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Cynipidae Parasitic Gall wasps

Gall wasps

Not all parasite wasps are invertabrate parasitoids, In fact the most obvious to the nature watcher are the Cynipidae family which are true parasites of plants. Although their host is totally different from the other families of parasitoids they still inject venom with their eggs which modifies the host tissues to form strangely shaped growths that are known as galls. These galls provides the larvae with nourishing food and protection from the outside world. 

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Ritchie (1993) noted that all phytophagous Cynipoidea are placed in this family.  Members of Cynipidae either form galls on various plants or live as inquilines in the galls of other gall-forming insects.

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Many gall wasps have two generations per year with each generation having a slightly different body shape and may well also reproduce differently, eg. It is quite common for the winter/ early spring generation to be parthenogenic (entirely female), and for those females to lay unfertilised eggs as well as normal fertilised ones that will develop into sexual males and females accordingly, which will appear in the summer/autumn generation.

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It is well known that wasps have a haplo-diploid genetic structure that will produce females from fertilised eggs and males from the unfertilised eggs but the trick with these wasps is the ability to make females from unfertilised eggs. This is accomplished by the chromosomes inside the nucleus of the egg replicating as normal and then the two daughter cells coming into contact with each other, and the cellular membrane breaking down with the two haploid chromosones pairing up to form the full compliment of chromosomes and a diploid cell is created. Once this is done then the egg can divide as normal.

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Some cynapids will produce different shaped galls in different plants according to which generation they are born in.

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As well as fungus mycilia, which they can not run away from, gall wasp larvae frequently fall prey to a huge array of parasite parasitoids known as hyperparasites and cleptoparasites known as inquilines, which rather than attacking the larvae directly, will live in the gall along with them and steal their food. These communities can become very complex, involving many layers of inter-dependancy.

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Andricus kollari agamic wasp IMG_3605_ed

An example of the above is the Oak Marble gall (Andricus kollari) that in one gall can contain the cynapid wasp as the host that made the gall; upto 5 species of inquilines (Ceroptres arator, Synergus gallaepomiformis, S. pallidipennis, S. reinhardi and S. umbraculus) eating the hosts food; aswell as upto 13 parasitoid species (Eurytoma brunniventris, Sycophila biguttata, S. variegata, Megastigmus dorsalis, M. stigmatizans, Torymus geranii, T. auratus, Caenacis lauta, Hobbya stenonota, Mesopolobus amaenus, M. fasciiventris, M. sericeus, Eupelmus urozonus) living on the host, inquilines and each other.

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Also the hyperparasitoids of species that have two generations per year can also change their body shape to follow their hosts through the generations, for example; by one generation to have a longer ovipositor than the previous one to reach the host.

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There is disagreement about the limits of Cynipidae.  Most European authors (eg., Quinlan 1979) have a broad concept of the family and recognise 4 subfamilies:  Himalocynipinae, Austrocynipinae, Pycnostigmatinae and Cynipinae.  In North American authors usually have a narrower concept, limiting the family to gall formers and inquilines.  Ritchie (1993) considered that if this is accepted, then the other subfamilies should be given family status.  However, this awaits further work on higher classification.

Cynipidae (Charipinae) (Cynipoidea).

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Ritchie (1993) considered the Charipinae as a distinct family, Charipidae.  Species are defined by host association and negative morphological attributes (eg., the lack of sculpture).  Adults are usually very small (1-3 mm) and are difficult to identify.

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Two subfamilies, Alloxystinae and Charipinae were identified by Ritchie (1993).  In this report we have considered the Charipinae as a subfamily of Cynipidae.  The former considered species of Alloxystinae as hyperparasitoids of Aphidae (Homoptera) through Braconidae (Aphidiinae) and Aphelinidae.  The early stages of the hyperparasitoid are spent inside its primary host, which is inside an aphid, but after the braconid or aphelinid pupates, the last larval instar of the alloxystine feeds externally and pupates inside the inflated aphid skin.  Members of Charipinae are hyperparasitoids of Psylloidea (Homoptera) through Encyrtidae (Hérard 1986). There are 4 genera and circa 200 species worldwide. 

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Cynipidae (Cynipinae) (Cynipoidea)

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Ritchie (1993) noted that many female Cynipinae have the hypopygial spine extremely well developed.  However, males are difficult to identify, and it is often impossible to place them to genus.  Gall-forming species on oaks and roses can be recognized by the extremely narrow pronotum (median dorsal length less than 1/7th the lateral height).  Most species of Cynipinae form galls on Quercus (Fagaceae) in warm-temperate and subtropical regions.  Sexual and agamic generations alternate in many oak gall formers.  A few genera form galls on Compositae and Rosaceae, but rarely other plants. Cynipinae is restricted to the Holarctic region except for one species in South Africa (Rhoophilus loewi Kieffer) and 8 species in South America (Myrtopsen spp., Plagiotrochus suberi Weld).  The Cynipinae contains circa 93 genera and 1,200 species worldwide.

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Checklist of UK Recorded: Cynipidae   (Family) cynipids

 

  • Andricus albopunctatus  f. agamic (Schlechtendal, 1870) a gall wasp

  • Andricus ambiguus  f. agamic (misident) Trotter a gall wasp

       Andricus ambiguus  f. sexual (misident) Trotter a gall wasp

  • Andricus amenti  f. sexual Giraud, 1859 Hairy-catkin gall

  • Andricus anthracina  f. agamic (Curtis, 1838) Oyster-gall causer

       Andricus anthracina  f. sexual (Curtis, 1838) a gall wasp

       Andricus callidoma  f. sexual (Hartig, 1841) Tufted-gall causer

       Andricus corruptrix  f. sexual (Schlechtendal, 1870) a gall wasp

       Andricus curvator  f. sexual Hartig, 1840 Curved-leaf gall causer

       Andricus fecundator  f. sexual (Hartig, 1840) Hairy-catkin gall causer

  • Andricus glandulae  f. agamic (Hartig, 1840) Thatched gall causer

       Andricus glandulae  f. sexual (Hartig, 1840) a gall wasp

  • Andricus grossulariae  f. sexual (Giraud) a gall wasp

  • Andricus inflator  f. agamic Hartig, 1840 Globular gall causer

       Andricus inflator  f. sexual Hartig, 1840 Twig gall causer

       Andricus kollari  f. sexual (Hartig, 1843) a gall wasp

       Andricus lignicola  f. sexual (Hartig, 1840) a gall wasp

  • Andricus lucidus  f. agamic (Hartig, 1843) Hedgehog gall causer

  • Andricus nudus  f. agamic Adler, 1881 Malpighi�s gall causer

       Andricus nudus  f. sexual Adler, 1881 Bald-seed gall causer

  • Andricus ostreus  f. agamic (Hartig, 1840) Oyster-gall causer

  • Andricus quadrilineatus  f. agamic Hartig, 1840 Furrowed-catkin gall causer

  • Andricus quercuscalicis  f. agamic (Burgsdorf, 1783) Knopper gall causer

       Andricus quercuscalicis  f. sexual (Burgsdorf, 1783) a gall wasp

  • Andricus quercuscorticis  f. agamic (Linnaeus, 1761) Bark-gall causer

       Andricus quercuscorticis  f. sexual (Linnaeus, 1761) The bud-gall causer

  • Andricus quercusradicis  f. agamic (Fabricius, 1798) Truffle gall causer

       Andricus quercusradicis  f. sexual (Fabricius, 1798) Knot gall causer

  • Andricus quercusramuli  f. agamic (Linnaeus, 1761) The autumn-gall causer

       Andricus quercusramuli  f. sexual (Linnaeus, 1761) Cotton-wool gall causer

  • Andricus rhizomae  f. agamic (Hartig, 1843) a gall wasp

  • Andricus seminationis  f. agamic (Giraud, 1859) Spindle gall causer

  • Andricus solitarius  f. agamic (Fonscolombe, 1832) Hairy-spindle gall causer

       Andricus solitarius  f. sexual (Fonscolombe, 1832) Bisected gall wasp

  • Andricus testaceipes  f. agamic Hartig, 1840 Red barnacle-gall causer

       Andricus testaceipes f. sexual Hartig, 1840 Leaf-vein gall causer

  • Aphelonyx cerricola  f. agamic Giraud a gall wasp

  • Aulacidea hieracii  (Bouch�, 1834) a gall wasp

  • Aulacidea nibletti  Quinlan, 1969 a gall wasp

  • Aulacidea pilosellae  (Kieffer, 1901) a gall wasp

  • Aulacidea subterminalis  Niblett, 1946 a gall wasp

  • Aulacidea tragopogonis  (Thomson, 1877) a gall wasp 

  • Aylax minor  Hartig, 1840 a gall wasp

  • Aylax paperveris  (Perris, 1839) a gall wasp

  • Biorhiza pallida  f. agamic (Olivier, 1791) a gall wasp

       Biorhiza pallida  f. sexual (Olivier, 1791) Oak-apple causer

  • Callirhytis bella  (Dettmer, 1930) a gall wasp

  • Callirhytis erythrocephala  (Giraud, 1859) a gall wasp 

  • Callirhytis erythrostoma  Dettmer, 1933 a gall wasp

  • Ceroptres arator  Hartig, 1841 a cynipid inquiline

  • Cynips agama  f. agamic Hartig, 1840 Yellow-pea gall causer

  • Cynips disticha  f. agamic Hartig, 1840 Two-cell gall causer

       Cynips disticha  f. sexual Hartig, 1840 a gall wasp

       Cynips divisa  f. sexual Hartig, 1840 Red-current gall causer

  • Cynips longiventris  f. agamic Hartig, 1840 Striped-pea gall causer

       Cynips longiventris  f. sexual Hartig, 1840 Green velvet-bud gall causer

  • Cynips quercusfolii  f. agamic Linnaeus, 1758 Cherry gall causer

       Cynips quercusfolii  f. sexual Linnaeus, 1758 Violet-egg gall causer

       Diplolepis eglanteriae sens. lat. Smooth Pea Gall

  • Diplolepis spinosissimae  (Giraud, 1859) a gall wasp 

  • Isocolus fitchi  (Kieffer, 1898) a gall wasp

  • Isocolus jaceae  (Schenck, 1862) a gall wasp

  • Isocolus rogenhoferi  Wachtl, 1880 a gall wasp

  • Isocolus scabiosae  (Giraud, 1859) a gall wasp

  • Liposthenus glechomae  (Kieffer, 1898) a gall wasp

  • Neuroterus albipes  f. agamic (Schenck, 1863) Smooth-spangle causer

       Neuroterus albipes  f. sexual (Schenck, 1863) Schencks a gall causer

  • Neuroterus aprilinus  f. agamic (Giraud, 1859) a gall wasp
    Neuroterus aprilinus  f. sexual (Giraud, 1859) April-bud gall causer

  • Neuroterus numismalis  f. agamic (Geoffroy in Fourcroy,1785) Silk-button causer

       Neuroterus numismalis  f. sexual (Geoffroy in Fourcroy,1785) Blister-gall         causer

       Neuroterus quercusbaccarum  f. sexual (Linnaeus, 1758) Currant gall causer

  • Neuroterus tricolor  f. agamic (Hartig, 1841) Cupped spangle causer

       Neuroterus tricolor  f. sexual (Hartig, 1841) Hairy-pea causer

  • Periclistus brandtii  (Ratzeburg, 1832) a cynipid inquiline

  • Periclistus caninae  (Hartig, 1840) a cynipid inquiline

  • Periclistus spinosissimae  Dettmer, 1924 a cynipid inquiline

  • Phanacis caulicola  (Hedicke, 1939) a gall wasp

  • Phanacis centaureae  Foerster, 1860 a gall wasp

  • Phanacis hypochoeridis  (Kieffer, 1887) a gall wasp

  • Saphonecrus connatus  (Hartig, 1840) a cynipid inquiline

  • Synergus albipes  Hartig, 1841 a cynipid inquiline

  • Synergus apicalis  Hartig, 1841 a cynipid inquiline

  • Synergus clandestinus  Eady, 1952 a cynipid inquiline

  • Synergus crassicornis  Curtis a cynipid inquiline

  • Synergus evanescens  Mayr, 1872 a cynipid inquiline

  • Synergus gallaepomiformis  (Boyer de Fonscolombe, 1832) a cynipid inquiline 

  • Synergus incrassatus  Hartig, 1840 a cynipid inquiline

  • Synergus nervosus  Hartig, 1840 a cynipid inquiline

  • Synergus pallicornis  Hartig, 1841 a cynipid inquiline

  • Synergus pallidipennis  Mayr, 1872 a cynipid inquiline

  • Synergus reinhardi  Mayr, 1872 a cynipid inquiline

  • Synergus rotundiventris  Mayr, 1872 a cynipid inquiline

  • Synergus ruficornis  Hartig, 1840 a cynipid inquiline

  • Synergus thaumacerus  (Dalman, 1823) a cynipid inquiline

  • Synergus umbraculus  (Olivier, 1791) a cynipid inquiline

  • Trigonaspis megaptera  f. agamic (Panzer, 1801) Kidney gall causer

       Trigonaspis megaptera  f. sexual (Panzer, 1801) Pink-wax gall causer

  • Xestophanes brevitarsis  (Thomson, 1877) a gall wasp

  • Xestophanes potentillae  (Retzius in Degeer, 1783) a gall wasp

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Andricus kollari larvae IMG_8680_edited.
Andricus fecundator larvae IMG_0060_edit

mature larvae of (left) Andricus kollari and (right) Andricus fecundator

Cynipidae (Eucoilinae) (Cynipoidea)

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Ritchie (1993) treated Eucoilinae as a distinct family Eucoilidae.  They noted that the scutellum is medially with a characteristic round or teardrop-shaped raised plate ("cup" of old literature).  They considered this the largest "family" of Cynipoidea.

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Eucoilids are internal parasitoids of calyptrate Diptera larvae, emerging from the puparium.  Many species are associated with dung or rotting fruit, but the family is not restricted to these habitats.  The number of species worldwide is unknown, but most are found in the tropics, where they are abundant.  In North America there are 15 genera and 78 species (10 in Canada).  Specialists believe that 2-3X as many species exist.

Immature Stages of Cynipoidea

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The Egg.

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The eggs of the parasitic members of the superfamily are uniformly of the stalked type, with the stalk, which is situated at the anterior end, ranging in length from less than that of the main body, as in Charips sp., to several times its length.  That of Figites anthomyiarum is elongated and somewhat con­stricted in the middle and has a stalk of about equal length.  In Eucoila keilini, the stalk is twice the length of the egg body, and in Ibalia leucospoides it is about four times as long.  The chorion is thin and transparent and without surface ornamentation.

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First instar Larva.

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The known first instar larvae of the superfamily are of several distinct types.  The "polypodeiform" larva of I. leucospoides is unusually elongated, with a relatively large head, which is somewhat flattened dorsoventrally and is provided with falcate mandibles.  Of the 13 body segments, all except the last bear a pair of fleshy, finger like processes ventrally, which are of uniform length.  There are no integumentary spines or setae.  The last abdominal segment is prolonged into a dorsally curved tail equal in length to the four preceding segments. 

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The first instar larvae of the several genera of Eucoilinae that have been described are distinctive and readily recognized and have been designated as "eucoiliform."  The essential characters of this larval type that distinguish it from the polypodeiform are the long, paired, fleshy ventral processes on the thoracic segments only and an exceptionally long tapering tail.  In Eucoila keilini, the head is large and somewhat conical, and the body segmentation is indistinct.  The fleshy thoracic processes are about half the length of the body and taper to a blunt point.  The posterior segments of the abdomen are very much narrowed, being only very slightly wider than the base of the tail, and the tail itself is appreciably longer than the thoracic processes, is curved ventrally, and terminates in a sharp point.  The segment immediately preceding the tail bears a fleshy conical lobe on the median ventral line.  The integument of the posterior abdominal segments bears numerous short sclerotized spicules.

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The larva of Cothonaspis rapae described by James and Malchanova is similar to that of Eucoila.  There are at least seven distinct abdominal segments in addition to the two or more that make up the tail, and the latter organ is nearly as long as the entire body.  There are a few setae upon the thoracic processes, and the distal third of the tail bears numerous spines. 

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The eucoiliform larva of Kleidotoma marshalli figured and described by James has 12 apparent body segments, and the body is much more slender than that of Eucoila or of Cothonaspis.  The head is large, with the mouth opening distinctly ventral.  The fleshy thoracic processes are much shorter, being equal only to one segment in length, and they bear no setae.  Stout setae are present on the distal portion of the tail.  The anal opening is indicated on the dorsum of the eighth abdominal segment. 

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The first instar larva of Figites anthomyiarum is modified eucoiliform the thoracic processes being even more reduced than in Kleidotoma  with the prothoracic pair only as long as wide, and the tail is short, almost cylindrical, and bluntly rounded rather than pointed at the tip.  The segmentation is distinct, and the 12 body segments preceding the tail bear fleshy spines on the dorsum, these being largest in the mid abdominal region. 

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In the Charipinae, the only first instar larva that was described by 1940 was that of Charips sp. by Haviland.  This larva has few characters in common with other members of the family and must be considered as a modified caudate form.  The head is large, equaling the thoracic segments in width, and is produced anterioventrally into a conical "proboscis."  There are three pairs of sclerotized "nodules" ventrally and one pair dorsally, which presumably are sensory organs.  The mandibles are long and slender.  There are 13 body segments, of which the first 12 diminish gradually in length and width caudad.  The last abdominal segment is broad at its base, tapers sharply, and terminates in a cylindrical, ventrally directed tail.  This last segment is equal in length to the 9 abdominal segments preceding it.  The anal opening is large, situated dorsally at the base of the last segment, and encircled by a sclerotized ring.  The head, and the body segments except the last, are heavily sclerotized, and each of the body segments telescopes into the one preceding it. 

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None of the first instar larvae of the family that has been studied possesses any indication of a tracheal system, and respiration is consequently by diffusion only.

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Second instar Larvae.

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The second instar larvae of the Cynipoidea reveal differences that are almost as great as those in the first instar.  In Ibalia, the paired ventral processes have disappeared, and the tail is some­what reduced.  The second instar larva of Cothonaspis rapae retains the eucoiliform characters of the first instar, but the segmentation is more distinct, whereas in Kleidotoma marshalli there is a change to the polypodeiform.  The head of the latter is very large, exceeding the body segments in width and length, and the segmentation is exceptionally distinct.  The minute paired processes occur ven­trally on the first 10 body segments.  The larva of Figites anthomyiarum is similar to Kleidotoma in all essential respects though the head is small, the segmentation indistinct, and the tail situated ventrally and at right angles to the axis of the body.  An internal tracheal system is present, but there are no spiracles.  The second instar larva of Charips sp.  is still of the caudate form, though the heavy sclerotization of the integument is lacking.  Each of the thoracic segments bears a pair of small processes ventrally, and a pair of large conical structures is present at the posterior ventral margin of the head. 

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James studied the early instars of cynipoid larvae and came to the conclusion that the eucoiliform first instar larvae of Eucoila and Cothonaspis are derived from eggs which hatch in the middle of the protopod stage of embryonic development, whereas the polypodeiform larvae correspond, as the name implies, to the polypod phase.  The form of the Figites larva, with its reduced appendages and distinct segmentation, indicates hatching in a later embryonic stage than does any of the other species discussed.  The Charips larva, being devoid of appendages, is regarded as preceding or as being a very early form of the protopod stage.  This view of the stage of development at the time of hatching is borne out by the occurrence of polypodeiform second instar larvae following the eucoiliform first instar in Figites and Kleidotoma.

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Only three instars have been distinguished in the species studied, with the excep­tion of Ibalia leucospoides, which has four.  The third instar larva of Ibalia is cylindri­cal in form with the tail still further reduced and may be readily recognized by the presence of spiracles on the second and third thoracic segments. 

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Mature Larvae.

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The mature larvae of the various species differ in only relatively minor characters.  That of Ibalia has the integument smooth and shining except in the pleural areas of the second to eleventh body segments, which bear rounded "bases" studded with minute spines.  The mandibles are tridentate, whereas they ere bidentate in Figites and Charips.  There is a somewhat surprising variation in the number and position of the spiracles.  Ibalia has 10 pairs, situated on the second and third thoracic and the first eight abdominal segments; Eucoila keilini and Figites anthomyiarum have nine pairs, on the last two thoracic and the first seven abdominal segments; Cothonaspis rapae eight pairs, on the third thoracic and the first seven abdominal segments; and Charips sp. has only six pairs, on the second and third thoracic and the first, second, fourth, and sixth abdominal segments. 

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Information courtesy of  University of California, Riverside

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