Kokkocynips imbricariae (agamic)

Kokkocynips coxii (agamic)

Kokkocynips rileyi (agamic)

Kokkocynips difficilis (agamic)

Kokkocynips attractans (sexgen)

Kokkocynips decidua (agamic)

Kokkocynips panamensis (agamic)

Kokkocynips doctorrosae (agamic)

Kokkocynips imbricariae (Ashmead, 1896) n. comb.

Andricus imbrecariae Ashmead, 1896. Proc. U.S. Nat. Mus., vol. 19, p.122.
Holcaspis fasciata Bassett, 1900. Trans. Amer. Ent. Soc., vol, 26, p.328. [Syn. by Weld, 1926b: 43].
Dryocosmus fasciatus (Bassett): Weld, 1922a. Proc. U.S. Nat. Mus., vol. 61, art. 18, p.7.
Dryocosmus imbricariae* (Ashmead): Weld, 1926b. Proc. U.S. Nat. Mus., 68, 43.* The original spelling ‘imbrecariae’ is a typographical error and Ashmead had corrected it in his own copy of the paper to align it with the name of the host oak (Weld, 1926b)

Gall: according to the original description (Ashmead 1896) the gall is globular, brownish and hard, 7-10 mm in diameter. The surface of the gall has a striped pattern, and is hence commonly known as the banded bullet gall. The galls typically grow in clusters from a fissure in twigs from the previous year’s growth. The larval cell is nearly always closely cemented to the thick, hard, gall epidermis, but in a single instance there were a few fibers separating it.

Host plant: the species was described from galls collected on Quercus imbricaria and Q. ilicifolia. Weld (1926b) added Quercus marilandica, Q. catesbaei, Q. texana and Q. laurifolia to the host plant records, and later mentioned that it galled all eastern US red oak species (Weld 1959).

Biology: known only from an asexual generation. Adults emerge from galls from September to October. Galls collected in the autumn produced adults in the following year, but some emerged a year later after diapause (Ash-mead 1896).

Distribution:  USA  and  Canada;  originally  described  from  Missouri,  and  also  recorded  from  many  localities from Ontario southwards to Florida and westwards to Texas (Weld 1926b).

Kokkocynips coxii (Bassett, 1881) n. comb.

Cynips coxii Bassett, 1881. Canad. Ent., vol. 13, p.112. [Syn. by Kinsey, 1922: 123-125].
Andricus coxii (Bassett), Ashmead, 1885. Trans. Amer. Ent. Soc., vol. 12: 295.
Andricus coxi Bassett, 1900. Trans. Amer. Ent. Soc., vol. 26, p.320. 
Andricus bassetianus Dalla Torre and Kieffer, 1902. in Wytsman Gen. Ins. Hym. Cynipidae, p.61. [Syn. by Kinsey, 1922: 123-125].
Plagiotrichus (sic.) coxii (Bassett), Kinsey, 1922. Ind. Univ. Stud. 53, pp.122-5.
Plagiotrochus coxii var translatus Kinsey, 1922: 126 n. syn.
Dryocosmus coxii (Bassett): Weld, 1926b. Proc. U.S. Nat. Mus., 68(10): 41.
Dryocosmus coxii var translatus (Kinsey) Weld, 1951: 646.

Gall: the galls are elongate to globose twig swellings up to 20 mm. in diameter and 65 mm in length; on older twigs from previous years’ growth. Multilocular, smooth surface covered with normal bark. Larval cells closely embedded in the less solid part of the tissue (Kinsey 1922).

Host plant: Quercus emoryi and Quercus hypoleucoides (=hypoleuca Engelm.).

Biology: only known from an asexual generation. The original source is somewhat imprecise about gall phenology, saying that the whole life cycle takes more than one year, as the insects emerge in midwinter and the young galls appear immediately.

Distribution: Arizona, New Mexico and possibly Texas (Kinsey 1922).

Remarks: Kinsey (1922) transferred this species described by Bassett to the genus Plagiotrichus (sic.) (=Plagiotrochus Mayr) under the mistaken belief that this genus, described from the Western Palaearctic, was also present in North America. In his account of this species Kinsey gave a short re-description, summarized the data on biology and distribution, and described two new varieties. Kinsey (1922) emphasized the particularity of this species as he had some doubts about its correct generic placement, but his placement was finally made based on the structure of the gall. Weld later transferred the species to Dryocosmus, on the basis that “the flies run to Dryocosmus in the Tierreich key” (a reference to the work by Dalla Torre and Kieffer (1910); Weld 1926b).

Kokkocynips rileyi (Ashmead, 1896) n. comb.(Figs. 8, 10D, 11F)

Andricus rileyi Ashmead, 1896. Proc. U.S. Nat. Mus, vol. 19, p. 121.
Dryocosmus rileyi (Ashmead):Weld, 1926b. Proc. U.S. Nat. Mus., 68(10): 44.

Gall: the main body of the gall is globular, small (around 3 mm.), with a detachable yellowish fleshy knob on the dorsal side of the gall (Fig. 11F). Galls are attached to the midrib or lateral veins of a leaf by a slender stalk. The young gall body is yellowish, and gradually turns brownish following maturation. The larval cell occupies the whole of the interior (Ashmead 1896).

Host plant: this species was originally described from galls collected on Quercus rubra. It has subsequently been found on all eastern US red oaks (Weld 1926b, Weld 1959).

Biology: known only from an asexual generation. Galls collected in the autumn produced adults in September and October of the following year; in other samples adults emerged from January to March next year.

Distribution:  this  species  was  originally  described  from  Missouri. Weld  (1926b)  added  many  localities  from New York to Florida and westwards to Missouri and Texas.

Kokkocynips difficilis (Ashmead, 1887) n. comb.

Andricus difficilis Ashmead, 1887. in: Tr. Amer. Ent. Soc, V. 14 p.143.
Callirhytis difficilis (Ashm.): Mayr, 1902. Verh. Ges. Wien, v. 52, 289.

Gall: small (12 X 15 mm), irregularly rounded, densely rugose, reddish when young and turns grayish following gall maturation, slightly flattened at sides, the rugosities arranged transversally in five to six rows. These galls occur in clusters, emerging in rows from fissures or slits in the terminal twigs; when mature fall to the ground (Ashmead 1887).

Host plant: Quercus laurifolia, Q. cinerea [incana], Q. catesbaei [laevis], Q.nigra, Q. myrtifolia and Q. phellos (Weld 1959)

Biology: only known from the asexual generation. According to Ashmead (1887), the adults emerge from galls from August to September; Weld (1959) noted that adults also emerged later in the year up to December.

Distribution:  South-eastern  North  America,  from  Maryland  southwards  to  Florida  and  westwards  to  Texas (Burks 1979, pers. obs.).

Kokkocynips attractans (Kinsey, 1922) n. comb.

Andricus attractans Kinsey, 1922. in: Bull. Amer. Mus. Nat. His., vol. 46 p.281.
Callirhytis uvellae Weld, 1944. in: Proc. U. A. Nat. Mus., vol. 95 p.23 [Syn. in Dailey, 1969: 132].
Dryocosmus attractans (Kinsey): Weld, 1951: 644.
Callirhytis attractans (Kinsey): Dailey, 1969: 132

The molecular results placed one additional species, Callirhytis attractans (Kinsey 1922), within the same clade as species from Kokkocynips with 100% posterior probability support (Fig. 1). This is an interesting placement, as this species is known only from its sexual form, unlike other species in the genus. Accordingly, we transfer this species to Kokkocynips although we have not yet been able to study the type material. In the near future we hope to have the opportunity to re-describe this species.

Callirhytis uvellae Weld was synonymized with Dryocosmus attractans (Kinsey) by Dailey (1969) but Burks (1979) did not mention this synonymy, which we consider to be a mistake. The galls (Fig. 11G) are unilocular, about 2 mm in diameter and develop gregariously in spring on buds (Dailey 1969). The host oaks are of all the section Lobatae present in California, Q. agrifolia, Q. kelloggii and Q. wislizeni.

Kokkocynips decidua (Beutenmueller, 1913) n. comb.

Andricus decidua Beutenmueller, 1913. Insecut. Inscit. Menst., 1: 131-132.
Dryocosmus deciduus (Beutenmueller):Weld, 1926b. Proc. U.S. Nat. Mus., 68: 41.

Gall: galls are multilocular clusters of rounded or elongated swellings on the midribs of the leaves that contain from a few to about 40 oblong larval chambers, depending upon the size of the gall (Beutenmueller, 1913).

Host plant: initially described from red oak (Quercus rubra) and black oak (Q. velutina), but Weld (1959) lists this species from most section Lobatae oaks in eastern North America.

Biology: only known  from an asexual generation. The gall is  fully-grown  by  September or  October,  when  it falls to the ground.

Distribution: USA, from New England states southwards at least as far as North Carolina and westwards to the eastern Midwest.

Kokkocynips  panamensis induces  an  elongated  multilocular  swelling  gall  in  young  stems,  oval  to  spindle-shaped, with the same colour and surface texture as the bark of twigs (Figs. 4D & E). The galls have an average length of 35 mm (range 20 to 60 mm) and diameter 51 mm (range 30 to 85 mm) (n = 12). The internal structure of the gall is highly lignified and is multilocular, containing several larval cells (Figs. 4C, 4D and 11H). The outer layers crack and peel in older galls.

Host plant: Quercus salicifolia Née (Fagaceae, Quercus sect. Lobatae).Distribution: Kokkocynips panamensis was found from 1,400–1,800m above sea level at Volcán Barú and El Salto, Chiriquí Province, Panama.

Biology: Only the asexual generation of Kokkocynips panamensis is known. The galls are found between October and November, during the rainy season. The adult insects emerge from mature galls in November.

Kokkocynips doctorrosae Pujade-Villar, 2013

Gall: the gall (Fig. 11A) is unilocular on twigs, nearly spherical, 8–10 mm in diameter, slightly flattened basally, with a smooth surface, lacking pubescence, yellowish with concentric circular red-brown lines, apically reddish-pink. The gall stays soft when mature, but not fragile. Larval chamber ovate, 1.4 × 1.8 mm, with thin walls (0.16 mm), surrounded by a sponge-like tissue, located in the centre of the gall. Exit hole usually located laterally. There is always a longitudinal crack in the bark where the gall is inserted. 

Host plant: Quercus acutifolia, Q. castanea, Q. crassifolia and Q. crassipes (section Lobatae within Quercus). 

Biology: only known from an asexual generation. 

Distribution: currently known only from Mexico. Originally recorded only from the state of Michoacan. New records from Oaxaca, Puebla and Mexico states added here extend the distribution of this species.

Remarks: K. doctorrosae can be easily distinguished from the other known species of Kokkocynips by the regular well-marked reticulate sculpture on the metasomal terga. Congruent with the morphological similarity of both adults and galls, this species is sister to the Nearctic species K. imbricariae in the multi-locus phylogenetic analysis (Fig. 1), and forms a close cluster with K. imbricariae and K. difficilis based on cytb distances (all just over 5% divergent; Table 1).

Re-description and systematic re-appraisal of the genus Kokkocynips Pujade-Villar & Melika, (Hymenoptera: Cynipidae: Cynipini), including new combinations of Nearctic species and the description of a new species from Panama