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430 Accepted by M. Marinov: 9 Apr. 2018; published: 17 May 2018 ZOOTAXA ISSN 1175-5326 (print edition) ISSN 1175-5334 (online edition) Copyright © 2018 Magnolia Press Zootaxa 4420 (3): 430438 http://www.mapress.com/j/zt/ Article https://doi.org/10.11646/zootaxa.4420.3.8 http://zoobank.org/urn:lsid:zoobank.org:pub:BC7F05F3-BAEE-4AAF-B028-D039EDE088EF Females of two species of Argia from Chapada dos Guimarães National Park, Brazil (Odonata: Coenagrionidae) DIOGO SILVA VILELA 1,2,5 , RHAINER GUILLERMO-FERREIRA 2 , KLEBER DEL-CLARO 3 & ADOLFO CORDERO-RIVERA 4 1 Graduate Program in Entomology, Department of Biology, University of São Paulo (USP), Ribeirão Preto, Brazil. 2 Laboratory of Ecological Studies on Ethology and Evolution (LESTES), Department of Hydrobiology, Federal University of São Car- los, Brazil. 3 Laboratório de Ecologia Comportamental e Interações (LECI), Biology Institute, Federal University of Uberlândia, Uberlândia, Brazil. 4 Grupo de Ecoloxía Evolutiva e da Conservación, Departamento de Ecoloxía e Bioloxía Animal, Universidade de Vigo, Pontevedra, Spain. 5 Corresponding author. E-mail: [email protected] Abstract The female of Argia tupi Calvert, 1909 (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Cachoeira do Marimbondo (15.4330° S, 55.7198° W, 370 m), 01 xi 2015) is described, illustrated and diagnosed based on comparison with sympatric species of Argia Rambur, 1842. We also augmented the description of Argia bicellulata (Calvert, 1909) female (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Rio Paciencia (15.3438° S, 55.8322° W, 280 m), 25 x 2015). Key words: Damselfly, Cerrado, Neotropical, Savannah, Palm Swamp Introduction Argia Rambur, 1842 is one of the most speciose genus within Coenagrionidae with over 130 described species (Garrison & von Ellenrieder 2017), almost 50 occurring in Brazil (Lencioni 2006). Descriptions are complete for most species that includes full diagnoses for both sexes. Among Brazilian species only six remain with undescribed females: A. clausseni Selys, 1865, A. cyathigera Navás, 1934, A. subapicalis Calvert, 1909, A. tamoyo Calvert, 1909 and A. tupi Calvert, 1909 (Lencioni 2006). Fortunately, all females mentioned above are being described by Rosser Garrison and Natalia von Ellenrieder (pers. comm.). Here we describe the female of A. tupi and also augment Calvert's description of the female of A. bicellulata, based on the second series of females collected since the original single female was described by Calvert (1909). Materials and methods The specimens were collected during an expedition to the Chapada dos Guimarães National Park, Mato Grosso State, Brazil, in late 2015. We follow Garrison et al. (2010) for body morphology nomenclature. All measurements are in mm. Laboratory photographs were taken with a Canon Eos M and Eos M5 with a Canon 28mm macro lens by ACR. Field photographs were taken with a Canon Eos 7D Mark II fitted with a Canon 100mm macro lens by ACR. Drawings were executed by Rosser W. Garrison. Abbreviations for structures used throughout the text are as follows: S1–10: abdominal segments 1 to 10, Fw: forewing, Hw: hindwing. For wing venation we followed Riek & Kukalová-Peck (1984).

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Page 1: 4420 (3): 430 438 Article ZOOTAXA Vilela et al Argia females.pdf · The female of Argia tupi Calvert, 1909 (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Cachoeira do

ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2018 Magnolia Press

Zootaxa 4420 (3): 430–438

http://www.mapress.com/j/zt/Article

https://doi.org/10.11646/zootaxa.4420.3.8

http://zoobank.org/urn:lsid:zoobank.org:pub:BC7F05F3-BAEE-4AAF-B028-D039EDE088EF

Females of two species of Argia from Chapada dos Guimarães National Park,

Brazil (Odonata: Coenagrionidae)

DIOGO SILVA VILELA1,2,5, RHAINER GUILLERMO-FERREIRA2,

KLEBER DEL-CLARO3 & ADOLFO CORDERO-RIVERA4

1Graduate Program in Entomology, Department of Biology, University of São Paulo (USP), Ribeirão Preto, Brazil.2Laboratory of Ecological Studies on Ethology and Evolution (LESTES), Department of Hydrobiology, Federal University of São Car-

los, Brazil.3Laboratório de Ecologia Comportamental e Interações (LECI), Biology Institute, Federal University of Uberlândia, Uberlândia,

Brazil.4Grupo de Ecoloxía Evolutiva e da Conservación, Departamento de Ecoloxía e Bioloxía Animal, Universidade de Vigo, Pontevedra,

Spain.5Corresponding author. E-mail: [email protected]

Abstract

The female of Argia tupi Calvert, 1909 (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Cachoeira do

Marimbondo (15.4330° S, 55.7198° W, 370 m), 01 xi 2015) is described, illustrated and diagnosed based on comparison

with sympatric species of Argia Rambur, 1842. We also augmented the description of Argia bicellulata (Calvert, 1909)

female (BRAZIL, Mato Grosso, Chapada dos Guimarães National Park, Rio Paciencia (15.3438° S, 55.8322° W, 280 m),

25 x 2015).

Key words: Damselfly, Cerrado, Neotropical, Savannah, Palm Swamp

Introduction

Argia Rambur, 1842 is one of the most speciose genus within Coenagrionidae with over 130 described species

(Garrison & von Ellenrieder 2017), almost 50 occurring in Brazil (Lencioni 2006). Descriptions are complete for

most species that includes full diagnoses for both sexes. Among Brazilian species only six remain with undescribed

females: A. clausseni Selys, 1865, A. cyathigera Navás, 1934, A. subapicalis Calvert, 1909, A. tamoyo Calvert,

1909 and A. tupi Calvert, 1909 (Lencioni 2006). Fortunately, all females mentioned above are being described by

Rosser Garrison and Natalia von Ellenrieder (pers. comm.). Here we describe the female of A. tupi and also

augment Calvert's description of the female of A. bicellulata, based on the second series of females collected since

the original single female was described by Calvert (1909).

Materials and methods

The specimens were collected during an expedition to the Chapada dos Guimarães National Park, Mato Grosso

State, Brazil, in late 2015.

We follow Garrison et al. (2010) for body morphology nomenclature. All measurements are in mm.

Laboratory photographs were taken with a Canon Eos M and Eos M5 with a Canon 28mm macro lens by ACR.

Field photographs were taken with a Canon Eos 7D Mark II fitted with a Canon 100mm macro lens by ACR.

Drawings were executed by Rosser W. Garrison. Abbreviations for structures used throughout the text are as

follows: S1–10: abdominal segments 1 to 10, Fw: forewing, Hw: hindwing. For wing venation we followed Riek &

Kukalová-Peck (1984).

430 Accepted by M. Marinov: 9 Apr. 2018; published: 17 May 2018

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Acronyms used for collections are as follows:

LESTES—Laboratory of Ecological Studies on Ethology and Evolution, UFSCar, Brazil

Results

Remarks on the female of Argia bicellulata Calvert, 1909

Figs. 1–11.

Material examined. 3 ♀ (LESTES, Cod. ACR 03297, ACR 3322, ACR 3323), BRAZIL, Mato Grosso, Chapada

dos Guimarães National Park, Rio Paciencia (15.3438° S, 55.8322° W, 280 m), 25 x 2015, R. Guillermo-Ferreira

leg. [RGF].

Measurements. Fw 14.3, Hw 13.8, abdomen 18.8, total length 23.4.

Variations on the specimens examined (n = 3). The mesostigmal plates illustrated here (Figs. 2‒3) are from

Calvert's single female. No variation on mesostigmal plate morphology was detected, comparing our specimens to

figures of Calvert's female (Figs. 2‒3); Wing venation: no variation RP2 branching and postquadrangular cells;

postnodal varied as follows: 10–11in Fw and 9–11 in Hw (Table 1); size varied as follows: abdomen 18.3–20.2

(19.1±0.9), total length 23.3–25.9 (24.2±1.4), Fw 14.3–14.9 (14.5±0.3), Hw 13.8–14.6 (14±0.4). Body coloration

had little variation on the specimens examined (Figs. 4–11), being more visible on the slightly different post ocular

spots coloration (Figs. 10–11).

TABLE 1. Venation statistics for Argia bicellulata examined by Calvert (1909) and in this study.

Differential diagnosis. Argia bicellulata is the smallest known species of this genus (Garrison & von

Ellenrieder 2015). Females are similar to males (Figs. 1a‒b) in having the black ventral thoracic coloration, a

unique trait distinguishing A. bicellulata from its sympatric congeners (Figs. 8‒9). The narrow posterodistally

pointed digit-like mesostigmal lobe (Figs. 2‒3) differs from other sympatric species including the broadly arcuate

lobe in A. botacudo Calvert, 1909 (Figs. 12‒13) and broad thumb-like lobe in A. tupi (Figs. 15‒16). As stated by

Calvert (1909), A. bicellulata females have a coloration that resembles the male and by examining the females with

living colors (Figs. 4‒11), we noticed some differences from its original description. The dorsal spot on the S2,

blue in Calvert’s female, is pale on the examined females of this study. On the contrary, the S9 dorsal spot was pale

in Calvert’s female, and our females have it in blue. The postocular spots are smaller than in males, and its

coloration is violet, instead of the blue that is found in males. A remarkable trait is the number of Fw

postquadrangular cells (Table 1), numbered three in all our specimens (a frequent occurrence to Argia species),

which enforces the synonymy of Diargia Calvert, 1909 with Argia, where Diargia does not present strong enough

characteristics to be considered a different genus (Gloyd 1968). All other morphological traits on our specimens

agree with Calvert’s description, such as the lack of vulvar spines on S9, hyaline wings with a reddish venation at

the base, darkening distally.

Habitats and ecology. Both sexes were collected in palm swamp (i.e. veredas, Vilela et al. 2016) areas, in the

same locality from where a new Argia species was recently discovered (Vilela et al. 2018). At Paciência River the

palm swamp had dense vegetation and was connected to the stream although we found A. bicellulata only within

palm swamp including small slow streams within the palm swamps. This species can be confused with Ischnura

Calvert, 1909

(9♂1♀)

This study

(3♀)

Fw Hw Fw Hw

Postquadrangular cells 2 (90%) or 3 (10%) 2 (95%) or 3 (5%) 3 (100%) 2 (100%)

RP2 branching 5 (80%), 6 (10%) or

midway (10%)4 (40%), 5 (45%), midway (10%) or in just one wing (5%)

6 (100%) 5 (100%)

Postnodal cells 9–12 7–11 10–11 9–11

Zootaxa 4420 (3) © 2018 Magnolia Press · 431DESCRIPTION OF TWO FEMALE ARGIA FROM COLOMBIA

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Charpentier, 1840 or Argentagrion Fraser, 1948 species due to its small size and flight behavior, flying through the

dense grass vegetation of the swamp.

FIGURE 1 a‒b. Argia bicellulata (Calvert) BRAZIL: Chapada dos Guimarães; 1a. ♀ (ACR 3322); 1b. ♂ (ACR 3326)

VILELA ET AL.432 · Zootaxa 4420 (3) © 2018 Magnolia Press

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FIGURES 2‒3. Argia bicellulata (Calvert) BRAZIL: "Chapada" (paralectotype) mesostigmal lobes; 2. anterior view; 3.

dorsal view.

FIGURES 4‒11. Argia bicellulata (Calvert) BRAZIL: Chapada dos Guimarães; 4. S1‒10, ♀ (ACR 3322); 5. S1‒10, ♀ (ACR 3326); 6. S8‒10, ♀ (ACR 3326); 7. S8‒10, ♀ (ACR 3322); 8. thorax, ♀ (ACR 3322); 9. thorax, ♀ (ACR 3326); 10. head, ♀ (ACR 3326); 11. head, ♀ (ACR 3322).

FIGURES 12‒13. Argia botacudo (Calvert) BRAZIL: “Chapada” mesostigmal lobes; 12. ♀ dorsal view; 13. ♀ anterior view.

Zootaxa 4420 (3) © 2018 Magnolia Press · 433DESCRIPTION OF TWO FEMALE ARGIA FROM COLOMBIA

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Description of the female of Argia tupi Calvert, 1909

Figs. 14–24, 29b.

Material examined. 2 ♀ (LESTES, Cod. ACR 8184 (described below), ACR 8197), BRAZIL, Mato Grosso,

Chapada dos Guimarães National Park, Cachoeira do Marimbondo (15.4330° S, 55.7198° W, 370 m), 01 xi 2015,

A. Cordero-Rivera leg. [ACR] and R. Guillermo-Ferreira leg. [RGF].

Female (Fig. 14). Head. Labium, labrum, clypeus, base of mandibles, genae, frons and front of head to level of

antennae tan, epicranium black, except for small dark cupreous spots lateral to each lateral ocelli and at base of

antennae, postocular spots tan, tan occipital bar separated from postocular spots by a thin black line, border of

postocular lobes black; eyes (in life) black dorsally and white ventrally; rear of head largely black, a pale line

bordering eye margin.

FIGURE 14. Argia tupi (Calvert) BRAZIL: Chapada dos Guimarães; ♀ (ACR 8184).

Thorax. Anterior lobe of prothorax tan; middle lobe black with two lateral pale circular spots; propleuron tan,

posterior lobe tan, except for rectangular black spot at base, almost separated medially; mesostigmal plate black,

mesostigmal lobes strongly erect, almost perpendicular to the mesepisternum, lying somewhat diagonally with

medio-apical portion of lobe bent posteriorly (Figs. 15‒16); pterothorax pale with usual pair of black thoracic

stripes as follows: mid-dorsal stripe slightly narrower than tan antehumeral stripe, this followed by a broad deeply

forked humeral stripe, its upper 0.60 a narrow line along mesopleural suture slightly enlarged at mesopleural pit,

posterior arm thicker, extending diagonally to base of wings and connected along crest with mid-dorsal strip,

mesinfraepisternum black, pale tan on 0.25 lower portion, remainder of thorax ivory with narrow metapleural stripe

ending ventrally at metastigma; venter of the thorax ivory.

Wings. Hyaline, venation black, pterostigma brown; postnodals 16/15 in Fw, 14/13 in Hw, postquadrangular

cells 4/4 in Fw and 4/4 in Hw, RP2 branching at postnodal 8 in Fw and at postnodal 7 in Hw.

Abdomen. Tan above, ivory below with following areas black: S1 with a dorsal black spot covering basal half;

VILELA ET AL.434 · Zootaxa 4420 (3) © 2018 Magnolia Press

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S2 with a dorso-lateral stripe narrowed above medially followed by an expanded portion dorsally but not

connecting above, ventrally with an ill-defined black stripe bordering margin; S3‒7 with a similar pattern but mid-

dorsal stripe narrower medially and connecting above at apical 0.10 thus isolating a broad tan spindle-shaped

dorsal stripe, dorso-lateral stripe apically almost (S3) or (S4‒6) connecting with ventral stripe; S7 with black more

expansive thus losing pale spindle-shaped stripe laterally; S8‒9 black apically with a pale blue trident shaped

dorsal spot, thicker on S9, S10 mostly pale blue, remainder black; appendages black, ovipositor and styli black,

except for white basis of the styli.

Measurements. Fw 28, Hw 27.3, abdomen 32.4, total length 42.

FIGURES 15‒16. Argia tupi (Calvert) BRAZIL: Serra do Cipó mesostigmal lobes; 15. ♀ dorsal view; 16. ♀ anterior view.

FIGURES 17‒24. Argia tupi (Calvert) BRAZIL: Chapada dos Guimarães; 17. S1‒10, ♀ (ACR 8184); 18. S1‒10, ♀ (ACR 8197); 19. head, ♀ (ACR 8184); 20. head, ♀ (ACR 8197); 21. S8‒10, ♀ (ACR 8184); 22. S8‒10, ♀ (ACR 8197); 23. thorax, ♀ (ACR 8197); 24. thorax, ♀ (ACR 8184).

Zootaxa 4420 (3) © 2018 Magnolia Press · 435DESCRIPTION OF TWO FEMALE ARGIA FROM COLOMBIA

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FIGURES 25‒26. Argia claussenii (Selys) BRAZIL: Serra do Cipó mesostigmal lobes; 25. ♀ dorsal view; 26. ♀ anterior view.

FIGURES 27‒28. Argia cyathigera (Navás) BRAZIL: Nova Teutonia mesostigmal lobes; 27. ♀ dorsal view; 28. ♀ anterior view.

FIGURE 29 a‒b. Habitats of Argia tupi. 29a. Cachoeira do Marimbondo, natural habitat of Argia tupi; 29b. couple of Argia

tupi at its natural habitat.

VILELA ET AL.436 · Zootaxa 4420 (3) © 2018 Magnolia Press

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Variations on the specimens examined (n = 2). The other female showed no variations on mesostigmal plates

morphology; wing venation showed no variation in postquadrangular cells, RP2 branching or postnodal; size varied

as follows: abdomen 31.8‒32.4 (32.1±0.4), total length 41.1‒42 (41.5±0.6), Fw 27.3–28 (27.6±0.4), Hw 26.6–27.3

(26.9±0.4). Body coloration variation for both females is shown in Figs. 17‒24. The tan and blue coloration were

darker in the other female.

Differential diagnosis. In contrast to A. bicellulata, A. tupi is one of the largest Brazilian species occuring

within Chapada. Apart from the large body size, A. tupi females can be separated from other Brazilian species,

including A. clausseni and A. cyathigera (females yet to be described by Rosser Garrison and Natalia von

Ellenrieder) by the morphology of its mesostigmal lobes. In A. tupi the short narrow mesostigmal lobe is strongly

erect (Fig. 16), with a less rounded shape when compared to A. claussenii (Figs. 25‒26) and A. cyathigera (Fig.

27‒28). The medio-apical portion of the lobe of A. tupi lies somewhat diagonally in relation to the mesepisternum

and is bent posteriorly, whereas in A. claussenii (Fig. 25) and A. ciathygera (Fig. 27) these lobes are not bent

posteriorly and have a larger foliate shape.

Habitats and Ecology. This species was collected at a waterfall area in Chapada dos Guimarães National Park

(Fig. 29a). It inhabits the exposed streams above the waterfall and was found perched on the stones and vegetation

around stream margin (Fig. 29b). Oviposition was observed in tandem, on the rocky walls of the waterfall, in the

roots and other vegetal debris found on the rapid areas of the stream.

Discussion

Described by Calvert (1909) as member of the new genus Diargia, D. bicellulata (later synonymized with Argia by

Gloyd, 1968) is the smallest member of the genus. According to Rosser Garrison (pers. comm.) the other known A.

bicellulata specimen was a male received by Frederico Lencioni, from Mato Grosso State, in 2011. Only one

female was previously known and described, however with only a small outline drawing of dorsal mesostigmal

plate (Calvert 1909); Figs. 2‒3 are renderings of the mesostigmal plates of the female described by Calvert (1909).

Furthermore, this is an unusual somewhat atypical species not only due to its small size, but also due to apparent

unique genital ligula morphology (to be treated in a further study by Rosser Garrison and Natalia von Ellenrieder,

pers. comm.) and black venter of the thorax in both sexes. Argia tupi is a poorly known species, only treated by

Calvert (1909) in his original description. This species was only cited in one other study (Longfield 1929).

Females of A. bicellulata and A. tupi were discovered within a National Park, thus insuring these two species.

Argia bicellulata seems to be restricted to lentic habitats within palm swamp habitats. This may account for the

apparent rarity of this species as adults may likely be overlooked or mistaken for a species of Ischnura or

Argentagrion. On the other hand, A. tupi inhabits lotic water bodies, near waterfalls, easily recognizable by its

larger size.

Acknowledgments

DSV thanks Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for a scholarship grant

(Proc.140732/2016-0). RGF thanks CNPq and Fundação de Amparo à Pesquisa do Estado de São Paulo (FAPESP)

for constant support. KDC thanks CNPq for regular financial support (Proc. 301605/2013-0). ACR funding was

provided by the Spanish Ministry of Economy and Competitiveness (Grant number CGL2014-53140-P, including

FEDER funds), and Ministry of Education, Culture and Sports (Grant number PHBP14/00069). We thank CAPES-

DGPU (Proc. 31815) for funding our expedition to Chapada dos Guimarães. We are very grateful to Rosser W.

Garrison for his continuous support, review and for all the drawings of this manuscript. We also thank Thais

Regina de Almeida for field assistance, Dennis Paulson and Milen Marinov for valuable comments on the

manuscript.

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References

Calvert, P.P. (1909) Contributions to a knowledge of the Odonata of the Neotropical region, exclusive of Mexico and Central America. Annals of the Carnegie Museum, 6 (3), 73–280.

Garrison, R.W., von Ellenrieder, N. & Louton, J.A. (2010) Damselfly Genera of the New World: An Illustrated and Annotated

Key to the Zygoptera. The Johns Hopkins University Press, Baltimore, MD, 490 pp.Garrison, R.W. & von Ellenrieder, N. (2015) Damselflies of the genus Argia of the Guiana Shield (Odonata: Coenagrionidae).

Zootaxa, 4042 (1), 1‒134. https://doi.org/10.11646/zootaxa.4042.1.1

Garrison, R.W. & von Ellenrieder, N. (2017) New species of the damselfly genus Argia from Mexico, Central America and Ecuador with an emphasis on Costa Rica (Insecta: Odonata: Coenagrionidae). Zootaxa, 4235 (1), 1‒93. https://doi.org/10.11646/zootaxa.4235.1.1

Gloyd, L.K. (1968) The synonymy of Diargia and Hyponeura with the genus Argia (Odonata: Agrionidae: Argiinae). The

Great Lakes Entomologist, 1 (8), 271‒274.Lencioni, F.A.A. (2006) Damselflies of Brazil: an illustrated identification guide. All Print Editora, São Paulo, 419 pp.Longfield, C. (1929) A list of Odonata of the state of Matto Grosso, Brazil. Transactions of the Entomological society of

London, 77 (1), 125‒139. https://doi.org/10.1111/j.1365-2311.1929.tb00682.x

Riek, E.F. & Kukalová-Peck, J. (1984) A new interpretation of dragonfly wing venation based upon Early Upper Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings. Canadian Journal of Zoology, 62 (6), 1150–1166. https://doi.org/10.1139/z84-166

Vilela, D.S., Guillermo-Ferreira, R. & Del-Claro, K. (2016) The Odonata community of a Brazilian vereda: seasonal patterns, species diversity and rarity in a palm swamp environment. Bioscience Journal, 32, 486–495. https://doi.org/10.14393/BJ-v32n2a2016-30491

VILELA ET AL.438 · Zootaxa 4420 (3) © 2018 Magnolia Press