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On Morphology of Tanyderus Pictus (Diptera: Tanyderidae) Pupa and Adult from Chile

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On Morphology of Tanyderus Pictus (Diptera: Tanyderidae) Pupa and Adult from Chile Russian Entomol. J. 25(1): 79–95 © RUSSIAN ENTOMOLOGICAL JOURNAL, 2016 On morphology of Tanyderus pictus (Diptera: Tanyderidae) pupa and adult from Chile Î ìîðôîëîãèè êóêîëêè è èìàãî Tanyderus pictus (Diptera: Tanyderidae) èç ×èëè Elena D. Lukashevich, Dmitry E. Shcherbakov Å.Ä. Ëóêàøåâè÷, Ä.Å. Ùåðáàêîâ Borissiak Paleontological Institute, Russian Academy of Sciences, 117997 Moscow, Russia. [email protected], [email protected] Палеонтологический институт им. А.А. Борисяка РАН, 117997 Москва, Россия. KEY WORDS: living fossils, rearing, xylophagy, chaetotaxy, mouthparts, SEM. КЛЮЧЕВЫЕ СЛОВА: живые ископаемые, выведение, ксилофагия, хетотаксия, ротовые органы, СЭМ. ABSTRACT. The male and female pupae of Tany- ны по вершинному краю. У T. pictus и некоторых derus pictus Philippi, 1865 are described, diagnosed, других танидерид обнаружен короткий вырост на and illustrated for the first time from the specimens жужжальце, соответствующий прехальтеру Ptycho- collected in Chile. The pupa is distinct from the others pteridae. known for this family in the cylindrical thoracic horns with the cup-shaped plastron plate at apex, middle leg sheaths shortest, and pairs of protuberances with one Introduction central seta surrounded by numerous shorter chaetoids on tergites II–VII. The pupation always occurs inside The family Tanyderidae, or primitive crane flies, is a log or branch, in the same chamber where larva an ancient and archaic nematoceran group, known resided and at least sometimes adult emerges inside since the Early Jurassic [Ansorge, 1994; Skibińska et the trunk, just where pupation occurred. Several larvae al., 2014]. Tanyderids have an amphitropical distribu- pupated in laboratory and two females emerged, con- tion with higher diversity in the Southern Hemisphere, firming our earlier identification of the larvae as and were even more widespread in the Mesozoic [Es- T. pictus and their xylophagy. Photographs of live kov, Lukashevich, 2015]. The family includes 38 ex- pupae and female of T. pictus are included. The female tant species in 10 genera. Only three monotypic genera mandibles of T. pictus are well developed, sclerotized, are known in South America, all in the temperate zone and finely toothed along apical margin. The halter of and all described from Chile: Tanyderus Philippi, T. pictus and some other tanyderids is provided with a 1865, Neoderus Alexander, 1927 and Araucoderus short anterobasal projection corresponding to the pre- Alexander, 1929. halter of Ptychopteridae. Knowledge of tanyderid immatures is sparse (for larvae see a review in [Lukashevich, Shcherbakov, РЕЗЮМЕ. Куколки самцов и самок Tanyderus 2014]). The larvae were first discovered and reared to pictus Philippi, 1865 впервые описаны и проиллюст- pupa and adult in Protoplasa fitchii Osten-Sacken, 1860 рированы по собранным в Чили экземплярам. Ку- by Crampton [1930] and Alexander [1930a]. Later the колка отличается от остальных, известных для се- pupae collected together with larvae were described for мейства, цилиндрическими торакальными рогами с Peringueyomyina barnardi Alexander, 1921 and Pro- пористой пластинкой на вершине, сравнительно tanyderus margarita Alexander, 1948 [Wood, 1952; короткими средними ногами и наличием на терги- Knight, 1964]. However, larvae are usually found with- тах II–VII пары выростов со щетинкой, окружённой out pupae [Exner, Craig, 1976; Podeniene, Gelhaus, хетоидами. Окукливание происходит внутри брев- 2013] and only sometimes successfully reared to adults на или ветки, в той же камере, в которой обитала [Pilgrim, 1990]. To date, larvae are not described for личинка, и там же может вылупляться имаго. В the genera Araucoderus, Neoderus and Nothoderus лаборатории несколько личинок окуклились и вы- Alexander, 1927, and pupae for these three plus Tany- велись две самки, что подтвердило нашу предвари- derus, Radinoderus Handlirsch, 1909, Eutanyderus тельную идентификацию личинок как T. pictus и их Alexander, 1928, and Mischoderus Handlirsch, 1909 ксилофагию. Приведены фотографии живых куко- (pupae of Eutanyderus wilsoni Alexander, 1928 and лок и самок T. pictus. У самок T. pictus мандибулы Mischoderus sp. were obtained but remain undescribed хорошо развиты, склеротизованы и мелко зазубре- [Hinton, 1966; Pilgrim, 1990]). Tanyderid larvae, but 80 Elena D. Lukashevich & Dmitry E. Shcherbakov not yet pupae, are found in the fossil state [Lukashevich, boxes with Nothofagus branches (10–20 cm long and 2– Krzemiński, 2009]. 6 cm in diameter, with or without bark); the water The larvae of all three genera, which pupae are covered only the container bottom, without any circula- described, Protoplasa Osten-Sacken, 1860, Protany- tion; most part of branches were wet, but out of water. derus Handlirsh, 1909 and Peringueyomyina Alexander, We maintained containers at ambient room temperature 1921, occur in the hyporheic zone of sand- and cobble- of the laboratory at day and kept cool at night (15º C); bottom rivers and streams; these pupae were found in the containers were checked daily for pupated larvae drier ridges above the water, to where larvae migrate (later for reared adults) and misted with water. prior to pupation. No one pupa of xylobiotic genera was Photographs were taken using a Leica M165C ste- described yet. reomicroscope with a Leica DFC425 digital camera, The type genus of the family Tanyderidae was de- Z-stacked using Helicon Focus 4.10 Pro and adjusted in scribed a century and a half ago [Philippi, 1865], how- Adobe Photoshop® CS3 10.0. For scanning electron ever up to now this monotypic genus is very rare in microscopy (SEM) two pupal exuvia were dehydrated collections and only a dozen of adults is known [Alex- in graded alcohols, soaked in hexamethyldisilazane, ander, 1936; I.R. Madriz, pers. comm., 2014]. In 2014 and dried. Scanning electron micrographs of uncoated we discovered and described tanyderid larvae from logs and gold-coated specimens were taken with a Tescan submerged in streams among Nothofagus forests in the Vega microscope using backscattered electron (BSE) national parks Alerce Andino and Nahuelbuta, Los and secondary electron (SE) detectors. Lagos and Araucania regions, Chile [Lukashevich, The morphological terminology generally follows Shcherbakov, 2014]. The larvae were identified as that of Crampton [1930] and Wood [1952]. ?Tanyderus pictus based on their COI gene sequences being strongly different from those of adult Araucoderus Abbreviations used in the figures gloriosus (Alexander, 1920), the only other tanyderid al anterior lateral or pleural setae and seta-bearing processes species known from the region. However no adults or ant antenna pupae of T. pictus were collected by us in 2014. In apa anterior parietal setae and processes January 2015 in the national parks Alerce Andino and as antesternite Alerce Costero, Los Lagos and Los Rios regions we asp postabdominal spiracle again collected many larvae and also in the former park at antetergite a pupal cast and a mature pupa of T. pictus. Of numer- atg anterior tergal seta ous larvae from the latter park four pupated in laborato- b bubbles ba raised basalar area and setae ry, one very soon and three several months later, and bc basicercus two T. pictus females emerged (Figs 1–4). Our new data cl clypeus confirm the earlier identification of the larvae as T. pictus dc disticercus and their xylophagy. es epistomal setae f frontal setae and processes fl fore leg Material examined fs frontal clefts corresponding to frontal sutures gc gonocoxite Tanyderus pictus Philippi, 1865 gs gonostyle LARVAE: Alerce Andino National Park, sector Sargazo, Lenca h thoracic horn River, 340 masl (41°30' S 72°37’W), 10–18.01.2015, E. Luka- hl hind leg shevich; Alerce Costero National Park, sector Chaihuin, 2–4 km E hp hypopharynx Chaihuin, 20–30 masl (39°57' S 73°32' W), 20–26.01.2015, M. l labrum Chertoprud, E. Lukashevich. lp labial palpus PUPAE: Alerce Andino National Park, sector Sargazo, Lenca lpr lateral processes River, 340 masl, 1 male pupal exuvium #1 and 1 female pupa #2, md mandible 17.01.2015, E. Lukashevich; Alerce Costero National Park, sector Chaihuin, 2 km E Chaihuin, 20 masl, 1 male pupa #3, 25.01.2015 ml middle leg (larva pupated in laboratory); 2 female pupal exuvia #4 and #5, 8.03. mn metanotum 2015, 12.03.2015 (larvae pupated in laboratory); 1 female pupa #6, mp maxillary palpus 1.05.2015 (larva pupated in laboratory), E. Lukashevich. mx maxilla ADULTS: Alerce Costero National Park, sector Chaihuin, 2 km pf prefrontal setae E Chaihuin, 20 masl, 2 females #4 and #5, 15.03.2015 and 19.03.2015 ph prehalteral setae (emerged as adults from larvae pupated 8.03.2015 and 12.03.2015 pl posterior lateral or pleural setae and processes in laboratory). po preocular setae ppaposterior parietal setae and processes Methods ps posterior sternal setae pt posterior tergal setae sc scutal setae All field-sampled and reared specimens were put in sg genal or subgenal setae 70% ethanol, one reared female was pinned. sp rudimentary spiracle The larvae were transferred to the laboratory along tg tegular region and setae with their native substrate. They were kept in plastic vp verriculate protuberance On morphology of Tanyderus pictus pupa and adult from Chile 81 Localities, habitats and behaviour Some of these branches were with bark and some with- out, some were solid and some saturated, decomposed, The larvae were collected in the national parks easily disintegrating into whitish fibres. The most pro- Alerce Andino and Alerce Costero (Figs 5–6), in Notho- ductive log
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