow, remainder of clypeus pale brown; ventral half of antefrons pale brown medially and pale yellow laterally, dorsal portion of antefrons, postfrons, and vertex pale to dark reddish brown, with medial furrow of postfrons and vertex around ocelli with metallic reflections in some females; occipital triangle pale to dark reddish brown with yellow posterior surface, rear of head reddish brown with two yellow spots behind eyes. Postfrons with wide shallow medial furrow; vertex with a pair of low tubercles; posterior margin of occipital triangle slightly bilobate.
Thorax. Prothorax reddish brown except anterior lobe, mediodorsal area of middle lobe, and posterior margin of posterior lobe yellow. Pterothorax ( Figure 19d) reddish brown with pale yellow stripes as follows: mesepisternum with medial longitudinal stripe adjacent to mediodorsal carina, longitudinal stripe parallel to mesepisternal-mesepimeral carina, and narrow stripe along medial third of mesepisternal-mesepimeral carina; mesepimeron with wide stripe along posterior half narrowing dorsally; metepisternum with narrow stripe along ventral margin and another sinuous one ventral to metastigma; metepimeron with a stripe along posterior half, narrowing dorsally, and subtriangular spot on anterodorsal corner; venter of pterothorax pale yellow with lateral margins from base of leg to posterior margin reddish brown. -Posterior surface of coxa, trochanter, and femur of front leg and posterior surface of coxa of middle and hind leg pale yellow, remainder dark reddish brown; tibia, tarsus, pretarsus, and spination black; metafemur armed with 14-22 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings entirely infumated in 45% of the females, to hyaline with restricted amber areas, from base to level of first row of anal cells in FW and to third row in HW, and apex across distal three rows to distal end of pt. One cubito-anal crossvein and arculus between to Anx 2 and The levis group of Orthemis revisited 157 Anx 3, usually closer to Anx 2, sometimes midway, rarely opposite to Anx 2; sectors of arculus stalked; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4 cells, rarely with 5 cells; one bridge crossvein; Fw discoidal field with 3 rows of cells at base to 4-6 rows at hind margin, Hw with 2-3 at base, then 2, then increasing to 12-16 at hind margin; 3 rows of cells between wing margin and anal loop at level of anal angle of triangle; anal loop enclosing 19-25 cells. Pt orange to reddish brown, overlying 4-6 cells. Anx: 16-20 in Fw, 13-15 in Hw; Pnx: 13-17 in Fw, 13-17 in Hw.

Variation in males
Head. As for female but labial palp with medial black stripe as wide as about 0.4-0.5 of palp width ( Figure 1c); labrum as in females in teneral specimens black; clypeus and basal portion of antefrons pale to dark reddish brown to black medially; dorsal portion of antefrons, postfrons, and vertex metallic purple with blue; occipital triangle pale reddish brown to dark reddish brown to black.
Thorax. Prothorax as for female to dark reddish brown to black. Pterothorax as in female in teneral males, dark reddish brown with yellow diffuse stripes, to black on mesepisternum with very faint yellowish stripes on metepimeron and metepisternum ( Figure 19c) to almost entirely black with metallic blue reflections and brown dashes along mid-dorsal carina and proximal third of mesepisternum (De Marmels, 1989). Venter of pterothorax as in female or black with central area brown. -Legs as in female or black with lateral and extensor surfaces of procoxa and protrochanter pale brown; metafemur armed with 18-35 short spurs, usually followed by 1 longer spur, rarely by two. -Anal loop enclosing 18-23 cells. Pt pale to dark reddish brown. Fw Anx: 16-19; Hw Anx: 12-16; Fw Pnx: 14-17; Hw Pnx: 13-19.
Abdomen. As for female but S4 longer than three times its apical width (ratio apical width/length = 0.21-0.31). Dorsal terga S1-9 mostly red with posterodorsal faint black spot and S10 black with red mid-dorsal spot, to red with black posterodorsal spot extending to posterior margin on S3-7 or along most of S5-7 and S8-9 black with ventrolateral red stripe, to abdomen almost entirely black; caudal appendages black. Ventrolateral carina on S2 vestigial, on S3-8 well developed and black, on S9 well developed along basal two thirds, absent at distal third. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga ( black to dark reddish brown or black with yellow stripe along ventrolateral carinae on S1-7; S8-10 dark reddish brown to black. Anterior lamina in lateral view shorter than hamule and as high as genital lobe ( Figure 10d); hamule bifid with small inner branch forming short pointed spine, and larger outer branch bent ventrally over inner branch with blunt square tip; outer corner of outer branch rounded (Figure 11d). Distal segment of vesica spermalis with basal portion trapezoidal in ectal view (Figure 13d), and with long flagella on ental surface; distal lobes represented on each side by medial longitudinal semicircular membranous lobe with sclerotized lateral projection margined with denticles, and elongate rhomboidal outer lobe longer than basal portion of distal segment in lateral view (Figure 12d). Cercus markedly curved ventrally in lateral view, with tip upturned, with a row of 4-6 tubercles along distal dilated portion ( Figure 15e); in dorsal view converging along basal 0.70 with opposite cercus, then approximately parallel to it along distal 0.30 (Figure 14d). Epiproct extending to 0.65-0.74 of cerci length, as long as 0.77-0.84 of its maximum width, with apex bifid and as wide as 0.72-0.80 of its maximum width (Figure 16d).  The levis group of Orthemis revisited 161 recent examination of a relatively large series of specimens from the lowland Amazon forest from Peru, Surinam, and Brazil fitting closely the description of O. anthracina and differing from the male from Panama in morphology of vesica spermalis and hamule allowed me to conclude that the male studied previously was misidentified, representing the species I describe here as O. aciculata. Several of the specimens of O. anthracina found in collections were misidentified as O. cultriformis [UMMZ] and O. flavopicta [RMNH].

Distribution and biology
Orthemis anthracina is known from Venezuela and Trinidad south through Surinam and N Peru ( Figure 25). Habitats include forested creeks, streams, rivers, and lakes.

Diagnosis
Male of O. attenuata has metallic purple frons and vertex, labial palp with medial black stripe as wide as 0.33-0.40 of palp width, pterothorax reddish brown with yellow stripes (Figure 19f), which can be obscured in mature males (Figure 19e), dorsal terga of S1-3 mostly reddish brown to orange, and S4-10 mostly black with narrow yellow to orange stripes along S4-7 medial longitudinal carinae and along S4-5 to S4-8 ventrolateral carinae. Abdomen narrows abruptly from S2 to S4 (Figures 19e-f), and S4 is about as wide at level of apical carina as 0.22-0.33 of its length. Hamule inner and outer branches are located in the same plane, with both ending at about the same level (Figures 9b, 10e); inner corner of outer branch is smoothly rounded (Figure 11e). Distal segment of vesica spermalis is ca. oval, with sides gradually converging to distal end in ectal view (Figure 13e), paired distal short medial longitudinal membranous lobes beset with denticles along distal margin on ectal side, paired outer lateral lobes shorter than basal sclerotized portion of distal segment, folded medially to the sides (Figure 12e), and paired flagella. Cercus gradually curves ventrally ( Figure 15f) and its tip is slightly upturned. Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16e).
Male of O. attenuata shares the combination of an abruptly narrowed abdomen at base and epiproct tip narrow and entire only with O. concolor and O. levis. These three species are morphologically very similar, differing almost solely by color pattern. Orthemis attenuata differs from O. concolor by its metallic purple frons and vertex, in teneral males reddish brown with some metallic purple reflections on medial furrow of frons and around ocelli, and by the pterothorax with yellow stripes, which can be obscured in mature males; versus frons and vertex reddish brown and pterothorax concolorous reddish brown (Figure 19g-h), sometimes with ill-defined diffuse yellow areas on mesepisternum, more extensive in teneral males. Orthemis attenuata can be distinguished from O. levis by its abdominal color pattern, with S1-3, or S1-4 in teneral males, mostly reddish brown to orange, and S4-10 mostly black with only narrow yellow to orange medial longitudinal stripes on S4-7 and ventrolateral stripes on S4-5 to S4-8 (versus abdomen mostly red, with only S8-10 dorsally black in O. levis), and by shape of abdomen base between apex of S2 and base of S3, which is more markedly globose in O. attenuata (Figures 19e-f) than it is in O. levis (as in Figure 7m).
Female shares with male: labial palp with narrow medial black stripe (0.33-0.40 of palp width), pterothorax reddish brown with yellow stripes, and slender abdomen narrowing abruptly from S2 to S4 (Figures 7e, 8e). In some females there are metallic reflections on medial furrow of frons and on vertex surrounding ocelli. Color of dorsal abdominal terga is similar to that of male. Ventrolateral flap on S8 is yellow and relatively narrow, ca. 0.23-0.26 as wide as its length, with a smoothly curved contour (Figure 17e). Sternum S8 is smooth, with vulvar lamina consisting of The levis group of Orthemis revisited 163 posterior margin of S8 forming a shallow inverted U-shaped rim, with medial excision narrower than its lateral margins (Figure 18e).
Combination of relatively slender abdomen narrowing abruptly at base, with base of S4 less than half as high as base of S3 in lateral view (Figure 7e), and vulvar lamina consisting of shallow inverted U-shaped rim of S8 posterior margin is shared only with O. concolor and O. levis. Orthemis attenuata differs from O. concolor by its pterothorax with yellow stripes (as Figure 19f), versus pterothorax concolorous reddish brown (Figure 19h) with variable extent of ill-defined diffuse yellow areas on mesepisternum in teneral females, and by the position of the paired depressions anterior to vulvar lamina rim, which in O. attenuata are located directly anterior to the lateral margins of the medial excision of the rim (Figure 18e), whereas in O. concolor they are located laterally to the medial excision ( Figure 18i). Differentiation of the female from that of O. levis is based on shape of abdomen base between apex of S2 and base of S3, which is more markedly globose in O. attenuata (Figure 7e) than in O. levis ( Figure 7m). As in the male, female of O. attenuata also differs by dorsal terga S4-7 mostly dark reddish brown to black with narrow mediodorsal and ventrolateral pale stripes versus mostly red in O. levis. Red color on abdominal dorsal terga S4-7 in females of O. levis is not as pronounced as it is in males, being sometimes replaced by reddish brown, and unlike in males, in females of O. levis there are yellow to orange pale stripes along mediodorsal and ventrolateral carinae of S1-7 to S1-8, so that abdominal color of poorly preserved females can look very similar to that of females of O. attenuata.
Remarks Ris (1919Ris ( , p. 1105) indicated that Dr. Grünberg was unable to find the type of this species in the collection of the Berlin Museum [ZMHB]. However, Michael Ohl, current curator at that institution, confirmed that the holotype is in fact hosted there, and Mélanie Turriault kindly photographed it (Figure 19f), allowing for confirmation of the correct association of this name by Ris and further workers.

Distribution and biology
Extends from Colombia, Venezuela, and Guyana south to Peru,Bolivia,and Brazil (Figure 24). Habitat includes rivers, streams, and ponds in forested areas. sp. nov. Figures 1e,2f,4b,5b,6d,9c,10f,11f,12f,13f,14e,15f,16f,23 Etymology From Latin celata (concealed), the female gender of the adjective celatus, referring to the position of hamule inner branch, concealed from sight in lateral view. Head. Labium black, with palps pale yellow with medial black stripe as wide as 0.40 of palp width and anterior margin narrowly black ( Figure 1e); labrum black with pair of rounded laterobasal pale yellow spots; base of mandibles yellow; lateral portion of clypeus along eyes pale yellow, remainder of anteclypeus pale brown, postclypeus pale brown with medial dark brown spot and postclypeal lobes narrowly fringed with dark brown line along distal margin; ventral half of antefrons pale brown medially and pale yellow laterally, dorsal portion of antefrons, postfrons, and vertex metallic purple with blue reflections, occipital triangle dark reddish brown, rear of head reddish brown. Postfrons with wide shallow medial furrow; vertex with pair of low tubercles; posterior margin of occipital triangle slightly bilobate.

Type specimens examined
Thorax. Prothorax reddish brown. Pterothorax (Figure 2f) reddish brown with pale yellow stripes as follows: wide stripe along posterior half of mesepimeron narrowing dorsally; metepisternum with narrow stripe along ventral margin and another sinuous one ventral to metastigma, and oval spot on posterodorsal corner; metepimeron with stripe along posterior half, narrowing dorsally, and subtriangular spot on anterodorsal corner; venter of pterothorax pale yellow with triangular reddish brown spot on each side extending from base of leg and narrowing to posterior margin. -Legs with coxa, trochanter, and base of femur reddish brown; femur and tibia dark reddish brown; tarsus, pretarsus, and spination black; metafemur armed with 25 (right) to 27 (left) short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur.
-Wings infumated with small amber spots at base extending to level of first row of anal cells in FW and HW, very narrow reddish tinge around FW nodus, and distal amber spot extending from level of distal end of pterostigma to apex. One cubito-anal crossvein and arculus distal to Anx 2 in Fw and HW; sectors of arculus stalked; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4 (right) to 6 (left) cells; one bridge crossvein in Fw and Hw; Fw discoidal field with 3 rows of cells at base to 4 (right) or 5 (left) rows at hind margin, Hw with 3 at base, then 2, then increasing to 16 (right) or 13 (left) at hind margin; 3 rows of cells between wing margin and anal loop at level of anal angle of triangle; anal loop enclosing 22 cells. Pt reddish brown, 4.6 (right) and 4.65 (left) long in Fw and 4.5 (right) and 4.35 (left) in Hw, overlying 6 cells in Fw and 5-6 cells in Hw. Anx: 16 in Fw; 13 in Hw; Pnx: 13 (right) and 14 (left) in Fw; 14 in Hw.
Abdomen. Sides linear, gradually narrowing from S3 to S4 in ventral view (Figure 6d), S4 less than three times as long as wide (ratio width/length = 0.38). Dorsal terga (Figure 5b) with middorsal carina narrowly pale yellow in all segments; S1 dark reddish brown; S2-3 orange red; S4-7 mostly orange red, except for small oval posterodorsal faint black spot on posterior 0.15 of segment; S8 orange red with pair of small rounded black spots at mid-length, and dark diffuse mid-dorsal stripe; S9 dark reddish brown; S10 black; caudal appendages dark reddish brown. Ventrolateral carina on S2 vestigial, on S3-8 well developed and black, on S9 well developed along basal two thirds, absent at distal third. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 6d): S1 and S2 dark reddish brown with yellow along ventrolateral carinae; S3-7 dark reddish brown with reddish stripe along lateral margin on basal two thirds of segment; S8 dark reddish brown with reddish stripe along lateral margin on basal half of segment; S9-10 dark reddish brown. Anterior lamina in lateral view shorter than hamule and as high as genital lobe (Figure 10f); hamule bifid with small inner branch forming short pointed spine, parallel to larger outer branch, directed anteriorly (Figure 9c), and not visible in lateral or frontal views of hamule (Figures 10f, 11f); outer branch tip bent ventrally at right angle in lateral view, separated from inner branch by a distance shorter than inner branch length ( Figure 9c); outer corner of hamule smoothly curved (Figure 10f). Distal segment of vesica spermalis with basal portion trapezoidal in ventral view (Figure 13f), and with a pair of long flagella on ental surface; distal lobes represented on each side by an elongate outer oval-shaped lobe ca. as long as basal portion of distal segment in lateral view, and medioectal sclerotized lobe margined with denticles distally and lacking any lateral sclerotized projections; medioectal lobe distinctly shorter than outer lobe (Figure 12f). Cercus markedly curved ventrally in lateral view, with row of 6 ventral tubercles along distal half and upturned tip ( Figure 15h); in dorsal view converging along basal 0.66 with opposite cercus, then approximately parallel to it along distal 0.33 (Figure 14e). Epiproct extending to 0.75 of cerci length, with apex bifid and about as wide as 0.73 of its basal width (Figure 16f).  Figure 11a, d, h, k, m) and by paired medioectal lobes of vesica spermalis distal segment almost entirely membranous, with only some marginal denticles along distal margin (Figures 12f, 13f; versus with some kind of lateral sclerotized projection, Figures 12a, d, j, m, 13a, d, g-h, m). It further differs from O. anthracina by its narrower epiproct apex, 0.70 mm measured subapically (Figure 16f; versus 0.80-0.85 mm measured subapically, Figure 16d), from O. cinnamomea by frons and vertex metallic purple with blue reflections (versus reddish brown), and from O. harpago by abdomen gradually narrowed at base in lateral view and dorsal terga of S1-7 mostly red (Figures 4b, 5b; versus constricted abruptly and dorsal terga S1-7 mostly black in mature males, with mediodorsal yellow stripe, Figure 4i).

Dimensions
Female unknown.

Remarks
The holotype male of O. celata in the collection of Naturalis Leiden [RMNH] was identified as Orthemis flavopicta by Geijskes' in 1970.

Diagnosis
Male of O. cinnamomea has pale to dark reddish brown frons and vertex, labial palp with medial black stripe as wide as ca. 0.33 of palp width, pterothorax reddish brown with pale yellow stripes, dorsal terga mostly reddish brown, with ventrolateral margins of S1-3 pale greenish-yellow, and narrow longitudinal orange stripes mediodorsally on S1-7 and both dorsal and ventral to ventrolateral carinae on S3-7 (Figure 4c). Abdomen narrows abruptly from S2 to S4 (Figure 4c), and S4 is about as wide at level of apical carina as 0.28-0.37 of its length. Hamule inner branch is small and ends in a short point, outer branch is larger, with a blunt tip bent ventrally over inner branch; outer corner of outer branch is smoothly rounded (Figures 10g, 11h). Basal portion of distal segment of vesica spermalis is ca. trapezoidal in ectal view (Figure 13g), with distal lobes represented on each side by a medioventral semicircular membranous lobe with a sclerotized lateral projection margined with denticles, and an elongate outer lobe approximately rectangular in lateral view, about as long as basal portion (Figure 12g), with paired flagella on ental surface. Cercus (Figure 14f) is markedly curved ventrally in lateral view, with tip upturned (Figure 15i). Epiproct apex is about as wide as 0.50-0.55 of its maximum width, and is slightly bifid (Figure 16g Figures 4a, b, e, 19c). It can be further diagnosed from O. harpago by shape of hamule, with inner branch shorter than outer branch and in the same plane (Figures 10g, 11h; versus inner branch longer then outer branch and parallel to it, Figures 9f, 10n, 11h).
Female shares with male: labial palp with narrow medial black stripe, ca. 0.33 of palp width, pterothorax reddish brown with yellow stripes, and relatively slender abdomen narrowing abruptly from S2 to S4 (Figure 7f), with S4 as wide at apex as 0.30-0.39 of its length. Color of dorsal abdominal terga is similar to that of male. Ventrolateral flap on S8 (Figures 17f, 19b) is white, as wide as 0.31-0.34 of its length. Vulvar lamina of S8 consists of a medial longitudinal ridge anterior to posterior margin of sternum S8, enclosed anterolaterally by semicircular ribs, as wide as 0.30 of sternum width, with anterior corners projected anterolaterally and convex (Figure 18h Figure 18l, o). It further differs from all of them except for O. harpago by abdomen narrowing abruptly at base, with base of S4 as high as less than half of base of S3 in lateral view (Figure 7f; versus narrowing gradually, with base of S4 as high as more than half of base of S3 in lateral view, Figure 7a, d, h, i), and from O. harpago by S8 lateral flap relatively wider, as wide as 0.31-0.34 of its length, and triangular in shape (Figure 17f, versus lower, as high as 0.29 of its length, and ca. smoothly convex, Figure 17m).

Remarks
Additional specimens examined since the original description of this species agree well with the type series.

Diagnosis
Male of O. concolor has pale to dark reddish brown frons and vertex, labial palp with medial black stripe as wide as 0.33-0.40 of palp width, pterothorax concolorous reddish brown (Figure 19g) with ill-defined diffuse yellow areas on sides of pterothorax in young males, dorsal terga S1-3 mostly reddish brown to orange, and S4-10 mostly black with narrow yellow to orange stripes along S4-7 medial longitudinal carinae and along S4-5 to S4-8 ventrolateral carinae (Figure 19g). Abdomen narrows abruptly from S2 to S4 (Figure 4d), and S4 is about as wide at level of apical carina as 0.25-0.30 of its length. Hamule inner and outer branches are located in the same plane, with both ending at about the same level ( Figure 10h); inner corner of outer branch is smoothly rounded (Figure 11i). Distal segment of vesica spermalis is ca. oval, with sides gradually converging to distal end in ectal view (Figure 13i), paired distal short medial longitudinal membranous lobes beset with denticles along distal margin on ectal side, paired outer lateral lobes shorter than basal sclerotized portion of distal segment folded medially to the sides (Figure 12h), and with paired flagella. Cercus (Figure 14g) curves ventrally gradually (Figure 15j) and its tip is slightly upturned. Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16i). Male of O. concolor shares combination of abruptly narrowed abdomen at base and epiproct tip narrow and entire only with O. attenuata and O. levis. These three species are very similar morphologically, differing almost solely by color pattern. Orthemis concolor differs from O. attenuata and O. levis by its frons and vertex reddish brown and pterothorax concolorous reddish brown (Figure 19g), sometimes with ill-defined diffuse yellow areas on mesepisternum, versus metallic purple frons and vertex, in teneral males reddish brown with some metallic purple reflections on frons furrow and around ocelli, and pterothorax with yellow stripes (Figure 19f), which can however be obscured in mature males (Figure 19e), and it further differs from O. levis by its abdominal color pattern, with S1-3 mostly reddish brown to orange and S4-10 mostly black with only narrow yellow to orange medial longitudinal stripes on S4-7 and ventrolateral stripes on S4-5 to S4-8 (Figures 4d, 19g), versus S1-7 mostly red, with only S8-10 dorsally black in O. levis, and by shape of abdomen base between apex of S2 and base of S3, which is more markedly globose (Figure 4d; versus less globose, as in Figure 7m).
Female shares with male: labial palp with narrow medial black stripe (0.33-0.40 of palp width), pterothorax almost uniformly reddish brown with ill-defined diffuse yellow areas on sides in young specimens, and slender abdomen narrowing abruptly from S2 to S4 (Figure 7g). Color of dorsal abdominal terga is similar to that of male. Ventrolateral flap on S8 is yellow and relatively narrow, ca. 0.27 as wide as its length, with a smoothly curved contour (Figure 17g). Sternum S8 is smooth, with vulvar lamina consisting of posterior margin of S8 forming a shallow inverted U-shaped rim, with medial excision narrower than its lateral margins (Figure 18i). Combination of relatively slender abdomen narrowing abruptly at base, with base of S4 as high as less than half of base of S3 in lateral view, and vulvar lamina consisting of shallow inverted U-shaped rim of S8 posterior margin is shared only with O. attenuata and O. levis. Orthemis concolor differs from both by its pterothorax concolorous reddish brown (Figure 19h) with ill-defined yellow areas in teneral specimens, versus pterothorax with yellow stripes, and by the position of the paired depressions anterior to vulvar lamina rim, which are located laterally to the lateral margins of the medial excision of rim (Figure 18i), versus located directly anterior to the lateral margins of the rim (Figure 18e). It further differs from O. levis by dorsal terga S4-7 mostly dark reddish brown to black with narrow mediodorsal and ventrolateral pale stripes versus mostly red, and by shape of abdomen base between apex of S2 and base of S3, which is more markedly globose (Figure 7g; versus less globose, Figure 7m). However, red color on abdominal dorsal terga S4-7 in females of O. levis is not as pronounced as it is in males, being sometimes replaced by orange to reddish brown, which especially in poorly preserved specimens looks then very similar to the pattern of O. concolor.

Distribution and biology
Orthemis concolor is found from Venezuela and Trinidad through the Guyanas to N Brazil (Figure 24), associated with forested streams and rivers.

Description of female
Head. Prementum black; labial palps pale yellow with medial black stripe as wide as about 0.40 of palp width and anterior margin narrowly black (as in Figure 1f); labrum black with a mediobasal reddish spot or with pair of rounded laterobasal pale yellow spots; base of mandibles yellow; lateral portion of clypeus along eyes pale yellow, remainder of anteclypeus pale reddish brown, postclypeus pale reddish brown with medial dark brown spot and postclypeal lobes narrowly fringed with dark brown line along distal margin; ventral half of antefrons reddish brown medially and pale yellow laterally, dorsal portion of antefrons, postfrons, and vertex reddish brown, occipital triangle red to reddish brown posteriorly, rear of head orange to reddish brown around occipital triangle and yellow along eyes. Postfrons with wide shallow medial furrow; vertex with pair of low tubercles; posterior margin of occipital triangle slightly bilobate.
Thorax. Prothorax reddish brown except pale yellow anterior and posterior lobes. Pterothorax ( Figure 2j) reddish brown with pale yellow stripes as follows: narrow diffuse stripe lateral to mediodorsal carina, wider diffuse stripe at mid width, and small spot at mesepisternalmesepimeral suture ventral kink; mesepimeron with wide stripe along posterior half narrowing dorsally; metepisternum with narrow stripe along ventral margin and another sinuous one ventral to metastigma, and oval spot on posterodorsal corner; metepimeron with stripe along posterior half, narrowing dorsally, and a triangular spot on anterodorsal corner; venter of pterothorax pale yellow with triangular reddish brown spot on each side extending from base of leg and narrowing to posterior margin. -Legs with coxa, trochanter, and basal 0.50 of inner surface of pro-and mesofemur pale brown to pale yellow; femur reddish brown; tibia dark reddish brown, tarsus, pretarsus, and spines black; metafemur with 11 to 20 short spurs which slightly and gradually increase in size towards apex, followed distally by one or two longer spurs. -Wings hyaline with small amber spots at base extending to level of first row of anal cells in FW, and to level of second row and distal end of membranula in HW, very narrow reddish tinge around nodus, and distal amber spot extending from level of distal end of pterostigma to apex. One cubito-anal crossvein and arculus distal to Anx 2 in Fw and HW; sectors of arculus stalked; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4 cells; one bridge crossvein in Fw and Hw; Fw discoidal field with 3 rows of cells at base to 5-7 rows at hind margin, Hw with 3 at base, then 2, then increasing to 14-15 at hind margin; 3 rows of cells between wing margin and anal loop at level Abdomen. Sides linear, gradually narrowing from S3 to S4 in ventral view (Figure 8h), S4 about three times as long as wide (ratio width/length = 0.30-0.33). Dorsal terga (Figure 7h) S1-7 orange red, except for mid-dorsal and ventrolateral pale yellow stripes, and pair of small faint dark oval spots on posterolateral corner of S3-6 and posterior 0.2 black on S7 in two females, in one female S4-7 orange red lacking yellow ventrolateral stripes; S8-10 black with mid-dorsal carina narrowly pale yellow. Ventrolateral carina on S2 vestigial, on S3-8 well developed and black, on S9 absent. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 8h): S1 and S2 dark reddish brown with yellow along ventrolateral carinae; S3-7 dark reddish brown with yellow stripe along lateral margin on basal two thirds of segment; S8 pale brown; S9 dark reddish brown; S10 black. Ventrolateral flap on S8 pale yellow (Figure 17h). Vulvar lamina on sternum of S8 extending along posterior 0.25 of S8 length, with free transverse anterior ridge projected into two triangular, concave points directed anteriorly, and medial longitudinal ridge transversally ridged extending to S8 posterior margin (Figure 18j). Appendages dark reddish brown.

Variation in males
Head. As for female but labial palp medial black stripe as wide as about 0.50-0.70 of palp width ( Figure 1f); labrum black; clypeus and basal portion of antefrons pale to dark reddish brown to black medially; dorsal portion of antefrons and postfrons in two teneral males reddish brown with purple reflections and vertex metallic blue; in four males antefrons, postfrons, and vertex metallic purple with blue reflections; in one male antefrons and postfrons metallic purple with blue reflections and vertex metallic blue; in three males antefrons, postfrons, and vertex bright metallic blue with purple reflections, and in two males bright metallic blue with turquoise reflections; occipital triangle pale reddish brown to dark reddish brown to black.
Thorax. Prothorax as for female to dark reddish brown to black. Pterothorax as in female in two teneral males, dark reddish brown with yellow diffuse stripes (as in Figure 2j), to black on mesepisternum and very faint yellowish stripes on metepimeron and metepisternum in five males (Figure 19i), to almost entirely black with greenish-blue metallic reflections with some restricted brown areas on ventral portion of sclerites in five males (Figures 2i, 19j). Venter of pterothorax as in female to black with central area brown. -Legs as in female to black with lateral and extensor surfaces of procoxa and protrochanter pale brown; metafemur armed with 23-33 short spurs, usually followed by 1 longer spur, rarely by two. -Anal loop enclosing 20-25 cells. Pt pale to dark reddish brown. Fw Anx: 16-20; Hw Anx: 12-15; Fw Pnx: 12-15; Hw Pnx: 13-17.
Abdomen. As for female but S4 as long as ca. twice to three times its apical width (ratio apical width/length = 0.33-0.48). Dorsal terga S1-7 mostly red with posterodorsal black spot in one teneral male, black spot extending to posterior margin on S3-7 in two males (Figure 19i), along most of S5-7 length in two males, and S3-7 almost entirely black (Figure 19j) in three males; S8 black with ventrolateral red stripe to all black; S9-10 and caudal appendages black. Ventrolateral carina on S2 vestigial, on S3-8 well developed and black, on S9 well developed along basal two thirds, absent at distal third. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 6e): mostly black to dark reddish brown or black with yellow stripe along ventrolateral carinae on S1-7; S8-10 dark reddish brown to black. Anterior lamina in lateral view shorter than hamule and as high as genital lobe (Figure 10i); hamule bifid with small inner branch forming a short pointed spine, and larger outer branch bent ventrally over inner branch with blunt square tip; outer corner of outer branch angled (Figure 11k). Distal segment of vesica spermalis with basal portion trapezoidal in ectal view (Figure 13h), and with long flagella on ental surface; distal lobes represented on each side by medioventral semicircular membranous lobe with sclerotized lateral projection margined with denticles, and elongate outer lobe approximately rectangular and ca. as long as basal portion in lateral view (Figure 12i). Cercus markedly curved ventrally in lateral view, with tip slightly upturned, with row of 5-7 tubercles along distal dilated portion ( Figure 15l (Figure 17h; versus relatively wider, with ratio width/length of 0.33-0.37, Figure 17i).

Remarks
Male color pattern was found to be variable, probably related to age, from very similar to that of females in teneral males -as indicated by their shiny wing membranes and abdomen crushed due likely to softness of integument at time of fixing -with yellow stripes on thorax and abdomen mostly red, except for metallic reflections of frons and vertex metallic and wider extension of black on labial palp, to very dark, probably corresponding to older mature males, in which the black on labial palp is the widest, frons and vertex are metallic blue, and thorax and abdomen are almost entirely black. Variation in males above includes the holotype; holotype was borrowed and its vesica spermalis was redrawn (Figures 12i, 13h) after cleaning some dry substance that had previously obscured the shape of the lobes in ectal view.

Diagnosis
Male of O. cultriformis has metallic purple frons and vertex, labial palp with medial black stripe as wide as 0.33-0.50 of palp width, pterothorax reddish brown with yellow stripes (Figure 2k), which can be entirely obscured in mature males, and dorsal terga S1-7 mostly red to mostly dark reddish brown to black, with pale yellow to greenish ventrolateral stripes on S1-3, orange to red stripes along medial longitudinal carinae on S4-8 and along ventrolateral carinae on S4-5 to S4-8. Abdomen narrows gradually from S2 to S4, and S4 is about as wide at level of apical carina as 0.35-0.50 of its length. Hamule inner and outer branches are located in the same plane ( Figure 9d), with outer branch bent ventrally over inner branch (Figure 10j), and outer corner of outer branch roundly angled (Figure 11j). Basal portion of distal segment of vesica spermalis is rhomboidal in ectal view (Figure 13j), distal portion has paired distal short medial longitudinal membranous lobes beset with denticles along margins, and paired outer lateral lobes are folded medially to the sides and are shorter than basal sclerotized portion of distal segment (Figure 12j). Cercus gradually curves ventrally ( Figure 15l) and its tip is slightly upturned. Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16k). Male of O. cultriformis shares combination of hamule with outer branch bent ventrally over inner branch ( Figure 10j) and epiproct apex relatively narrow, subapically less than 0.45 of its basal width (Figures 16k), (Figure 2k) to entirely dark in mature males, inner and outer branches of hamule separated by a distance longer than length of inner branch (Figure 11j), and vesica spermalis distal segment rhomboidal in ectal view (Figure 13j), versus pterothorax marbled (Figure 2v), inner and outer branches of hamule separated by a distance shorter than length of inner branch (Figure 11q), and vesica spermalis distal segment parallel sided in ectal view, Figure 13r). Outer surface of outer branch of hamule is smooth in O. cultriformis (Figures 10j, 11j), but grooved O. tambopatae and O. teres .
Female has labial palp with medial black stripe as wide as 0.33-0.40 of palp width, pterothorax reddish brown with yellow stripes, and abdomen narrowing gradually from S2 to S4, with S4 about as wide at level of apical carina as 0.35-0.46 of its length. Color of dorsal abdominal terga is similar to that of male. Ventrolateral flap on S8 is yellow, ca. 0.33-0.37 as wide as its length (Figure 17i

Remarks
The illustrations of the vulvar lamina and flap of S8 included in von Ellenrieder (2009, pp. 372, 373, figures 16f, 18d) were based on a misidentified female of O. aciculata; illustrations from specimens of O. cultriformis collected in copula with males are provided here (Figures 17i, 18k).
The levis group of Orthemis revisited 177

Distribution and biology
Orthemis cultriformis is a widespread species, occurring from Costa Rica in Central America south to Paraguay and N Argentina in South America (Figure 21). It is found near streams, rivers, and temporary ponds and swamps in the forest. sp. nov. Figures 1g,4f,6f,7i,8i,9e,10m,11l,12k,13k,14i,15n,16l,17j,18l,20 Etymology This species is named faaseni (noun in the genitive case) after Tim Faasen, who collected the first two specimens of this species that I examined and kindly allowed me to study and describe them.

Description of male holotype
Head. Prementum black; labial palps pale yellow with medial black stripe as wide as 0.33 of palp width and anterior margin narrowly black to the sides (as in Figure 1g); labrum yellow margined with black; base of mandibles and clypeus yellow; ventral half of antefrons pale brown medially and pale yellow laterally, dorsal portion of antefrons, postfrons, and vertex metallic blue; occipital triangle dark reddish brown; rear of head dark reddish brown with two yellow spots behind eye. Postfrons with wide shallow medial furrow; vertex with pair of low tubercles; posterior margin of occipital triangle slightly bilobate.
Thorax. Prothorax dark reddish brown, with anterior margin yellow. Pterothorax dark reddish brown to black, with diffuse whitish-blue pruinescence on dorsum and venter, and pale yellow stripes as follows (as in Figure 19k): wide stripe along posterior half of mesepimeron and stripe along posterior margin of metepimeron; venter of pterothorax brown with yellowish ill-defined medial longitudinal stripe along basal half, bifurcated into two diverging stripes on distal half. -Legs dark reddish brown, with tarsi and armature black; metafemur armed with 17 (left) to 18 (right) short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur (left hind tibia and tarsus missing). -Wings hyaline with amber apex. One cubito-anal crossvein in Fw and HW; sectors of arculus stalked; arculus between Anx 2-3 but closer to Anx 2, midway on left Hw; Fw triangles crossed, Hw triangles free; Fw subtriangles with 3 cells; one bridge crossvein in Fw and Hw; left Fw discoidal field with 3 rows of cells at base to 6 rows at hind margin ( Figures 4f, 6f), with no evident constriction between S3 and S4, S4 about twice as long as wide (ratio apical width/length = 0.52). Dorsal terga black, S2-8 with mid-dorsal carina pale yellow and margined by medial longitudinal pale yellow stripe; S1-3 with ventrolateral longitudinal pale yellow stripe; caudal appendages black. Ventrolateral carina on S2 and basal half of 3 vestigial, on distal half of S3 and S4-5 absent (as in Figure 6f), on S6-8 well developed, on S9 incomplete, extending along basal 0.66 of segment. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (as in Figure 6f) dark reddish brown (S2-4) to black (S5-10), S1-2 with pale yellow stripe along ventrolateral carinae; S3-5 with yellow stripe along lateral margin on basal two thirds (S3) to basal half (S4-5) of segment. Anterior lamina in lateral view about a third of hamule height and about as high as genital lobe (as in Figure 10m); hamule bifid with longer inner branch forming pointed hook directed anteriorly, surpassing outer branch ventrally (as in Figure 11l); outer branch tip separated from inner branch by distance about as long as inner branch length (as in Figure 9e); outer corner of hamule outer branch rounded (as in Figure 11l). Distal segment of vesica spermalis with basal portion trapezoidal in ectal view, with sides parallel along basal half and converging to tip along distal half (Figure 13k), and with long flagella on ental surface; distal lobes represented on each side by subtriangular membranous outer lobe with transverse lateral fold, shorter than basal portion of distal segment in lateral view, and medial longitudinal sclerotized spatulate lobe longer than outer lobe, provided with a lateral longitudinal ridge, slightly expanded dorsoventrally at apex, and margined ventroapically with denticles ( Figure 12k). Cercus markedly curved ventrally at base in lateral view, with row of 6 ventral tubercles along distal half and tip upturned (as in Figure 15m); in dorsal view about parallel sided to opposite cercus with tips converging (as in Figure 14i). Epiproct extending to 0.74 of cercus length, with apex slightly bifid and about as wide as 0.4 of its maximum width (as in Figure 16l).

Variation in male paratype
Head. As for holotype but occipital triangle black.
Abdomen. As for holotype but ventrolateral carina on S3 vestigial, and absent on S4-5 but its position indicated by row of denticles ( Figure 6f); cercus with row of 5 ventral tubercles along distal half.
The levis group of Orthemis revisited 179

Variation in female paratype
Head. As for holotype but clypeus pale brown; antefrons yellow laterally, reddish brown medially; postfrons reddish brown; vertex reddish brown with metallic blue reflections; occipital triangle dark reddish brown; rear of head reddish brown.
Thorax. As for holotype but lacking any pruinescence; mesepisternum with yellow medial longitudinal stripe adjacent to mediodorsal carina and yellow longitudinal stripe parallel to mesepisternal-mesepimeral carina ( Figure 19l); metafemur armed with 17 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings hyaline, with infumated apex; arculus between Anx 2-3 but midway in Hw; Fw discoidal field with 6 cells at hind margin, Hw with 15-16; anal loop enclosing 26 cells. Pt 4.6 long in Fw and 4.5 in Hw, overlying 5-6 cells in Fw and 6 cells in Hw. [15][16]18 in Hw.

Diagnosis
Male of O. faaseni differs from all known species of Orthemis by ventrolateral carina on S4-5 absent (Figures 4f, 6f) and distal segment of vesica spermalis with medial longitudinal sclerotized spatulate lobe longer than outer lobe (Figure 12k). Within the levis group, it approaches O. harpago by hamule shape, with inner branch longer than outer branch, forming a pointed hook directed anteriorly, surpassing outer branch ventrally (Figures 9e-f, 10m-n, 11l-m). It differs from O. harpago by narrower epiproct apex, as wide as 0.4 of epiproct maximum width (Figure 16l; versus as wide as 0.5-0.6 of maximum width, Figure 16o), abdomen narrowed gradually at base, with base of S4 more than half as high as base of S3 in lateral view (Figure 4f; versus narrowed abruptly, with base of S4 less than half as high as base of S3, Figure 4i), and S4 relatively shorter, with S4 width/length ratio of 0.52 (Figure 6f; versus longer, with S4 width/length ratio of 0.19-0.23, Figure 6i

Type specimens examined
Lectotype : Brazil, "Pará" [BMNH]. Paralectotypes: 3 same data as lectotype [BMNH].  Thorax. Prothorax dark reddish brown. Pterothorax dark brown with yellow stripes as follows: medial longitudinal stripe adjacent to mediodorsal carina; longitudinal stripe parallel to mediodorsal carina at mesepisternum mid width (as in Figure 2m); stripe along posterior half of mesepimeron narrowing dorsally; stripe along ventral margin of metepisternum and another sinuous one ventral to metastigma, and oval spot on posterodorsal corner; metepimeron with stripe along posterior half, narrowing dorsally, and subtriangular spot on anterodorsal corner ( Figure 2l); venter of pterothorax brown with pale yellow medial longitudinal stripe along basal half, bifurcated into two diverging stripes on distal half. -Legs with coxa pale yellow, trochanter of foreleg pale yellow, of middle and hind leg reddish brown; femur of foreleg pale yellow along posterior half, remainder reddish brown, of middle and hind leg reddish brown; tibia dark reddish brown; The levis group of Orthemis revisited 183 tarsus, pretarsus, and spines black; metafemur armed with 19 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings hyaline with tip amber from level of distal end of pterostigma to apex. One cubito-anal crossvein in Fw and HW; sectors of arculus stalked; arculus between Anx 2-3 but closer to Anx 2 in right FW, opposite to Anx 2 in left FW, midway between Anx 2-3 in HW; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4 cells; one bridge crossvein in Fw and Hw; Fw discoidal field with 3 rows of cells at base to 5 rows at hind margin, Hw with 3 rows of cells at base, then 2, then increasing to 15 at hind margin; 3 rows of cells between wing margin and anal loop at level of anal angle of triangle; anal loop enclosing 27 cells. Pt reddish brown, 5.0 long in Fw and 4.9 in Hw, overlying 6 cells in Fw and 5 cells in Hw. Anx: 17 in Fw, 14 (right) and 15 (left) in Hw; Pnx: 15 (right) and 17 (left) in Fw; 17 in Hw.
Abdomen. Sides linear, parallel sided in dorsal and ventral view (Figures 4g, 6g), with no evident constriction between S3 and S4, S4 almost two times as long as wide (ratio apical width/length = 0.53). S1-7 dorsal terga dark reddish brown (S1-4) to dark brown (S5-7) with mid-dorsal carina pale yellow and margined by medial longitudinal pale yellow stripe, and ventrolateral longitudinal pale yellow (on S1-3) to pale orange (on S4-7) stripe; S8 black with ventrolateral longitudinal pale orange stripe; S9-10 black; caudal appendages dark reddish brown. Ventrolateral carina on S2 vestigial, on S3-9 well developed and black, on S9 incomplete, extending along basal 0.75 of segment. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 6g): S1 and S2 reddish brown with yellow along ventrolateral carinae; S3-4 reddish brown, S5-8 dark reddish brown, with yellowish stripe along lateral margin on basal two thirds of segment; S9-10 dark reddish brown. Anterior lamina in lateral view slightly shorter than hamule and about as high as genital lobe ( Figure 10k); right hamule (left hamule missing) bifid with small inner branch forming short pointed spine directed posteriorly, in the same plane as and larger than outer branch; tip of outer branch separated from inner branch by distance about as long as inner branch length; outer corner of hamule roundly angled (Figure 11o). Distal segment of vesica spermalis with basal portion parallel sided in ectal view (Figure 13l), and with pair of long flagella on ental surface; distal portion represented on each side by outer lobe, with sclerotized subtriangular part about as long as basal portion of distal segment in lateral view, folded medially into transverse membranous part, approximately oval in ectal view, as long as half of basal portion or shorter, and medioectal sclerotized lobe with medial ridge margined with denticles laterally; medioectal lobe about as long as half the length of outer lobe (Figure 12l). Right cercus (left cercus missing) in lateral view markedly curved ventrally at base and with tip not upturned; distal dilated portion with row of 6 ventral tubercles along its base ( Figure 15n); in dorsal view converging along basal 0.66 with opposite cercus, then approximately parallel to it along distal 0.33 (Figure 14j). Epiproct extending to 0.77 of cerci length, with apex entire and about as wide as a third of its basal width (wooden probe inserted in abdomen precludes an exact measurement). Dimensions. Total length 50.1; abdomen length 32; Fw length ca. 41 (apical cells missing); Hw length 36.97; Hw maximum width 11.5; cercus length 2.2; epiproct length 1.7; epiproct maximum width 1.2.

Variation in males
Head. As for lectotype but labial palp medial black stripe as wide as 0.3-0.4 of palp width; postclypeus pale brown with postclypeal lobes narrowly fringed with dark brown line along distal margin in one male; frons and vertex metallic purple lacking blue reflections in one male, and metallic blue lacking purple reflections in two males; occipital triangle dark reddish brown to reddish brown dorsally, yellow posteroventrally; rear of head reddish brown, with two yellow spots behind eye.
Thorax. As for lectotype but prothorax reddish brown with anterior margin of anterior lobe pale; yellow longitudinal stripe parallel to mesepisternal-mesepimeral carina confluent at anterior 0.20 to narrow yellow stripe along mesepisternal-mesepimeral carina in one paratype; bluish white pruinescence on mesepisternum and venter of pterothorax in two males; mesepisternum entirely covered by pruinescence rendering pale stripes invisible in some paratypes (Figures 2n, 19m); metafemur armed with 16-19 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur; Fw subtriangles with 3 cells in one wing, with 5 cells in one wing; Fw discoidal field with 3-4 rows of cells at base to 4-7 rows at hind margin, Hw with 3 rows of cells at base, then 2, then increasing to 13-18 at hind margin; anal loop enclosing 21-27 cells. Pt overlying 4-6 cells in [12][13][14] in Fw, 14-18 in Hw.

Variation in females
Head. As for lectotype but labrum yellow margined with black; palp medial black stripe as wide as 0.33-0.4 of palp width; dorsal portion of antefrons, postfrons, and vertex pale brown; occipital triangle dark brown; rear of head dark reddish brown with two yellow spots behind eye.
Abdomen. As for lectotype but ventrolateral carina on S9 absent; dorsal tergum S8 black with mediodorsal carina narrowly yellow; ventrolateral flap on S8 pale yellow (Figure 19n). Vulvar lamina extending along posterior 0.20 of S8 length, forming three-dimensional raised ridge shaped as an inverted U, rounded in cross-section (Figure 18m). Caudal appendages dark reddish brown.  (Figure 12l; versus bearing a strong sclerotized lateral spine in O. garrisoni, Figure 12m). It differs from O. regalis by wings lacking dark markings along C-RA space, smaller size (Hw < 45 mm), presence of three rows of cells between anal margin of hind wing and AA2 at level of heel of anal loop, outer corner of outer branch of hamule not triangular and projected ventroposteriorly in lateral view (Figure 10k), and basal sclerotized portion of vesica spermalis ca. parallel sided in ectal view ( Figure 13l); versus wings with dark markings on C-RA space, larger size (Hw > 50 mm), four rows of cells between anal margin of hind wing and AA2 at level of heel of anal loop, outer corner of outer branch of hamule triangular and projected ventroposteriorly in lateral view (Figure 10r), and basal sclerotized portion of vesica spermalis deeply constricted medially in ectal view (Figure 13q) in O. regalis.
Female shares a similar vulvar lamina consisting of an inverted U-shaped ridge anterior to posterior margin of S8 with females of O. ambinigra and O. garrisoni. It differs from O. ambinigra by the ridge being swollen, rounded in cross-section (Figure 18m; versus thin and laminar, Figure 18c), and from O. garrisoni by the narrower transverse bar of the inverted "U" ridge, shorter than the longitudinal arms of the ridge (Figure 18m; versus wider transverse bar of the inverted "U" ridge, as long as the longitudinal arms of the ridge, Figure 18n).

Remarks
Orthemis flavopicta Kirby, 1889 was described based on a series of four males and one female from "Pará", with no further data. Kimmins (1969) designated the male bearing Kirby's type label as lectotype but did not provide any descriptive notes for it. Kirby's (1889) description stated that the epiproct was "broad and bifid, or rather crescent-shaped at the apex"; however, examination of the type series shows the epiproct to be narrow at tip in all four males. All type specimens have been preserved with a thin piece of wood inserted through the abdomen and extruding from sternum S8 blocking a full ventral view of caudal appendages. The lectotype has a partially crushed mesepisternum, for which reason I selected a paralectotype male to illustrate the dorsal view of the thorax. Vesica spermalis of lectotype was extracted and placed on a separate pin by Buchholz, and I extruded the vesica spermalis -without detaching it -from all paralectotypes as well for comparison. I believe that the female included by Kirby in the type series of O. flavopicta does not correspond to this species, because it differs from females found flying together with males of O. flavopicta in both Pará and Bahia which closely match the male of O. flavopicta based on color pattern and size. Kirby's female instead agrees well with females of O. aciculata found flying with males of that species in Trinidad and Surinam.
Examination of the lectotype of O. flavopicta showed that specimens from Panama identified and depicted as O. flavopicta in Garrison (1984) andvon Ellenrieder (2009) (Calvert 1899), and consequently I list his publication including this species in the synonymic list as "possibly in part".
Seemingly only in O. flavopicta does the membranous area on inner surface of hamule extend along the entire height of the hamule (Figure 11o); in all the specimens of other species that I examined, the area that remains membranous seems to be limited to the distal portion of inner surface of the hamule, between inner and outer branches, and the degree of intraspecific variability of the distance between both branches and their position relative to the base of the hamule is very low in comparison to that observed in O. flavopicta.

Etymology
This species is named garrisoni (noun in the genitive case) after my dear husband Rosser W. Garrison, who collected the first specimens of this species which I was able to examine, in recognition of his constant support to all students of neotropical odonates and his numerous outstanding contributions to the knowledge of New World odonate taxonomy.

Description of male holotype
Head. Prementum black; labial palps pale yellow with medial black stripe as wide as 0.33 of palp width anteriorly extending to the sides along almost entire anterior margin of palp ( Figure 1i); labrum black with pair of rounded laterobasal pale yellow spots; base of mandibles yellow; lateral portion of clypeus along eyes whitish, anteclypeus pale brown, postclypeus pale brown with medial dark brown spot and postclypeal lobes narrowly fringed with dark brown line along distal margin; ventral half of antefrons pale brown medially and whitish laterally, dorsal portion of antefrons, postfrons, and vertex metallic blue with purple reflections, occipital triangle dark reddish brown, rear of head reddish brown, with two yellow spots behind eye. Postfrons with wide shallow medial furrow; vertex with pair of low tubercles; posterior margin of occipital triangle slightly bilobate.
Thorax. Prothorax reddish brown with anterior lobe anterior margin pale yellow and medial lobe covered with bluish white pruinescence. Pterothorax dark reddish brown, with mesepisternum, antealar sinus, and terga covered with bluish white pruinescence, and pale yellow stripes as follows (Figure 2p): diffuse mesepisternal stripe adjacent to mesepisternal-mesepimeral carina; wide stripe along posterior half of mesepimeron narrowing dorsally; narrow stripe ventral to metastigma, and oval spot on posterodorsal corner; metepimeron with stripe along posterior margin and subtriangular spot on anterodorsal corner; venter of pterothorax brown with yellowish ill-defined medial longitudinal stripe along basal half, bifurcated into two diverging stripes on distal half. -Legs with base of coxa pale yellow, apical portion of coxa, trochanter, and posterior surface of fore and mid leg pale reddish brown, remainder dark reddish brown; tarsus, pretarsus, and spines black; metafemur armed with 22 (right) or 18 (left) short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings deeply infumated in gold. One cubito-anal crossvein in Fw and HW; sectors of arculus stalked; arculus between Anx 2-3 but closer to Anx 2; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4 cells; one bridge crossvein in Fw and Hw; Fw discoidal field with 3 rows of cells at base to 6 rows at hind margin, Hw with 3 rows of cells at base, then 2, then increasing to 18 at hind margin; 3 rows of cells between wing margin and anal loop at level of anal angle of triangle; Abdomen. Sides almost parallel in dorsal and ventral view (Figures 4h, 6h), with no evident constriction between S3 and S4, S4 two times as long as wide (ratio apical width/length = 0.5). S1-7 dorsal terga dark reddish brown (S1-4) to dark brown (S5-7) with mid-dorsal carina pale yellow and margined by medial longitudinal pale yellow stripe, and ventrolateral longitudinal pale yellow stripe; S8 black with ventrolateral longitudinal pale yellow stripe; S9-10 black; caudal appendages dark reddish brown. Ventrolateral carina on S2 vestigial, on S3-9 well developed and black, on S9 incomplete, extending along basal 0.66 of segment. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 6h): S1 and S2 reddish brown with yellow along ventrolateral carinae; S3-4 reddish brown, S5-8 dark reddish brown, with yellowish stripe along lateral margin on basal two thirds of segment; S9-10 dark reddish brown. Anterior lamina in lateral view slightly shorter than hamule and about as high as genital lobe ( Figure 10l); hamule bifid with smaller inner branch forming short pointed spine directed posteriorly, in the same plane as larger outer branch; outer branch tip separated from inner branch by distance about as long as inner's branch length; outer corner of hamule roundly angled (Figure 11p). Distal segment of vesica spermalis with basal portion parallel sided in ectal view (Figure 13m), and with pair of long flagella on ental surface; distal portion represented on each side by outer lobe, with sclerotized oval portion about as long as basal portion of distal segment in lateral view, folded medially into transverse membranous portion, approximately oval in ectal view, as long as half of basal portion or shorter, and medioectal sclerotized lobe with strong lateral spine directed anteroventrally on each side; medioectal lobe about as long as 0.75 of the length of outer lobe (Figure 12m). Cercus in lateral view markedly curved ventrally at base and with tip slightly upturned; distal dilated portion with row of 7 ventral tubercles along its base ( Figure 15p); in dorsal view converging along basal 0.66 with opposite cercus, then approximately parallel to it along distal 0.33 (Figure 14k). Epiproct extending to 0.78 of cerci length, with apex entire and about as wide as a third of its basal width (Figure 16n).  190 N. von Ellenrieder Variation in male paratypes Head. As for holotype but medial black stripe on labial palp as wide as 0.3 of palp width in four paratypes; lateral portion of clypeus along eyes, ventral half of antefrons laterally, and postclypeus along clypeal-frontal suture pale yellow in two paratypes; vertex with yellow spot from ocellus to medial line across posterior surface in one paratype; occipital triangle with pair of yellow spots posteroventrally in one paratype.
Thorax. As for holotype but mesepisternum with narrow medial longitudinal stripe adjacent to mediodorsal carina visible in one paratype, and with narrow longitudinal pale yellow stripe parallel to mediodorsal carina at mesepisternum mid width visible in five males with weak pruinescence; metafemur armed with 14-23 short spurs which slightly and gradually increase in size towards apex, followed distally by one, or rarely two, longer spurs. -Wings infumated only slightly in some paratypes; arculus between Anx 2-3 but midway in some wings; Fw discoidal field with 3 rows of cells at base to 4-7 rows at hind margin, Hw with 3 rows of cells at base, then 2, then increasing to 14-33 at hind margin; anal loop enclosing 20-24 cells. Pt overlying 5-7 cells in Fw and 4-6 cells in Hw. [13][14][15][15][16][17][18] Abdomen. As for holotype but with slight constriction between S3 and S4 in three paratypes; apical width/length S4 0.31-0.56; tip of outer branch of hamule separated from tip of inner branch by distance shorter than inner branch length in five paratypes ( Figure 11p); S9-10 reddish brown laterally in two males; with row of 6-8 ventral tubercles along distal half of male cercus; tip of cercus not upturned in three paratypes.

Variation in female paratypes
Head. As for holotype but labrum reddish brown with black anterior margin and a pair of lateral pale yellow spots; postclypeus yellow with postclypeal lobes narrowly fringed with a brown line along distal margin; antefrons yellow laterally, reddish brown medially; postfrons reddish brown with a small yellow rounded spot anterolateral to base of antenna, vertex reddish brown, dark reddish brown around lateral ocellus, with a yellow spot from ocellus to medial line across posterior surface; occipital triangle reddish brown dorsally, yellow posteroventrally.
Thorax. As for holotype but lacking any bluish white pruinescence; mesepisternum with yellow medial longitudinal stripe adjacent to mediodorsal carina and yellow longitudinal stripe parallel to mesepisternal-mesepimeral carina confluent at anterior 0.20 to narrow yellow stripe along mesepisternal-mesepimeral carina ( Figure 2q); metafemur armed with 12-17 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings hyaline, with infumated apex; arculus between Anx 2-3 but midway in some Hw; Fw discoidal field with 3 rows of cells at base to 5-7 rows at hind margin, Hw with 3 rows of cells at base, The levis group of Orthemis revisited 191 then 2, then increasing to 15-17 at hind margin; anal loop enclosing 20-23 cells. Pt overlying 5-6 cells in Fw and 4-6 cells in Hw. [12][13][15][16] in Hw.
Abdomen. As for holotype but ventrolateral carina on S9 absent. Ventrolateral flap on S8 pale yellow (Figure 17l). Vulvar lamina extending along posterior 0.20 of S8 length, consisting of an inverted U-shaped ridge, three-dimensional, rounded in cross-section (Figure 18n), with transverse bar of the inverted U-shaped ridge as long as longitudinal arms of the ridge. Appendages dark reddish brown.

Remarks
This species was misidentified as O. flavopicta by Garrison (1984) and von Ellenrieder (2009) (Figure 18n; versus narrower transverse bar of the inverted U-shaped ridge, shorter than the longitudinal arms of the ridge, Figure 18m).

N. von Ellenrieder Distribution and biology
This species is so far known only from Panama (Figure 20), where adults were found along forested streams and trails. It seems to be the northern counterpart to its nearest ally, O. flavopicta, being allopatric with that species.

Type specimens examined
Thorax. Prothorax reddish brown except anterior lobe, mediodorsal area of middle lobe, and posterior margin of posterior lobe yellow. Pterothorax (Figure 2s) dark reddish brown with pale yellow stripes as follows: mesepisternum with medial longitudinal stripe adjacent to mediodorsal carina, longitudinal stripe parallel to mesepisternal-mesepimeral carina, and two spots at midlength and at distal fourth of mesepisternal-mesepimeral carina; mesepimeron with wide stripe along posterior half narrowing dorsally; metepisternum with narrow stripe along ventral margin and another sinuous one ventral to metastigma; metepimeron with stripe along posterior half, narrowing dorsally, and oval spot on anterodorsal corner; venter of pterothorax pale yellow with lateral margins from base of leg to posterior margin dark reddish brown. -Posterior surface of coxa, trochanter, and femur of front and middle leg and posterior surface of coxa of hind leg pale yellow, remainder dark reddish brown; tibia, tarsus, pretarsus, and spination black; metafemur armed with 15-17 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings hyaline, amber from base to level of second row of anal cells in FW and to first Anx in HW and at apex across distal three rows to distal end of pt. One cubitoanal crossvein and arculus between to Anx 2 and Anx 3, closer to Anx 2 in Fw, midway in Hw; sectors of arculus stalked; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4-5 cells; one bridge crossvein; Fw discoidal field with 3 rows of cells at base to 5-6 rows at hind margin, Hw with 2-3 at base, then 2, then increasing to 15 at hind margin; 3 rows of cells between wing margin and anal loop at level of anal angle of triangle; anal loop enclosing 23-24 cells. Pt reddish brown, overlying 4-6 cells. [14][15][15][16][17] Abdomen. Base swollen, narrowing abruptly from S3 to S4 (Figure 7l), S4 ca. three times as long as wide (ratio apical width/length = 0.34). Dorsal terga with mid-dorsal carina bordered by yellow on S1-7; S1-3 reddish brown with ventral third yellow; S4 reddish brown anteriorly to black posterodorsally, with orange stripe along ventrolateral carina; S5-7 black with orange stripe along ventrolateral carina; S8 black with ventrolateral flap yellow; S9-10 and caudal appendages black. Ventrolateral carina on S2 vestigial, on S3-7 well developed and black. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga (Figure 8l) dark reddish brown with yellow to orange stripe along ventrolateral carinae on S1-7; S8 pale brown; S9-10 dark reddish brown. Ventrolateral flap on S8 pale yellow (Figure 17m Figure 11a, d, f, h), but differs from it by inner branch longer than outer branch (Figure 11m; versus ca. as long as outer branch, Figure 11k). It further differs from them except for O. cinnamomea by abdomen narrowing abruptly at base (Figure 4l; versus narrowing more gradually, Figures 4a-b , 19a, c-d). Mature males can be separated further from those of O. cinnamomea by frons and vertex metallic purple with blue reflections (versus reddish brown).
Female shares a similar vulvar lamina, characterized by the presence of a medial longitudinal ridge anterior to posterior margin of sternum S8, with O. anthracina, O. aciculata, O. cinnamomea, O. coracina, O. cultriformis, O. faaseni, and O. flavopicta Figures 17j, 19l), and from O. cinnamomea by S8 lateral flap relatively narrower, as wide as 0.29 of its length, and ca. smoothly convex (Figure 17m; versus S8 flap relatively wider, as wide as 0.31-0.34 of its length, and triangular in contour, Figure 17f). It can be distinguished from O. coracina by abdomen narrowing abruptly at base, with base of S4 less than half as high as base of S3 in lateral view (Figure 7l; versus narrowing gradually, with base of S4 more than half as high as base of S3 in lateral view, Figure 7h).

Remarks
I borrowed the holotype for direct comparison with the new species described in this paper, and provide here some further illustrations not included in its original description (Figures 6i, 14l). One additional specimen of O. harpago from Rondônia State, Brazil was found among the examined material. Shape of hamule, vesica spermalis, abdominal ventral terga contour, cerci, and epiproct fully agree with those of holotype and paratype. As indicated by its milky wing membranes and partially crushed head and abdomen, this specimen is a teneral male, much paler in color than holotype and paratype, with thorax pale brown with pale yellow stripes, and dorsal surface of frons and vertex brown with metallic purple and blue reflections. Its size is slightly larger than that of the types: total length 43; abdomen length 27.5; Fw length 33; Hw length 32.4; maximum Hw width 8.8.
I identify the female from Loreto, Peru, as belonging to O. harpago by supposition. Its size is larger than that of the three known males of this species, but similar sexual dimorphism in size is known for some other species ( This female could also belong to an undescribed species close to O. harpago, but I prefer to be conservative and treat it here as the female of O. harpago. Finding more specimens of O. harpago, especially pairs in copula, and DNA bar-coding of both males and females, will eventually confirm or reject this association.

Diagnosis
Male of O. levis has metallic purple frons and vertex, labial palp with medial black stripe as wide as 0.33-0.40 of palp width, pterothorax reddish brown with yellow stripes, which can be obscured in mature males, dorsal terga S1-7 mostly red, with ventrolateral and posterior carinae narrowly black, and S8-10 dorsally black. Abdomen narrows abruptly from S2 to S4, and S4 is about as wide at level of apical carina as 0.27-0.37 of its length. Hamule inner and outer branches are located in the same plane, with both ending at about the same level ( Figure 10o); inner corner of outer branch is smoothly rounded (Figure 11n). Distal segment of vesica spermalis is ca. oval, with sides gradually converging to distal end in ectal view (Figure 13o), paired distal short medial longitudinal membranous lobes beset with denticles along distal margin on ectal side, paired outer lateral lobes shorter than basal sclerotized portion of distal segment folded medially to the sides, and paired flagella (Figure 12o). Cercus curves ventrally gradually and its tip is slightly upturned. Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16p). Male of O. levis shares combination of abruptly narrowed abdomen at base and epiproct tip narrow and entire only with O. attenuata and O. concolor. It differs from both by its red abdominal color pattern, with only S8-10 dorsally black (versus S4-7 or S5-7 mostly black with narrow medial longitudinal and ventrolateral yellow to orange stripes), and by shape of abdomen base between apex of S2 and base of S3, which is less globose in O. levis (versus more globose,Figures 4d,. It further differs from O. concolor by metallic purple frons and vertex and pterothorax with yellow stripes, versus frons and vertex reddish brown and pterothorax concolorous reddish brown (Figure 19g).
Female shares with male: labial palp with narrow medial black stripe (0.25-0.40 of palp width), pterothorax reddish brown with yellow stripes, and slender abdomen narrowing abruptly from S2 to S4 (Figure 7m). Color of dorsal abdominal terga is similar to that of male. Ventrolateral flap on S8 (Figure 17n) is yellow and relatively narrow, ca. 0.22-0.28 as wide as its length, with a smoothly curved contour. Sternum S8 is smooth, with vulvar lamina consisting of posterior margin of sternum of S8 forming a shallow inverted U-shaped rim, with medial excision narrower than its lateral margins (Figure 18p). Combination of relatively slender abdomen narrowing abruptly at base, with base of S4 less than half as high as base of S3 in lateral view, and vulvar lamina consisting of shallow U-shaped rim of S8 posterior margin is shared only with O. attenuata and O. concolor. Orthemis levis differs from both by red color present on abdominal dorsal terga S4-7 and by shape of the abdomen base between the apex of S2 and the base of S3, which is less globose in O. levis (Figure 7m; versus more globose, Figure 7e, g). It further differs from O. concolor by pterothorax with yellow stripes and by paired depressions anterior to vulvar lamina rim located directly anterior to lateral margins of rim (Figure 18p; versus located laterally and outwards from lateral margins of rim in O. concolor, Figure 18i). Since red color on abdominal dorsal terga S4-7 is not as pronounced as it is in the males, being sometimes replaced by reddish brown on S1-4 to S1-7, and since unlike in males, there are yellow to orange paler stripes along mediodorsal and ventrolateral carinae of S1-7 to S1-8, poorly preserved females especially can look very similar in abdominal color to those of O. attenuata and O. concolor.

Remarks
Among the examined material, specimens from South America seem to have in general a more slender abdomen than those of North America, but there are intermediates in Central America, The levis group of Orthemis revisited 197 and I was unable to find any morphological differences justifying the recognition of two species or subspecies, and treat this variability as geographical variation.

Etymology
This species is named paulsoni (noun in the genitive case) after Dennis R. Paulson, who lent me the first specimens I examined, in recognition of his always available kind advice and encouragement to me and to other numerous odonate students and of his invaluable contributions to the knowledge of New World odonate diversity.

Male holotype
Head. Labium black, with palps pale yellow with medial black stripe as wide as ca. 0.40 of palp width and anterior margin narrowly black ( Figure 1j); labrum black with mediobasal reddish spot; base of mandibles yellow; lateral portion of clypeus along eyes pale yellow, remainder of anteclypeus pale brown, postclypeus pale brown with medial dark brown spot and postclypeal lobes narrowly fringed with dark brown line along distal margin; ventral half of antefrons pale brown medially and pale yellow laterally, dorsal portion of antefrons, postfrons, and vertex metallic blue with purple reflections, occipital dark reddish brown, rear of head reddish brown with two pale yellow spots behind eye. Postfrons with wide shallow medial furrow; vertex with pair of low tubercles; posterior margin of occipital triangle slightly bilobate.
Thorax. Prothorax reddish brown except anterior lobe margined with pale yellow. Dorsum and sides of pterothorax (Figure 2t) entirely covered with bluish white pruinescence, some exposed areas dark reddish brown, with pale yellow stripe along posterior half of metepimeron partly evident; venter of pterothorax reddish brown. -Legs with coxa, trochanter, and inner surface of femur pale reddish brown; remainder of femur reddish brown basally to dark reddish brown distally; tibia dark reddish brown; tarsus, pretarsus, and spines black; metafemur armed with 10 (right) to 11 (left) short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Wings (as in Figure 3a) slightly infumated. One cubito-anal crossvein and arculus opposite to Anx 2 in Fw and HW; sectors of arculus stalked; Fw triangles crossed, Hw triangles free; Fw subtriangles with 4 cells; two bridge crossveins in Fw and Hw; Fw discoidal field with 3 rows of cells at base to 5 rows at hind margin, Hw with 2 rows of cells at base, increasing to 14 (right) or 15 (left) at hind margin; 4 rows of cells between wing margin and anal loop at level of anal angle of triangle; anal loop enclosing 20 (right) cells (incomplete on left wing). Pt reddish brown, 4.7 long in Fw and 4.5 in Hw, overlying 5 cells in Fw and 4 cells in Hw. Anx: 16 (right) to 17 (left) in Fw; 12 (right) to 13 (left) in Hw; Pnx: 15 in Fw and Hw.
Abdomen. Gradually narrowing from base to apex in ventral view (Figure 4j), S4 slightly longer than wide (ratio apical width/length = 0.85). Dorsal terga: S1 and basal half of S2 pale reddish brown; distal half of S2 and S3 dark red; S4-9 bright red with mid-dorsal, ventrolateral, and posterior carinae margined in black, and with small diffuse laterodistal black spot; S10 dark reddish brown with mediodorsal dark red spot; caudal appendages dark reddish brown. Ventrolateral carina on S2 vestigial, on S3-8 well developed, on S9 well developed along basal two thirds, absent at distal third. Transverse carina well developed on S2-3, vestigial on S4-5. Ventral terga reddish brown, with orange pale lateral longitudinal stripes along outer two-thirds of S2-8 (Figure 6j). Anterior lamina in lateral view shorter than hamule and as high as genital lobe ( Figure 10p); hamule bifid with small inner branch forming short pointed spine, in same plane as rounded outer branch, separated by distance shorter than inner branch length (Figure 11q). Distal segment of vesica spermalis with basal portion approximately rectangular in ectal view, widest at apex, with sides concave near apex, and with long flagella on ental surface ( Figure 13p); distal portion represented on each side by ental lobe, with sclerotized subtriangular portion shorter than basal portion of distal segment in lateral view, folded medially into transverse membranous portion, approximately oval in ectal view, about as long as half of basal portion, and medioectal membranous lobe with lateral longitudinal sclerotized ridge covered with denticles; medioectal lobe almost as long as outer lobe (Figure 12p). Cercus slightly curved ventrally in lateral view, with row of 7 (left) to 8 (right) ventral tubercles along venter of dilated portion and tip slightly upturned ( Figure 15s); in dorsal view converging along basal 0.70 with opposite cercus, then approximately parallel to it along distal 0.30 (Figure 14m). Epiproct extending to 0.82 of cercus length, with apex slightly concave and as wide as 0.4 of its basal width (as in Figure 16q).

Variation in male paratypes
Head. As for holotype but labial palp medial black stripe as wide as about 0.33 of palp width in one paratype.

Variation in female paratypes
Head. As for holotype but medial black stripe as wide as ca. 0.33-0.40 of palp width labrum with basal pale yellow spot on medial half; antefrons medially and postfrons, vertex, and occipital triangle reddish brown, with dark blue metallic reflections along postfrons medial furrow and surrounding the ocelli on vertex.
Thorax. As for holotype but pruinescence extending onto venter of pterothorax, and thin on dorsum and sides, making pale diffuse yellowish stripes visible: narrow stripe along mid-dorsal carina, and wide stripes along mid width of mesepimeron, posterior edge of mesepimeron, and posterior edge of metepimeron ( Figure 2u). Metafemur armed with 10-13 short spurs which slightly and gradually increase in size towards apex, followed distally by one longer spur. -Arculus between Anx 2-3, closer to Anx 2; one paratype with 1 bridge crossvein on left wings, one paratype with 1 bridge crossvein on right wings and left Hw; Fw discoidal field with 4-7 cells at hind margin, Hw with 14-19; anal loop enclosing 19-23 cells. Pt overlying 5-6 cells in Fw and 4-7 cells in Hw. Anx: 15-18 in Fw, 12-14 in Hw; Pnx: 13-15 in Fw, 15-16 in Hw.
Abdomen. As for holotype but ratio S4 apical width/length = 0.89-1.0 (Figure 8m). Ventrolateral carina on S4-5 present, on S9 absent. Distal black spot on dorsum of S8 extended to posterior margin, on S9 forming a complete anteroposterior black stripe ( Figure 7n); S10 dark reddish brown. Ventrolateral flap on S8 yellow narrowly margined in black (Figure 17o). Vulvar lamina consisting of posterior margin of S8 forming a rim with a deep medial inverted U-shaped excision and paired depressions under convex lateral margins (Figure 18r).

Diagnosis
Orthemis paulsoni can be diagnosed from all species in the levis group by its relatively wider abdomen (Figures 6j, 8m, 19p), with S4 apical width/length ratio of 0.59-0.85 in males and 0.89-1.0 in females; versus abdomen more slender, with S4 apical width/length ratio of 0.19-0.56 in males and 0.3-0.57 in females (Figures 6a-i Within the ferruginea group, male of O. paulsoni is unique by its cerci markedly curved ventrally (Figure 15s-t). In other known species of this group cerci curve ventrally only slightly (i.e. Figure 15g). Orthemis paulsoni resembles O. biolleyi by combination of bright red abdomen and outer corner of male hamule not triangular and projected posteriorly (Figure 11g, q), but it can easily be distinguished from it by male postfrons and vertex metallic blue versus metallic purple in O. biolleyi, and pterothorax pruinescent blue in mature males and bearing four pale yellowish stripes in females (probably also in teneral males), versus with six bright yellow stripes in males and females of O. biolleyi. Female of O. paulsoni can be diagnosed from all species in the ferruginea group by rim of vulvar lamina with a deep inverted U-shaped medial excision and paired depressions under the convex lateral margins (Figure 18r), versus rim of vulvar lamina lacking a deep medial inverted U-shaped excision, either ca. straight, shallowly concave, or slightly bilobate (Figure 18f, q, t).

Remarks
I mistakenly treated and illustrated this species as O. plaumanni in my previous analysis of the levis group of Orthemis (von Ellenrieder, 2009). Buchholz (1950) diagnosed O. plaumanni from other species in this group solely on the basis of presence of two bridge crossveins in the wings, and all the wings of the examined males and some wings of the examined females of O. paulsoni have two bridge crossveins. Even though the two specimens of O. paulsoni from Peru available to me earlier seemed to fit the description of O. plaumanni well, the shape of the hamule differed from that illustrated by Buchholz (1950) for that species. In order to confirm the identification I borrowed the holotype of O. plaumanni from ZFMK, and its examination revealed that O. plaumanni is a junior synonym of O. ambinigra, and that O. paulsoni is in fact a new species.
Additional specimens of O. paulsoni from Ecuador examined here showed that the abdomen in this species is wider than in all species of the levis group, pointing to the correct placement of this species in the ferruginea group rather than in the levis group. This is also indicated by the presence of four rows of cells between anal margin of hind wing and AA2 at level of heel of anal loop (Figure 3a) in most wings (in two of the males and one female one of the Hw has three rows

Diagnosis
Male of O. philipi has metallic blue frons and vertex, labial palp with medial black stripe as wide as 0.20-0.33 of palp width (Figures 1k-l), pterothorax "marbled" reddish brown with pale yellow spots (Figure 2v), and dorsal terga S4-9 red to orange (Figure 4k). Abdomen narrows gradually from S2 to S4 (Figures 4k, 6k), and S4 is about as wide at level of apical carina as 0.47-0.49 of its length. Hamule inner and outer branches are located in the same plane (Figure 9h), with outer branch bent ventrally over inner branch ( Figure 10q); inner corner of outer branch is pointed and outer corner is rounded (Figure 11r). Basal portion of distal segment of vesica spermalis is parallel sided in ectal view (Figure 13r), and distal portion has paired short medial longitudinal membranous lobes beset with denticles on their margins; paired outer lateral lobes are folded medially to the sides and are shorter than basal sclerotized portion of distal segment (Figure 12q). Cercus (Figure 14n) curves ventrally gradually ( Figure 15r) and its tip is slightly upturned. Epiproct apex is about as wide as a third or less of its maximum width (Figure 16r).
Male of O. philipi shares combination of hamule with outer branch bent ventrally over inner branch and epiproct apex relatively narrow, subapically less than 0.45 of its maximum width, only with O. cultriformis, O. tambopatae, and O. teres. It differs from all by its marbled pterothorax (Figure 2v; versus pterothorax dark reddish brown to black with yellow stripes or entirely dark in mature males, Figure 2k, w-x). It further differs from O. cultriformis by inner and outer branches of hamule separated by a distance shorter than length of inner branch, and vesica spermalis distal segment parallel sided in ectal view (Figures 11r, 13r; versus inner and outer branches of hamule separated by a distance longer than length of inner branch, and vesica spermalis distal segment rhomboidal in ectal view, Figures 11j, 13j), and from O. tambopatae and O. teres by ventral surface of outer branch of hamule smooth (Figure 11r; versus grooved, Figure 11t-u).
Female unknown.

Diagnosis
Male of O. regalis has dark metallic purple frons and vertex, labial palp with medial black stripe as wide as 0.30-0.33 of palp width, pterothorax dark brown to purple with metallic reflections and well-defined yellow stripes, and dorsal terga S1-7 dark reddish brown to black, with yellow stripes along medial longitudinal and ventrolateral carinae, S8 black with yellow stripes along ventrolateral carina, and S9-10 black. Abdomen narrows gradually from S2 to S4, and S4 is about as wide at level of apical carina as 0.48-0.52 of its length. Hamule inner and outer branches are located in the same plane, ending at ca. the same level, with inner corner of outer branch rounded and outer corner triangular and projected ventroposteriorly (Figures 10r, 11s). Basal portion of distal segment of vesica spermalis is medially constricted in ectal view (Figure 13q), distal portion has paired distal short medial longitudinal membranous lobes, and paired outer lateral lobes which are folded medially to the sides and are shorter than basal sclerotized portion of distal segment (Figure 12r). Cercus curves ventrally only slightly ( Figure 15u) and its tip is slightly upturned. Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16s). Female shares color pattern with male, except for frons and vertex dark reddish brown with metallic blue reflections. Abdomen narrows gradually from S2 to S4 (Figure 7o), and S4 is about as wide at level of apical carina as ca. 0.54 of its length. Ventrolateral flap on S8 is black and narrow, as wide as ca. 0.1 of its length (Figure 17p). Vulvar lamina is represented by bilobate posterior margin of S8 (Figure 18s).
Both male and female of O. regalis are unique within the genus by wings with extensive dark markings along C-RA space (versus dark markings restricted to base [i.e. O. philipi] or base and nodus [i.e. O. nodiplaga], if present at all) and male further by basal sclerotized portion of vesica spermalis deeply constricted medially in ectal view (Figure 13q; versus basal portion with sides convex to concave or slightly constricted in ectal view, never deeply constricted, i.e. Figure 13ao, r-s). Within the levis group, O. regalis shares combination of a dark reddish brown to black abdomen with narrow medial longitudinal and ventrolateral yellow stripes on S4-7, gradually narrowing at base, male epiproct narrow at apex, and hamule with inner and outer branches in about the same plane only with O. flavopicta and O. garrisoni. It differs from them and from all other levis group species by its larger size (Hw > 50 mm), presence of four rows of cells between The levis group of Orthemis revisited 203 anal margin of hind wing and AA2 at level of heel of anal loop (as in Figure 3a), outer corner of outer branch of hamule triangular and projected ventroposteriorly in lateral view (Figure 10r), and female S8 ventrolateral flap extremely narrow (as wide as 0.1 of its length, Figure 17p); versus smaller size (Hw < 45 mm), presence of two or three rows of cells between anal margin of hind wing and AA2 at level of heel of anal loop (Figure 3b), outer corner of outer branch of hamule not triangular and projected ventroposteriorly in lateral view (Figure 10a-q, s-t), and female S8 ventrolateral flap relatively wider (as wide as 0.20-0.37 of its length, Figures 17a-o). It can be diagnosed from all species in the ferruginea group by its relatively more slender abdomen, with S4 width/length male ratio of 0.48-0.52 and female ratio of 0.54 (versus male and female ratio of 0.59 or more).

Remarks
As this species is so easily recognizable based on its unique wing color pattern and large size, study of other diagnostic characters has been neglected in the literature; in order to partially compensate for this I provide a brief characterization and diagnosis, including illustrations of vesica spermalis (Figures 12r, 13q), caudal appendages (Figures 15u, 16s), female S8 ventrolateral flap (Figure 17p), and vulvar lamina (Figure 18s).

Orthemis tambopatae
Type specimens examined 2 paratypes, Peru, Madre de Dios Dept., Tambopata-Cándamo Reserved Zone, Camp 3, the Collpa, Río Tambopata west bank, marsh (13 • 8 31 S, 69 • 36 4 W), 1 September 1992, leg. M. Butt [RWG]. Male of O. tambopatae has metallic purple frons and vertex, labial palp with medial black stripe as wide as 0.33-0.60 of palp width, pterothorax reddish brown with yellow stripes (Figure 2w), and dorsal terga S3-8 mostly red. Abdomen narrows gradually from S2 to S4 (Figure 4l), and S4 is about as wide at level of apical carina as 0.38-0.47 of its length. Hamule inner and outer branches form an acute angle between them, with outer branch bent ventrally over inner branch, inner corner of outer branch pointed, and outer surface of outer branch grooved (Figure 11t). Basal portion of distal segment of vesica spermalis is ca. rectangular and only slightly widening distally with sides slightly concave in ectal view (Figure 13t), distal portion has a membranous outer ental lobe ca. as long as basal sclerotized portion in lateral view (Figure 12t), and medioectal sclerotized lobe is ca. half as long as outer membranous lobe in ectal view (Figure 13t). Cercus curves ventrally abruptly ( Figure 15v) and its tip is strongly upturned. Epiproct apex is about as wide as a third or less of its maximum width, and is not bifid (Figure 16t). Male of O. tambopatae shares combination of hamule with outer branch bent ventrally over inner branch and epiproct apex relatively narrow, subapically less than 0.45 of its basal width (Figure 16t Figure 11u), and by distal segment of vesica spermalis with basal sclerotized portion much longer than its maximum width (Figure 13t; versus as long as its maximum width, Figure 13t) and distal outer membranous lobe ca. twice as long as distal medioectal membranous lobe and ca. as long as basal sclerotized portion (Figure 12t; versus distal outer membranous lobe ca. as long as distal medioectal membranous lobe and distinctly shorter than basal sclerotized portion, Figure 12t).

Other specimens examined
Female unknown.

Orthemis teres
Dimensions. Total length 48.9; abdomen length 31.1; Fw length 38.6; Hw length 37.1; Hw maximum width 11.4; cercus length 2.05; epiproct length 1.65; epiproct maximum width 1.1; epiproct subapical width 0.4.  Figure 11t), and by distal segment of vesica spermalis with basal sclerotized portion as long as its maximum width (Figure 13t; versus much longer than its maximum width, Figure 13t) and distal outer membranous lobe ca. as long as distal medioectal membranous lobe and distinctly shorter than basal sclerotized portion (Figure 12t; versus ca. twice as long as distal medioectal membranous lobe and ca. as long as basal sclerotized portion, Figure 12t). Female unknown.

Distribution and biology
Orthemis teres is so far known only from the holotype from Bolivia (Figure 22), and nothing is known about its preferred habitat.