Agriocnemis lepida sp. nov. from Lao PDR 1st June 2020177

Agriocnemis lepida sp. nov. from the Annamite Range in Lao PDR (: )

Malte Seehausen

Museum Wiesbaden, Naturhistorische Sammlungen, Friedrich-Ebert-Allee 2, 65185 Wiesbaden, Germany;

Received 4th February 2020; revised and accepted 5th April 2020

Abstract. Agriocnemis lepida sp. nov. is described and figured (holotype ♂: 20-ii-2003, Lao PDR, Khammouan Province, 2.5 km WNW Ban Tathot, Tham Kamouk, 17.6316°N, 105.1250°E, 200 m a.s.l., P. Jäger leg.; deposited at the Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany). Additionally, illustrations of the male append- ages and the posterior lobe of the prothorax of A. clauseni, A. minima, and A. nana as well as photographs and a Selys watercolour of the female holotype of A. carmelita are provided. Agriocnemis carmelita is shortly discussed with references to the genus Mortonagrion. Further key words. , Zygoptera, new species, cobra-hood, Agriocnemis carmelita, Mortonagrion

Introduction The genus Agriocnemis Selys, 1877, comprises 44 species (Schorr & Paul- son 2020) distributed in Africa, Asia, and Australia. However, some of the species are only known from the type series, and at least the status of A. cor­ beti Kumar & Prasad, 1978, is considered doubtful (Mitra 2010a). Most recently, A. kalinga Nair & Subramanian, 2014, had been described from India as well as A. canuango Dijkstra, 2015, and A. toto Dijkstra, 2015, from Angola. At least seven species of Agriocnemis are known to be native to mainland South-East Asia. These are A. pygmaea (Rambur, 1842), A. femina (Brauer, 1868), A. lacteola Selys, 1877, A. minima (Selys, 1877), A. carmelita Selys, 1877, which is only known from the female holotype collected in »Annam« (today’s Central Vietnam), A. nana Laidlaw, 1914, and A. clauseni Fraser, 1922 (Hämäläinen & Pinratana 1999; Dow 2011b). Yokoi & Souphan- thong (2014) listed A. dabreui Fraser, 1920, A. femina, A. minima, A. nana,

Odonatologica 49(1/2) 2020: 177-190Odonatologica – DOI:10.5281/zenodo.3823341 49(1/2) 2020: 177-190 178 M. Seehausen and A. pygmaea from Lao People’s Democratic Republic. However, the corre- sponding records of the Indian A. dabreui need verification because records from Thailand and Malaysia actually refer to A. minima, and it seems likely that this is the case for Lao PDR and Vietnam as well (Mitra 2010b ). Thus, at least four species of Agriocnemis are known to occur in Lao PDR. During studies of a collection from Lao PDR a male Agriocnemis remained unidentified Seehausen( 2018). Due to further research and comparisons it turned out to be an unknown species, which is described here.

Material and methods Only a single male of the new species could be studied, which is why pre- dominantly male specimens of congenerics were examined for comparison.

Collections with material studied: IRSNB – Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium SMF – Forschungsinstitut und Naturmuseum Senckenberg, Frankfurt am Main, Germany ZSM – Zoological State Collection, Munich, Germany

Other material studied: Agriocnemis carmelita Selys, 1877: 1♀ [Holotype], Annam [Central Vietnam], Coll. Selys (IRSNB). Agriocnemis clauseni Fraser, 1922: 11♂, viii-1952, Baragolai, Ober Assam, India, H. Neuhaus leg. (ZSM). Agriocnemis lacteola Selys, 1877: 1♂ [Syntype], Bengale, Coll. Selys (IRSNB). 2♂ [Syntypes], November 1868, Diuxna Ghats, Atkinson leg., Coll. Selys (IRSNB). 1♂, 30-vi-1914, »Fung Wahn«, China, ex. Coll. Mell, Coll. Ris (SMF). 1♂, »W6696«, China, ex. Coll. Mell, Coll. Ris (SMF). 1♂, 15-vi-1919, »Sumpfplatz«, China, ex. Coll. Mell, Coll. Ris (SMF). 1♂, 04-vi-1937, Shaowu, Fukien, China, 500 m a.s.l., J. Klapperich leg., Coll. G. von Rosen No. 7084 (ZSM). 1♂, 01-viii-1937, Shaowu, Fukien, China, 500 m a.s.l., J. Klapperich leg., Coll. G. von Rosen No. 7085 (ZSM). 1♂, 07-viii-1937, Shaowu, Fukien, China, 500 m a.s.l., J. Klapperich leg., Coll. G. von Rosen No. 7086 (ZSM). 8♂, viii-1952, Baragolai, Ober Assam, India, H. Neuhaus leg. (ZSM). (Selys, 1877): 1♂ [Holotype], Java, Coll. Selys (IRSNB). 1♂, v-1889, Süd Ceylon [likely mislabelled], Fruhstorfer leg., ex. Museum München, Coll. Ris (SMF). 1♂, 07-viii-1972, Ang Kep Nam-Bang Phra, S Chon Buri, Thai- land, G. von Rosen leg. (ZSM). 1♂, 27-v-2003, Prek Toal, Tonle Sap Lake, Battam- bang, Cambodia, J. Constant & K. Smets leg. (IRSNB).

Odonatologica 49(1/2) 2020: 177-190 Agriocnemis lepida sp. nov. from Lao PDR 179

Agriocnemis nana Laidlaw, 1914: 3♂, v-1889, Süd Ceylon [likely mislabelled], Fruhs­torfer leg., ex. Museum München, Coll. Ris (SMF). 4♂ 1♀, Burma [Myan- mar], ex. Coll. E.B. Williamson, Coll. Ris (SMF).

Abbreviations used: Hw – hind wing; Fw – fore wing; Pt – pterostigma; Ax – antenodal cross-veins; Px – postnodal cross-veins; S1, S2 etc. – abdominal segment 1, 2 etc.

Terminology used follows Dijkstra & Clausnitzer (2014).

Agriocnemis lepida sp. nov. (Figs 1–6)

Holotype ♂: Lao People’s Democratic Republic, Khammouan Province, 2.5 km WNW Ban Tathot, Tham Kamouk, 17.6316°N, 105.1250°E, 200 m a.s.l., 20-ii-2003, P. Jäger leg.; deposited at the Forschungsinstitut und Natur- museum Senckenberg, Frankfurt am Main, Germany (SMF).

Etymology Latin lepida means graceful as well as cute and jocose (feminine adjective); all attributes fit in my eyes to members of the genus Agriocnemis in general and to the new species in particular.

Male (holotype; Figs 1-6) Head – Labium pale; labrum and genae blue, upper margin towards ante­ clypeus reddish-brown, blackish in the middle; ante- and postclypeus blue; antefrons blue, broadly black in the midline with black margin towards postclypeus; postfrons black with two black, spot-like elongations on each side of the midline reaching towards antefrons; vertex and occiput black except for pale ocelli, blue comma-shaped post-ocular spots and blue oc- cipital bar; rear of head bluish; pale bluish spot at upper margin of antenna base; first segment of antenna brown with apical margin yellow-ocher; rest of antenna yellow-ocher.

Thorax – Anterior lobe of prothorax light blue; middle lobe black dorsally, the black colour reaching from posterior lobe to lateral half of middle lobe diagonally downward with wavy margin, lower half of middle lobe light blue

Odonatologica 49(1/2) 2020: 177-190 180 M. Seehausen laterally; posterior lobe in lateral view processed, smoothly curved, apical half light blue, basal half with black saddle; in dorsal view the apical margin of posterior lobe process is light blue, rounded and lifted so that it looks like a basin that is open towards the middle lobe; synthorax with mesepisternum black except a light blue ante-humeral stripe; mesepimeron almost entire- ly black; metepisternum and metepimeron light blue with posterior end of metapleural suture black; venter of synthorax pale; coxae, trochanters, and remainder of legs pale whitish-yellow, outer side of femora with black stripe broadened towards tibiae, inner side of femora with washed blackish stripe at the lower third; tibiae with upper half of inner side with washed blackish stripe; tip of tarsi and tarsal claws washed brownish; spines blackish.

Wings – Membrane hyaline; veins brownish; arculus in both wings situated about its own length beyond Ax2; medio-anal link zig-zagged; number of Px in Fw 5–6, number of Px in Hw 4; Pt diamond shaped, brown, each an- terior margin pale whitish.

Abdomen – S1 blue, black dorsally; S2 blue, black dorsally with an oval blue window at each side (“cobra-hood” mark), black narrowed down posteri- orly; S3–S6 dorsally blackish-brown with slight lateral extensions towards the posterior sixth on S3–S5, laterally blue but narrowing from S3 towards S6 and yielding to pale yellow; S7–S9 pale yellow-orange; S10 brownish, laterally lower third pale; cercus pale, laterally with brownish stripe, about as long as S10; in lateral view broadly triangular with apex rounded, ven- trally produced into a downward directed hook-like tip; hollowed in dorsal view; paraproct pale, very short and almost entirely hidden by S10; a tiny brownish spine directed backward is seen from lateral view, a stouter spine directed inside is seen only from dorsal and ventro-lateral view.

Genital ligula – Terminal segment ventrally somewhat rectangular with a pair of broad apical lobes, deeply V-shaped ventrally, tips laterally curved; terminal segment laterally with some broadly rounded extensions.

Measurements [mm] – Total length (with appendages) 21.5; abdomen length (without appendages) 17; cercus 0.25; Hw 10; Fw 10.9; Pt in Fw 0.5.

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Female Unknown.

Diagnosis and remarks Apart from Agriocnemis lepida sp. nov., the group of congenerics with a “co- bra-hood” mark on S2 in India and Southeast Asia comprises A. clauseni, A. corbeti Kumar & Prasad 1978, A. dabreui, A. kalinga, A. keralensis Pe-

Figure 1. Agriocnemis lepida sp. nov., holotype male in lateral view. Focus stacking photo: MS

Figure 2. Agriocnemis lepida sp. nov., holotype male. S2 with “co- bra-hood” mark in lateral view. Fo- cus stacking photo: MS

Odonatologica 49(1/2) 2020: 177-190 182 M. Seehausen ters, 1981, A. minima, and A. nana (Lieftinck 1930; Fraser 1933; Nair & Subra­manian 2014). Within this group, the morphology of the male ap- pendages of A. lepida is especially similar to that of A. nana. However, the cercus of A. nana appears hardly triangular because of a slightly curved up-

a b

c

Figure 3. Agriocnemis lepida sp. nov., holotype male, head. a – Frontal, b – dorsal, c – ventral view. Focus stack- ing photos: MS

a b

Figure 4. Agriocnemis lepida sp. nov., holotype male, posterior lobe of prothorax. a – Dorsal, b – lateral view. Focus stacking photos: MS

Odonatologica 49(1/2) 2020: 177-190 Agriocnemis lepida sp. nov. from Lao PDR 183 per margin, which gives it a humpy expression, and the downward directed hook is less prominent than in A. lepida (cf. Figs 7a, b with 7e, f). Strikingly different is the morphology of the posterior lobe of the prothorax of these two species: in A. nana it has a broad projection (at least as broad as a third of the prothorax) with a slightly notched margin (Figs 7g, h; cf. Lieftinck 1930). Furthermore, A. nana has a completely blue abdomen with black markings on each segment (might be lacking in S8) – and thus lacks the un- marked yellow-orange colour of S7–S9 as it is present in A. lepida.

a b

c

Figure 5. Agriocnemis lepida sp. nov., holotype male, appendages. a – Dor- sal, b – lateral, c – ventro-lateral view. Focus stacking photos: MS

a b

Figure 6. Agriocnemis lepida sp. nov., holotype male, genital ligula. a – Lateral, b – ventral view. Illustrations: MS

Odonatologica 49(1/2) 2020: 177-190 184 M. Seehausen

Compared to A. lepida the male cerci of A. clauseni are slightly longer and more slender, thus less triangular as well, and the paraprocts are with stouter spines, especially the backward directed spine (Figs 7i, j). Furthermore, the morphology of the posterior lobe of the prothorax (Figs 7k, l) is similar to that of A. nana, thus much broader and distinctly different from that of A. lepida, and the abdomen has black markings throughout with S8–S10 completely black (cf. Fraser 1933, Nair & Subramanian 2014).

Figure 7. Male appendages and posterior lobe of the prothorax in lateral and dor- sal view of: a–d – Agriocnemis lepida sp. nov., e–h – A. nana, i–l – A. clauseni, m–p – A. minima. Illustrations: MS

Odonatologica 49(1/2) 2020: 177-190 Agriocnemis lepida sp. nov. from Lao PDR 185

A description of the colour of Agriocnemis lepida sp. nov. is difficult. The specimen was previously conserved in alcohol, and it appears likely that the bluish colour of the thorax and abdomen might have been light greenish when alive. This alteration took place, for example, after alcohol preserva- tion in A. kalinga (Nair & Subramanian 2014). However, coloration and markings of A. lepida are superficially similar to A. minima and A. dabreui, but the male appendages as well as the morphology of the posterior lobe of the prothorax of the latter are distinctly different. Agriocnemis minima and A. dabreui have cerci longer than S10, in lateral view the upper margin is slightly convex, and additionally, the apex of A. minima is slightly curved downwards. Both species lack the downwardly directed hook that makes the cerci of A. lepida look broad and triangular (Figs 7m, n; cf. Fraser 1933). The posterior lobe of the prothorax of A. minima (Figs 7o, p) is dorsally more angular compared to that of A. lepida, and laterally, it is less elevated and straighter backwards instead of curved. I have not seen A. dabreui, but according to Fraser (1933), the posterior lobe of the prothorax has a square projection as in A. lacteola, while in A. lacteola, this square projection is broad and slightly notched, as in A. nana or A. clauseni. The other three Indian species within the “cobra-hood” group, namely A. corbeti, A. kalinga, and A. keralensis, differ distinctly in the morpholo­gy of male appendages, morphology of the posterior lobe of the prothorax as well as in the colour and markings of the abdomen (Kumar & Prasad 1978; Peters 1981; Nair & Subramanian 2014).

It appears unlikely to confuse A. lepida with species outside of the “cobra hood” group. Nevertheless, besides lacking the “cobra-hood”, A. femina and A. pygmaea differ in the morphology of the male appendages and the poste- rior lobe of the prothorax as well as in the black colour of the labrum with its metallic brilliance. Agriocnemis lacteola has a superficially approaching mor- phology regarding the male appendages, but the projection of the posterior lobe of the prothorax is broad and resembles that of A. clauseni and A. nana. Apart from the colour of the abdominal segments, S4–S10 are unmarked whitish and thus strikingly different from A. lepida (cf. Fraser 1933). The male Mortonagrion selenion (Ris, 1916) superficially has a similar coloration to A. lepida but the appendages are distinctly different (cf.Ris 1916).

Odonatologica 49(1/2) 2020: 177-190 186 M. Seehausen

Last but not least, there are three mysterious species that deserve some at- tention: the above-mentioned A. carmelita as well as A. pieli Navas, 1933 (described from a female collected in Chusan, China), and A. luteola Navas, 1936 (described from a male collected in Chusan, China). Agrio­cnemis car­ melita appears to be the most relevant because of the proximity of its type locality »Annam« in Central Vietnam to the Annamite Range of the new species. In addition to Selys (1877), the holotype at IRSNB (Figs 8a–d) and a watercolour painted by Selys (Fig. 9) were also studied. Unfortunately, the holotype lost S2–S10 in the past. However, there are two characters that separate it distinctly from A. lepida: first, the medio-anal link in the wing of A. carmeli­ta is less zig-zagged (Fig. 8b), thus resembling Mortonagrion Fraser, 1920, and second, the clypeus and the frons are dark with some reddish-purplish gloss, and in the watercolour they are even painted black

a b

c d

Figure 8. Agriocnemis carmelita, holotype female. a – Labels, b – wings, c – habi- tus in lateral view, d – head and posterior lobe of prothorax in dorsal view. Photos a, c, d: IRSNB; photo b: MS

Odonatologica 49(1/2) 2020: 177-190 Agriocnemis lepida sp. nov. from Lao PDR 187

(Figs 8d, 9) – this is a character found in A. femina, A. pygmaea, and in some Mortonagrion species. Furthermore, there are 8 Px in the wings, and the synthorax appears very pale without distinct markings (Figs 8b, c). The posterior lobe of the prothorax is inconspicuous but very broad and slightly waved (Fig. 8d). The identity of A. carmelita is not a topic of this paper and certainly needs more research, but some of the characters of A. carmelita actually resemble a teneral female of Mortonagrion falcatum Lieftinck, 1934 (cf. Lieftinck 1934; Asahina 1966).

Figure 9. Agriocnemis carmelita, holotype female. Watercolour by Selys (Ag150a; IRSNB).

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Agriocnemis luteola can easily be separated from A. lepida sp. nov. by the male appendages with paraprocts distinctly longer than the cerci (Navás 1936) – a character likewise found in A. femina only. Agriocnemis pieli was excluded because this species has a black frons and postclypeus (Navás 1933), characters that are found in A. femina and A. pygmaea as well. Also A. pieli as well as A. luteola have 7–8 Px in the fore and 6 in the hind wing (Navás 1933, 1936) whereas they are fewer in A. lepida.

Discussion Of course it is always preferable to describe a new species on the basis of more than a single specimen. However, there are only little data and collec- tions available from Lao PDR, and I was unable to find any further speci- men in collections even from surrounding countries. Therefore, without ad- ditional study, it must remain unclear whether Agriocnemis lepida sp. nov. is an endemic of the Annamite Range or whether it occurs within a wider area. At least it should be searched for in adjacent landscapes in Vietnam and Cambodia.

Figure 10. Habitat of Agriocnemis lepida sp. nov. at Tham Kamouk, Lao PDR. Pho- to: Peter Jäger (26-iv-2012)

Odonatologica 49(1/2) 2020: 177-190 Agriocnemis lepida sp. nov. from Lao PDR 189

The habitat in which A. lepida was found is at Tham Kamouk – a limestone cave in the midst of jungle and temporary agricultural fields within the karst region of the western Annamite Range. A river flows out of the cave forming an almost stagnant to slow-flowing pool of about 100 × 25 m (Fig. 10). The river has strong current and muddy water in the wet season but clear water in the dry season when the specimen was collected (P. Jäger pers. comm.). The relationship between Agriocnemis and the similar Mortonagrion still is uncertain. The new species is placed withinAgriocnemis in the current sense because of the zig-zagged medio-anal link in the wings. However, Dow (2011a) discussed wing venation as a character to separate the genera and found that it is not distinct. He stated that the structure of the genital ligula might be the best character to separate these genera – though the differences appear very slight. So in fact, there is no certain character to separate Agrio­ cnemis and Mortonagrion, and a full revision of both genera is needed.

Acknowledgements I am highly indebted to Dr. Peter Jäger (SMF) for giving the advice to his collection from Lao PDR as well as for information and photographs of the collecting site. Furthermore, I am grateful to Wolfgang Nässig and Massimo Terragni (both SMF), Lars Hendrich, and Katja Neven (both ZSM) and to Wouter Dekoninck and Jérôme Constant (both IRSNB) for the possibility to work on the collections and the loan of specimens in their care. Also, I acknowledge Wouter Dekoninck and Jérôme Constant for providing pho- tographs of the holotype of Agriocnemis carmelita and Wouter Dekoninck, Marcel Wasscher and Karin Verspui for scanning the painting of Selys and the permission to include it in this paper. I also thank Matti Hämäläinen and Martin Schorr for providing literature, Oleg Kosterin for his evaluation concerning the new species and critical reading of the manuscript, and Flo- rian Weihrauch for critical reading of the manuscript.

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