Xyelidae

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Xyelidae
Xyela julii

Xyela julii

Systematics
Class : Insects (Insecta)
Order : Hymenoptera (Hymenoptera)
Subordination : Plant wasps ("Symphyta")
Superfamily : Xyeloidea
Family : Xyelidae
Scientific name
Xyelidae
Newman , 1834

The Xyelidae are a species-poor family of plant wasps with around 80 recent species worldwide, 15 of them in Europe. The family has been proven since the Triassic and is therefore one of the oldest living insect families with an age of over 200 million years.

features

Xyelidae are small hymenoptera, the European species are about 3–5 mm long, the Macroxyela and Megaxyela living in East Asia and North America are significantly larger at 10–15 mm in length. As with most insects, the head has two compound eyes , three ocelli and biting-chewing mouthparts. The four wings are membranous. There is usually a large pterostigma on the front edge of the fore wing . The legs are designed as normal walking legs. At the front end of the rail ( tibia ) of the foreleg sits a redesigned spur that serves as a cleaning tool for the antennas. As is typical for all plant wasps, the abdomen sits broadly on the thorax without a constriction (wasp waist), on the upper side of which there are two so-called "cenchri" (adhesive pads) for holding the wings in rest position.

The body of the animals is sometimes monochrome black, but often colored with contrasting white, yellow or orange markings. The structure of the antennae of the Xyelidae is very striking and unmistakable . The third limb (i.e., the first limb of the antenna whip) is longer and wider compared to the following limbs, sometimes longer than all the other limbs combined. The third link is followed by at least five, in European species there are always nine or more narrow and short flagellum limbs. The elongated third antennae link is the result of the fusion of several original whiplash links. Such a "Synantennomer" occurs within the recent Hymenoptera except in the Xyelidae only in the Blasticotomidae . In Pleroneura , Xyelecia and most Xyela species, the maxillary palps are greatly enlarged and have specialized bristles on the distal limbs that are used to acquire food. The prothorax is narrow and straight at the rear edge. The wings which falls Flügelgeäder on having an unusually high for Hymenoptera number of cross wires and cells. The wings are hairless ( Xyela ) or hairy (e.g. Pleroneura ). The coloring of the wing membrane and veins varies. In the Xyelidae, the radial sector Rs , a vein in the front wing half, branches into the veins Rs1 and Rs2 , while other Hymenoptera Rs1 is absent. On the middle and rear legs there are three spurs in the middle and in front of the end. The females have a more or less long ovipositor at the end of the abdomen , which is straight or slightly curved and stretched backwards. In some species it can reach body length. Its shape and length are crucial for determining many species.

Larvae

As with many plant wasps, the larvae are very similar in body to butterfly caterpillars ("erucoid" larvae type). The species living in plants are white, the wild species are whitish-green or yellow in color. Megaxyela larvae are conspicuously dotted with black (Plate 21, Fig. 3 in) or resemble bird excrement. The rounded head with a heavily sclerotized head capsule has a larval eye (stemma or ocularium) on both sides, which is only preserved as a relic in miners, and very short, five-part antennae. He has strong, toothed mandibles . Labial and maxillary palps are tripartite. At the lip of the Labialdrüsen with which the larvae can spin silk-like threads that are used to build the cocoons lead. The reservoir of the glands is very large and extends almost through the whole body to the abdomen. The mouthparts are directed downwards (orthognath). There are three short, tripartite pairs of legs on the thorax . At the abdomen sit at all segments simple legs. The larvae of all other Hymenoptera and most of the butterfly caterpillars lack the pair of legs on the first segment. In free- living Xyelidae ( Macroxyela , Megaxyela ) the abdominal legs are clear and bipartite. In the herbivorous species ( Pleroneura , Xyela ) they are only formed as inconspicuous transverse ridges.

Dolls

Xyelidae have a so-called pupa dectica , in which the antennae, legs and mandibles are free and mobile. In fact, it is the already developed Imago, which is still enclosed by the doll's skin (“pharate”). The wings are not yet expanded at this stage and the ovipositor of the females is still curved over the abdomen. The pupa dectica opens the cocoon, digs itself to the surface and is able to walk around there, plate 21 fig. 4 in and to absorb fluid.

The pupa dectica is part of the basic plan of the holometabola and also occurs in numerous other subgroups of these, such as the reticulated flies (Neuropterida), beaked flies (Mecoptera), caddis flies (Trichoptera) and in basal butterfly families (Lepidoptera). The Hymenoptera to the exclusion of the Xyelidae have an immobile pupa adectica .

Way of life

All Xyelidae are herbivores ( phytophagous ) on trees. Larvae of the relatively species-rich genus Xyela , which is also common in Europe, live in the growing male inflorescences of pine species and feed on the immature pollen grains. The North American Xyela gallicaulis calls fresh shoots of pine bile produced in which they eat. Larvae of the Pleroneura species, which are also widespread in Europe, feed on the young shoots of fir trees ( Abies ). Only the Japanese Pleroneura piceae occurs on spruces ( Picea ). The larvae of the Macroxyelinae feed freely on the leaves of deciduous trees. The two North American Macroxyela TYPES feed on elms that the spread in the Far East and North America Megaxyela species on various walnut crops such as walnuts ( Juglans ), hickory ( Carya ) and wing nuts ( Pterocarya ). It is assumed that the larvae of Xyelecia nearctica have an internal way of life and are bound to fir trees.

For many Xyelidae species only a single larval food plant is known. Monophagy is presumably only present in Xyela species, while in the other genera a lack of data is probably often only a pretense of monophagy. In the larvae of the pollen-eating Xyela species, oviposition must be very precisely coordinated with the relatively short development time of male pine blossoms. This presumably prevents the transition to other host species. Some North American Xyela species have been reported to be bound to different pine species ( oligophagy ), but there may be taxonomic problems in separating the very similar Xyela species.

When the feeding phase is over, the larva drops to the ground and digs a small hole in the ground into which it weaves itself into a cocoon. Here she sheds her skin into a doll. The imago does not hatch until the spring of the following year to mate. Then the females lay the eggs with their ovipositor in the plant tissue of their host species. In many species, the pupae can linger for several years ( diapause ) and only hatch in the second or third year. In the case of the Xyela alpigena and X. obscura , which live on Swiss stone pine and mountain pine in the Alps , a diapause of at least two years is mandatory, as these pine species in the subalpine zone only bloom irregularly.

The adults are, as far as known, unspecialized pollen eaters on a variety of plant species. The enlarged maxillary palps of Xyela (and probably also of Pleroneura and Xyelecia ) are used to extract pollen from flowers. Mating takes place on the ground, with the animals facing away from each other, coupled to each other with their copulatory organs. In the subfamily Xyelinae, the mating organs of the male are therefore rotated by 180 ° in the abdomen (" strophandry "). In the Macroxyelinae, mating takes place in the same position, but here the male copulation organs only turn during the copulation process themselves (“facultative strophandry”).

Although the species occur on economically important tree species, they are mostly insignificant as pests. The larvae of Pleroneura piceae damage the growth of the Sakhalin spruce ( Picea glehnii ) by destroying the young shoots. Larvae of Megaxyela major (and presumably M. langstoni ) feed on the leaves of pecan nuts ( Carya illinoinensis ) and are considered pests in pecan cultures.

Systematics and taxonomy

The Xyelidae are the morphologically most primal group of the hymenoptera and almost certainly the sister group of all other living hymenoptera. This position results from numerous morphological features and also from molecular pedigrees based on the comparison of homologous DNA sequences. The old age of the family is supported by fossils. All hymenoptera fossils found in the Triassic are assigned to this family, representatives of other families can only be detected in the Jurassic at the earliest. In the Mesozoic and Tertiary the family was much more species-rich and more widespread than today, the few recent, living representatives can be viewed as a relic group.

Within the Xyelidae, a distinction is made between the Xyelinae and the Macroxyelinae. In the meantime it was assumed that the Xyelidae could be paraphyletic and that either the Xyelinae or the Macroxyelinae alone form the sister group of the rest of the Hymenoptera. But today this is considered unlikely. The living (recent) species can thus be classified in the following system:

The European species can be identified with “The Western Palaearctic Xyelidae” by Blank (2002), the entire Eurasian species with Blank et al. (2013). The North American Macroxyelinae were revised by Smith & Schiff (1998), the North American Xyela species by Burdick (1961), the North American Pleroneura species by Smith et al. (1977), the East Asian megaxyela species from Shinohara (1992), the East Asian pleroneura species from Shinohara (1995), and the megaxyela species of the world by Blank et al. (2017).

distribution

The recent Xyelidae occur exclusively in the northern hemisphere, whereby the moderate (temperate) latitudes are clearly preferred. Representatives of Megaxyela and Xyela are widespread as far as the northern (boreal) zone and the subtropics. No species are known from the tropics. Macroxyela occurs exclusively in North America, while the other genera have a Holarctic distribution . In the case of Megaxyela and Xyelecia , this is limited to East Asia and North America, while Xyela and Pleroneura are more common in Eurasia.

In Germany, Xyela curva and Xyela julii are widespread and common. Pleroneura coniferarum , Pleroneura dahlii and Xyela longula are very rare. Significantly more species are found in the Alps and in the Mediterranean region, as there are more autochthonous pine species, in which the individual Xyela species are specialized: Xyela alpigena on Pinus cembra , Xyela curva , Xyela graeca and Xyela menelaus on Pinus nigra , Xyela obscura on Pinus mugo . Xyela curva originally reached its northern limit in the Vienna Basin. With the black pines planted in gardens, parks and in the forest , it broadened its distribution secondary to Central Europe and the British Isles.

Fossils

The earliest finds of Xyelidae date back to the Middle Triassic. In the famous fossil deposit of Madygen in Kyrgyzstan, between 220 and 230 million years old, 25 species have already been found. In the Jurassic and Chalk, the family is species-rich and widespread with more than 36 genera in four subfamilies worldwide. In the Tertiary there are already comparatively fewer finds. The richest deposit here is the Rott fossil deposit near Bonn from the Oligocene with 11 species. The lack of family in Baltic amber, which is believed to have been caused by a species of pine, is a mystery. In some fossils from the Mesozoic and Tertiary periods, the fossilized intestinal contents from pollen were detected. This means that pollen nutrition can already be developed from this time.

supporting documents

Individual evidence

  1. ^ Blank, SM, Groll, EK, Liston, AD, Prous, M. & Taeger, A. 2012: ECatSym - Electronic World Catalog of Symphyta (Insecta, Hymenoptera). Program version 4.0 beta, data version 39 (December 18, 2012). Digital Entomological Information, Müncheberg.
  2. ^ Taeger, A., Blank, SM & Liston, AD 2006: European Sawflies (Hymenoptera: Symphyta) - A Species Checklist for the Countries. Pp. 399-504. In: Blank, SM, Schmidt, S. & Taeger, A. (eds): Recent Sawfly Research: Synthesis and Prospects. Goecke & Evers, wine presses.
  3. a b c d e Blank, SM 2002: Biosystematics of the extant Xyelidae with particular emphasis on the Old World taxa (Insecta: Hymenoptera). Dissertation, Free University of Berlin.
  4. a b Taeger, A .; Blank, SM; Liston, AD 2010: World Catalog of Symphyta (Hymenoptera). Zootaxa 2580: 1-1064.
  5. ^ Dyar, HG 1898: Description of an unusual saw-fly larva belonging to the Xyelinae. Psyche 8 (265): 212-214.
  6. a b c d e Burdick, DJ 1961: A taxonomic and biological study of the genus Xyela Dalman in North America. University of California Publications in Entomology 17 (3): 285-355.
  7. ^ Weber, H. & Weidener, H. 1974: Outline of Insectology. G. Fischer, Stuttgart, study edition, 640 pp.
  8. ^ Naumann, ID (ed.) 1991: The Insects of Australia. Melbourne Univ. Press, Carlton 1-2: 1-1137
  9. ^ Scoble, MJ 1995: The Lepidoptera. Form, function and diversity. Nat. Hist. Mus., London and Oxford Univ. Press, Oxford et al., 404 pp. (reprint with corrections)
  10. ^ Yates, HO III & Smith, DR 2009: History, Distribution, Damage, and Life Cycle of a Pine Shoot Gall Sawfly, Xyela gallicaulis (Hymenoptera: Xyelidae). Journal of Entomological Science 44: 276-283.
  11. a b c Shinohara, A. 1995: The Sawfly Genus Pleroneura (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 63 (4): 825-840.
  12. a b c d Smith, DR & Schiff, NM 1998: The genera Macroxyela Kirby and Megaxyela Ashmead (Hymenoptera: Xyelidae) in North America. Proceedings of the Entomological Society of Washington 100: 636-657.
  13. a b c Shinohara, A. 1992: The Sawfly Genus Megaxyela (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 60 (4): 783-796.
  14. a b Smith, DR 1964: Description of the male of Xyelecia nearctica , with comments on the genus (Hymenoptera: Xyelidae). Pan-Pacific Entomologist 40 (1): 54-56.
  15. ^ Blank, SM, Shinohara, A. & Byun, B.-K. 2005: The East Asian Xyela species (Hymenoptera: Xyelidae) associated with Japanese Red Pine ( Pinus densiflora ; Pinaceae) and their distribution history. Insect Systematics & Evolution 36: 259-278.
  16. a b c d Blank, SM, Shinohara, A. & Altenhofer, E. 2013: The Eurasian species of Xyela (Hymenoptera, Xyelidae): taxonomy, host plants and distribution. Zootaxa 3629: 1-106 PDF (abstract) .
  17. Schulmeister, p. 2001: Functional morphology of the male genitalia and copulation in lower Hymenoptera, with special emphasis on the Tenthredinoidea s. st. (Insecta, Hymenoptera, 'Symphyta'). Acta Zoologica 82: 331-349.
  18. Hara, H. 1996: Akaezomatsu no shin gaichu marunaginatahabachi. (In Japanese). [ Pleroneura piceae , a new pest of Picea glehnii .] Shinrin Hogo [= Forest Protection] 254: 25-26.
  19. ^ Ree, B. 2012. Insects. Texas Pecan Pest Management Newsletter 12 (1): [1-3].
  20. ^ Ree, B. 2014. Sawfly. Texas Pecan Pest Management Newsletter 14 (2): [1-3].
  21. ^ Ree, B. 2016: Insects. Texas Pecan Pest Management Newsletter 16 (2): [1-3]. PDF
  22. a b c d Blank, SM, Kramp, K., Smith, DR, Sundukov, YN, Wei, M. & Shinohara, A. 2017: Big and beautiful: the Megaxyela species (Hymenoptera, Xyelidae) of East Asia and North America. European Journal of Taxonomy 348: 1-46. doi : 10.5852 / ejt.2017.348
  23. Sharkey, MJ, Carpenter, JM, Vilhelmsen, L., Heraty, J., Liljeblad, J., Dowling, APG, Schulmeister, S., Murray, D., Deans, AR, Ronquist, F., Krogmann, L ., Wheeler, WC 2012: Phylogenetic relationships among superfamilies of Hymenoptera. Cladistics 28 (2012) 80-112. doi : 10.1111 / j.1096-0031.2011.00366.x
  24. Togashi, I. 1972: Discovery of the genus Xylelecia Ross (Hymenoptera, Xyelidae) from Japan. Kontyû 40: 87-89.
  25. Shinohara, A. 1998: Collection records of two rare xyelid sawflies (Hymenoptera, Symphyta) in Japan. Japanese Journal of Systematic Entomology 4: 389-390.
  26. Smith, DR 1967: A review of the larvae of Xyelidae, with notes on the family classification (Hymenoptera). Annals of the Entomological Society of America 60: 376-384.
  27. a b Blank, SM 2002: The Western Palaearctic Xyelidae (Hymenoptera). Pp. 197-233. In: Viitasaari, M. (ed.): Sawflies (Hymenoptera, Symphyta) I. A review of the suborder, the Western Palaearctic taxa of Xyeloidea and Pamphilioidea. Tremex, Helsinki.
  28. Shinohara, A. 2016: The sawfly genus Pleroneura (Hymenoptera, Xyelidae) of Japan: P. itoi n. Sp. and a key to species. Zootaxa 4121 (4): 495-500. doi : 10.11646 / zootaxa.4121.4.9
  29. ^ A b Smith, DR, Ohmart, CP & Dahlsten, DL 1977: The fir shoot-boring sawflies of the genus Pleroneura in North America (Hymenoptera: Xyelidae). Annals of the Entomological Society of America 70: 761-767.
  30. Vieillot, LP 1807–1809 Histoire naturelle des oiseaux de l'Amérique septentrionale, contenant un grand nombre d'espèces décrites ou figurées pour la première fois. Chez Desray, Paris, tome 1: [7] + [i] –iv + [1] –90, tabs 1–57, tome 2: [4] + [i] –ii + [1] –74, tabs 58 -124. doi : 10.3931 / e-rara-7221
  31. ^ Smith, DR 1990: A new Xyela (Hymenoptera: Xyelidae) from western United States. Entomological News 101: 9-12.
  32. Blank, SM, Kramp, K. & Shinohara, A. 2017: Xyela fusca spec. nov. from Japan elucidates East Asian – North American relationships of Xyela (Hymenoptera, Xyelidae). Zootaxa 4303 (1): 103-121. doi : 10.11646 / zootaxa.4303.1.6
  33. Blank, SM & Kramp, K. 2017: Xyela davidsmithi (Hymenoptera, Xyelidae), a New Pine Catkin Sawfly with an Unusual Host Association from the Sierra Nevada. Proceedings of the Entomological Society of Washington 119: 703-717.
  34. Hubert Pschorn-Walcher, Ewald Altenhofer: Long-term larvae collections and breeding of plant wasps (Hymenoptera, Symphyta) in Central Europe. In: Linz biological contributions. 32nd year, Linz 2000, pp. 273-327 ( PDF on ZOBODAT ).
  35. ^ Blank, SM & Burger, F. 1996: Notable Hymenoptera finds from East Germany (Hymenoptera, Symphyta and Aculeata). Contributions from the hymenopterist conference in Stuttgart, 1996, pp. 6-7.
  36. Liston, AD & Blank, SM 2006: New and little-known British Xyelidae and Tenthredinidae (Hymenoptera, Symphyta). Entomologist's Monthly Magazine 142: 219-227.
  37. Shcherbakov, DE 2008: Insect recovery after the Permian / Triassic crisis. Alavesia 2: 125-131.
  38. Rasnitsyn, AP 1995: Tertiary sawflies of the tribe Xyelini (Insecta: Vespida = Hymenoptera: Xyelidae) and their relationship to the mesozoic and modern faunas. Contributions in Science 450: 1-14. (PDF; 1.4 MB)
  39. Krassilov, V., Tekleva, M., Meyer-Melikyan, N., Rasnitsyn, AP 2003: New pollen morphotype from gut compression of a Cretaceous insect, and its bearing on palynomorphological evolution and palaeoecology. Cretaceous Research 24 (2003): 149-156. doi : 10.1016 / S0195-6671 (03) 00029-6

literature

  • Blank, SM 2002: Biosystematics of the extant Xyelidae with particular emphasis on the Old World taxa (Insecta: Hymenoptera). Dissertation, Free University of Berlin.
  • Blank, SM 2002: The Western Palaearctic Xyelidae (Hymenoptera). pp. 197-233. In: Viitasaari, M. (ed.): Sawflies (Hymenoptera, Symphyta) I. A review of the suborder, the Western Palaearctic taxa of Xyeloidea and Pamphilioidea. Tremex Publ., Helsinki.
  • Blank, SM & Kramp, K. 2017: Xyela davidsmithi (Hymenoptera, Xyelidae), a New Pine Catkin Sawfly with an Unusual Host Association from the Sierra Nevada. Proceedings of the Entomological Society of Washington 119: 703-717.
  • Blank, SM; Kramp, K. & Shinohara, A. 2017: Xyela fusca spec. nov. from Japan elucidates East Asian – North American relationships of Xyela (Hymenoptera, Xyelidae). Zootaxa 4303 (1): 103-121. doi : 10.11646 / zootaxa.4303.1.6
  • Blank, SM; Kramp, K .; Smith, DR; Sundukov, YN; Wei, M. & Shinohara, A. 2017: Big and beautiful: the Megaxyela species (Hymenoptera, Xyelidae) of East Asia and North America. European Journal of Taxonomy 348: 1-46. doi : 10.5852 / ejt.2017.348
  • Blank, SM, Shinohara, A. & Byun, B.-K. 2005: The East Asian Xyela species (Hymenoptera: Xyelidae) associated with Japanese Red Pine ( Pinus densiflora ; Pinaceae) and their distribution history. Insect Systematics & Evolution 36: 259-278. On-line
  • Blank, SM, Shinohara, A. & Altenhofer, E. 2013: The Eurasian species of Xyela (Hymenoptera, Xyelidae): taxonomy, host plants and distribution. Zootaxa 3629: 1-106 PDF (abstract) .
  • Burdick, DJ 1961: A taxonomic and biological study of the genus Xyela Dalman in North America. University of California Publications in Entomology 17 (3): 285-355.
  • Shinohara, A. 1992: The Sawfly Genus Megaxyela (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 60 (4): 783-796.
  • Shinohara, A. 1995: The Sawfly Genus Pleroneura (Hymenoptera, Xyelidae) in East Asia. Japanese Journal of Entomology 63 (4): 825-840.
  • Smith, DR, Ohmart, CP & Dahlsten, DL 1977: The fir shoot-boring sawflies of the genus Pleroneura in North America (Hymenoptera: Xyelidae). Annals of the Entomological Society of America 70: 761-767.
  • Smith, DR & Schiff, NM 1998: The genera Macroxyela Kirby and Megaxyela Ashmead (Hymenoptera: Xyelidae) in North America. Proceedings of the Entomological Society of Washington 100 (4): 636-657.
  • Taeger, A .; Blank, SM & Liston, AD 2010: World Catalog of Symphyta (Hymenoptera). Zootaxa 2580: 1-1064. (PDF; 23 kB)

Web links

Commons : Xyelidae  - Collection of images, videos, and audio files