Barucynips panamensis, a new genus and species of oak gallwasps (Hymenoptera, Cynipidae, Cynipini) from Panama, and description of one new species of Coffeikokkos

Abstract Barucynips panamensis Medianero & Nieves-Aldrey, a new genus and species of oak gallwasps (Hymenoptera: Cynipidae: Cynipini), is described from adults reared from galls on Quercus bumelioides in Panama. The new genus is taxonomically close to the recently described Coffeikokkos from Costa Rica, but differs from it and all of the described genera of Cynipini, by the shape and setation of the projecting part of the ventral spine of the hypopygium and by the sculpture of the propodeum. A new species of Coffeikokkos is also described from the same area, the Volcán Barú in Panama. Diagnostic characters, gall description, distribution, and biological data of the new genus and the two new species are given. The new genus is the first genus of oak gallwasps of the tribe Cynipini described in Panama.


Materials and methods
Study material. The adults studied were reared from galls collected on Quercus bumelioides Liebm. Samplings were made and material was collected from December 2007 to August 2010 at Volcán Barú, Chiriqui Province, Panama. The adult insects emerged from the galls in rearing cages under laboratory conditions. Voucher specimens and their galls were deposited in the entomology collections of the Museo Nacional de Ciencias Naturales, Madrid (Spain) and Maestria en Entomologia, Universidad de Panama (MEUP). The identification of the Quercus species was based on several key references (Burger 1977, D'Arcy 1987, Breedlove 2001) as well as on comparison with materials from the collection of the University of Panama and the Smithsonian Tropical Research Institute.
Specimen preparation. For observation under a scanning electron microscope (SEM), adult cynipids were dissected in 70% ethanol, air dried, mounted on a stub and coated with gold. Micrographs were taken with an FEI QUANTA 200 microscope (high vacuum technique) for several standardized views. Forewings were mounted in Euparal on slides and later examined under a Wild MZ8 stereo microscope. Images of adult habitus and gall dissections were taken with a NIKON Coolpix 4500 digital camera attached to a Wild MZ8 stereo microscope. Measurements were made with a calibrated micrometer scale attached to an ocular of the light microscope. The images will be deposited in the "morphbank.com" databank. Terminology of morphological structures and abbreviations follow Ronquist and Nordlander (1989), Ronquist (1995), Nieves-Aldrey (2001), and Liljeblad et al. (2008). For cuticular sculpture we follow Harris (1979). Measurements and abbreviations used include: POL (post-ocellar distance) is the distance between the inner margins of the posterior ocelli; OOL (ocellar-ocular distance) is the distance from the outer edge of a posterior ocellus to the inner margin of the compound eye.
Etymology. From Barú (the name of the volcano in Panama where the new genus was collected) and Cynips, referring to Cynips-groups, inside the tribe Cynipini.
Gender. Masculine Diagnosis and identification. By the 16 segmented antennae, head and mesosoma shape and sculpture, forewing venation and the association with Quercus bume-lioides, a tree common in the mountains of Central America, the new genus resembles the recently described genus Coffeikokkos Pujade-Villar & Melika, from Costa Rica. Barucynips differs, however, from that genus in some important characteristics which clearly separate the two genera. The main diagnostic characteristic that clearly allows the separation of Barucynips and Coffeikokkos is the shape of the hypopygial spine. The shape and setosity of the ventral spine of hypopygium of Barucynips is completely distinctive among all the exhibited by the described cynipids (Fig. 3). The projecting part of the hypopygial spine is lanceolate, pointed apically, and at least 4.0 times as long as broad, with a basal group of long setae, which reach the apex of the spine, forming a tuft, while Coffeikokkos present a projecting part of the hypopygial spine only 2.5 times as long as wide, uniformly broad, with parallel sides, rounded distally, with tuft of long supapical setae, reaching far beyond apex of spine. The new genus also differs from Coffeikokkos in the propodeal sculpture. The lateral carinae of the propodeum are subparallel in the anterior half and strongly divergent posteriorly in Coffeikokkos, without a median longitudinal carina, while the lateral propodeal carinae are much less strongly divergent posteriorly and a median longitudinal carina is present in Barucynips. Furthermore, the metatarsal claws of Coffeikokkos present a strong basal obtuse lobe, while the claws of Barucynips are toothed, with a small acute basal tooth. Other discriminant characteristics are as follows: the ventral margin of clypeus is strongly projected on mandibles (only slightly so in Coffeikokkos); the malar area has a strong alutaceous sculpture (Fig. 1B) and lacks irradiating striae from the clypeus (a few but clearly impressed striae are present in Coffeikokkos). The new genus is also taxonomically close to the genera of the Cynips group, Cynips Linnaeus (=Antron Kinsey, Besbicus Kinsey), Atrusca Kinsey, Philonix Fitch, Acraspis Mayr and Biorhiza Westwood (=Sphaeroteras Ashmead) but can easily be distinguished by the following character states: antenna with 16 antennomeres (Fig. 1E); projecting part of the ventral spine of the hypopygium is long, lanceolate, and at least 4.0 times as long as broad; from the point where the spine narrows, a dense group of long setae arises, which reach the apex of the spine, forming a tuft, ; lateral propodeal carinae poorly defined, fragmented, widely divergent posteriorly, with a median propodeal area bare, and narrower anteriorly, and a fragmented median longitudinal carina present (Fig.  2E); lateral propodeal area densely pubescent and forewing hyaline.
Of the genera included into the Cynips group, the new genus resemble the genus Atrusca, primarily because of the long ventral spine of the hypopygium, but it differs in the position, direction and length of the setae on the spine, which are subapical, reaching beyond the apex of the spine. Additionally, the species of genus Atrusca possesses, on the forewing, dark spots or dark stripes along veins, the radial cell is 2.0 -2.5 times as long as broad and the Rs is strongly angulate. More diagnostic characters are given in the generic key and the description below.
Description. Description of this genus is based in the asexual generation of the only known species. The eventual discovery of a sexual generation would imply the revision of the generic limits.
Lateral propodeal carinae moderately divergent ventrally, the median propodeal area narrow, bare, with a median longitudinal carina present (Fig. 2E). Metatarsal claws with an acute basal lobe. Forewing (Fig. 5B) hyaline, without fuscate spots or stripes, radial cell open along anterior margin; areolet triangular, closed and distinct. Apical margin of wing with short hair fringe. Metasoma with second metasomal tergite covering about two thirds of metasoma, with a patch of setae laterally in its anteromedial area. Projecting part of hypopygial spine long, lance shaped, at least 4.0 times as long as broad, with dense, long basal setae that reach the apex of the spine and form a dense tuft ( Fig. 3B-C).
Distribution. Based on our data, the new genus is found only to 2515-3045 m a.s.l. at Volcán Barú, Chiriqui, Panama, around the upper limit of the growth of Quercus species in Panama.   yellowish. Mandibles yellowish with black teeth. Antennal scape and pedicel yellowish in part. Half basal of coxae black; coxae apically, femora and tibiae ventrally yellowish. Forewing hyaline, slightly and uniformly darkened; veins dark brown.
Head alutaceous-reticulate, moderately pubescent, with relatively long white setae, except on vertex, upper frons and gena, in dorsal view about 2.7 times wider than long. POL 1.8 times longer than OOL, posterior ocellus separated from inner orbit of eye by 2.0 times its longest diameter. Head in anterior view (Fig. 1A) 1.2 times wider than high. Genae slightly expanded behind eyes, strongly alutaceus-reticulate (Fig. 1B). Clypeus more or less trapezoidal, 1.7 times wider than high moderately pubescent, ventral margin sinuate, moderately projecting over mandibles. Anterior tentorial pits visible; epistomal sulcus not visible, clypeo-pleurostomal lines visible. Malar space 0.2 times height of compound eye, without malar sulcus strongly alutaceus-reticulate, without irradiating striae from clypeus (Fig. 1B). Toruli situated slightly above mid-height of compound eye; distance between antennal rim and compound eye 0.8 times width of antennal socket including rim. Ocellar plate slightly raised. Head, posterior view ( Fig. 1C) without occipital carina. Gula short; distance between occipital and oral foramina as high as occipital foramen (Fig. 1C). Hypostomal sulci well separate at oral fossa.
Forewing (Fig. 5B) 1.2 times as long as body, without smoky spots or stripes and densely pubescent; basal cell pubescent; radial cell 4.2 times longer than wide, open along anterior margin; areolet large, triangular, closed. R1, Rs and M nearly straight not reaching wing margin. R1 forming a quite acute angle with anterior margin of wing; Rs+M not reaching basalis. Basalis slightly curved, 2r well pigmented. Apical margin of wing with moderately long hair fringe.
Metasoma (Fig. 3A) large, as long as head and mesosoma combined, in lateral view as wide as high. Second metasomal tergite covering about two thirds of metasoma, with a patch of setae in its lateral anteromedial area. Projecting part of hypopygial spine long, lanceolate, tapering from the base to apex, at least 4.0 times as long as broad, with dense long basal setae arising in the base of the projected part which reach the apex of the spine forming a tuft (Fig. 3B-C).
Gall (Fig. 5C-G). Irregular small formations (up to 4-9 mm) arising from crevices on the stems and on the petiole and midribs of leaves. The gall is displayed as a dense mass covered with light brown hairs, solitary, containing a single larval cell or more frequently forming clusters, and then appearing as polythalamous. Inside, the gall has a highly lignified core enclosing the larva (Fig. 5E).
Host plant. Quercus bumelioides Liebm. (section Quercus of Quercus; white oaks (Fagaceae), a species distributed from Mexico to Panama (Breedlove 2001). The gall resembles that of the species of the Nubila complex, established by Kinsey within the subgenus Acraspis of Cynips (Kinsey 1936), all known from Mexico. These galls were described as a mass of coarse hairs containing a spherical hard core, attached to mid-veins, on or under the surfaces of leaves. However, the insects are quite different in important characteristics such as the number of antennal segments and the shape of the hypopygial spine.
Distribution. Barucynips panamensis was found between 2515-3045 m a.s.l. at Volcán Barú, Chiriqui, Panama. Although currently known only at this locality, it is a species that is relatively abundant at the higher elevations of Volcán Barú.
Biology. Only the asexual generation is known, inducing galls on Quercus bumelioides Liebm (section Quercus). The galls are common and can be found at every time of year in different grades of maturation on stems and leaves. The galls frequently are found growing together with galls of the new species, also described in this paper, Coffeikokkos korytkowskii. The insects studied emerged from January to July.
Inquiline and parasitoid associated community. Diagnosis and comments. The new species represents the second species of the genus Coffeikokkos, which was recently described in Costa Rica. The species is closely related to C. copeyensis Pujade-Villar & Melika, 2012, being similar in color and a majority of morphological characteristics. The species differ in the pubescence of mesosoma, the shape of the mesoscutellum, propodeal sculpture and gall shape. C. korytkowskii has a moderately pubescent mesosoma with piliferous punctures, whereas the mesosoma is smooth in C. copeyensis. The new species has a more elongate mesoscutellum, clearly longer than is wide, and the propodeal carinae are complete, reaching the nucha, whereas in C. copeyensis, the mesoscutellum is as long as is wide or only slightly longer than wide, and the propodeal carinae is incomplete, not reaching the nucha. Additionally, the basal cell of the forewing in the new species is hairy, whereas the basal cell of C. copeyensis is bare. The new species induces a regular spherical gall (5 mm diameter) with a spotty surface, while the galls induced by C. copeyensis are similar in size, but irregular or slightly ovate and uniformly colored.
Description. Body length 3.62 mm (range 3.3-4.2; N = 5) for females. Body uniformly reddish brown and shiny; the toruli area, flagellomeres of antenna, area above clypeus, occiput, dorsolateral margin of pronotum, anteroadmedian signa area, parapsidal signa, mesopleuron, metapectal-propodeal complex and anteromedial area of scutellum dark brown to black. Legs with all coxae and femora yellowish; tibia and tarsomeres dark brown to black. Forewing hyaline with some very light infumation; veins dark brown to black.
Asexual female. Head moderately pubescent with piliferous punctures, in dorsal view about 3.5 times wider than long. POL 1.5 as long as OOL, posterior ocel-lus separated from inner orbit of eye by 1.7 times its longest diameter. Head in anterior view (Fig. 6A) transversely ovate, 1.15 times wider than high, gena not expanded behind eyes. Vertex frons and gena slightly alutaceous. Head moderately pubescent, with relatively long white setae, except vertex, frons with sparse, shorter setae. Clypeus more or less trapezoidal, 1.6 times wider than high, mostly smooth and moderately pubescent; ventral margin sinuate, slightly projecting over mandibles. Anterior tentorial pits visible; epistomal sulcus indicated, clypeo-pleurostomal lines visible. Malar space 0.3 times height of compound eye, without malar sulcus; some irradiating striae from clypeus present, reaching ventral margin of compound eye, absent medially above clypeus. Toruli situated slightly above mid-height of compound eye; distance between antennal rim and compound eye 1.1 times width of antennal socket including rim. Ocellar plate not raised. Head, posterior view (Fig. 6B) without occipital carina. Gula short; distance between occipital and oral  foramina 0.5 times height of occipital foramen (Fig. 6B). Hypostomal sulci well separate at oral fossa.
Legs. Densely pubescent; metatarsal claws with a large obtuse basal lobe (Fig. 7D). Forewing (Fig. 8B) slightly longer than body, strongly pubescent; basal cell with some rows of setae; radial cell 4.0 times longer than wide; open along anterior margin; areolet triangular, closed and distinct. R1 and Rs nearly straight, not quite reaching wing margin; R1 forming an acute angle with anterior margin of wing. Rs+M reaching basalis at its mid-height. 2r well pigmented, angulate and slightly projected medially. Apical margin of wing with moderately long hair fringe. Metasoma (Fig. 7E) large, as long as head and mesosoma combined, in lateral view as wide as high. Second metasomal tergite covering about 2/3 of metasoma, with a patch of dense setae in its anteromedial area. Projecting part of hypopygial spine short (Fig. 7F-G), shorter than basal height of spine (Fig. 7F); with parallel sides and pointed apically, with dense long subapical setae forming a patch, extending far beyond apex of spine.
Gall (Fig. 8D-H). Similar in location, shape, and size to the galls of C. copeyensis Pujade-Villar & Melika. However, the galls of this new species are much more regu-larly spherical and its surface is not uniformly colored, but spotty. Diameter of gall measures 5 to 8 mm. They are formed, solitary or more frequently in groups, in stems of Quercus bumelioides Liebm. The surface of the gall is smooth and shiny; whitish, green or yellowish when fresh with red spots, becoming brown when mature. Monothalamic, with compact woody tissue internally containing the single larval cell (Fig.  8H). Similar spherical and spotty galls are also induced by the Nearctic Cynips (=Besbicus) mirabilis (Kinsey 1922), but this galls are larger, pubescent, formed in leaves and with an internal structure of irradiant filaments (Kinsey 1930).
Biology. Only the asexual generation is known, inducing galls on Quercus bumelioides Liebm. (section Quercus). The galls are common and can be found at every time of year in different grades of maturation on stems. They often develop jointly with the galls of Barucynips panamensis, also described here. When the gall is mature, it falls to the ground (Fig. 8G), from which the adult emerges after a couple of months. The studied insects emerged in November.

Discussion
The majority of the morphological characters of Coffeikokkos and the new proposed genus Barucynips indicate their close affinity with the "Cynips group" of genera (Kinsey 1930, 1936, Liljeblad et al. 2008. Their association only with Quercus bumelioides, a species of "white oaks" (Quercus section Quercus), reinforce the morphological evidence. The genera of the Cynips group, which is represented primarily in the Nearctic Region, have had an unstable taxonomic status. Kinsey (1930Kinsey ( , 1936, in his revision of the genus Cynips, divided it into six subgenera: Cynips (European species) Antron, Besbicus, Atrusca, Philonix, and Acraspis (American species). Under the "mellea" species complex in the Acraspis subgenus, he included also all known Sphaeroteras species (Melika and Abrahamson 2002). Weld (1952) gave the status of genus to all subgenera of Kinsey, but Melika and Abrahamson (2002) consider Antron and Besbicus synonyms of Cynips, and Sphaeroteras a synonym of Biorhiza. Liljeblad et al. (2008) in one analyses of 308 characters, 283 from morphology and 25 from biology and distribution, suggested that these last synonymyzations were unfortunate, as they showed the close and unresolved phylogenetic relationships between the taxa forming the Cynips group, including the Trigonaspis, Belocnonema and Biorhiza species. However, because of the relatively poor taxon sampling in their analysis and the lack of convincing support values for the clades within the Cynips group, they refrained from proposing formal taxonomic changes in Cynips group. Pujade-Villar et al. (2010) described one new genus from Mexico (Kinseyella) related to Cynips group and suggested that the neartic genera Antron and Besbicus were erroneously synonymized to Cynips by Melika and Abrahamson (2002). Despite the unstable taxonomic status of this group of genera and the clear need for a complete revision, mainly in the Neartic fauna, the new genus proposed herein, as with the recently described Coffeikokkos, present clearly distinctive morphological characters that justify its establishment. Both genera are closely related and share a unique synapomorphy within the Cynipini, namely, the antenna with 14-15 flagellomeres. However, Barucynips is well distinguished from Coffeikokkos and from all known Cynipini genera by one distinctive morphological character, the shape and pubescence of the projecting part of the ventral spine of the hypopygium. Furthermore, a combination of other morphological character, such as the presence of a median propodeal carina and a weakly toothed metatarsal claw lend further support to the status of this proposed new genus. In Panama, we have found more species clearly included in the Cynips group of taxa but that are doubtfully assigned to any described genus. Future studies should clarify whether these species should be attributed to the genus Cynips (sensu lato) or to one or more new genera. These facts clearly show the taxonomic complexity of this fauna and note the necessity of more complete revisional studies.
The southern boundaries of distribution of the oak gall wasps in America (the Cynipidae associated with Fagaceae) are revealing an unexpectedly great taxonomic richness and phylogenetic diversity. As evidenced by gall diversity in Panama alone, we have demonstrated that a rich fauna of 45-65 species of Cynipidae do exist in this country, contrasting with the single species previously recorded (Nieves-Aldrey and Medianero 2011, Medianero and. Furthermore, many genera previously known only in the Nearctic region, such as Amphibolips, Disholcaspis, Odontocynips, Bassetia, and Loxaulus, have been found in recent years in the Neotropical region from Costa Rica to Colombia (Medianero and Nieves-Aldrey 2010a, 2010b, while representatives of undescribed species related to Neuroterus, Dryocosmus, Cynips, Trigonaspis and Callirhytis have also been found but have been not yet published (Medianero and Nieves-Aldrey unpub.). More interesting is the discovery of new genera such as Agastoroxenia (Nieves-Aldrey and Medianero 2010) from Panama, Coffeikokkos (Pujade-Villar et al. 2012a) and Zapatella (Pujade et al. 2012b) from Costa Rica and Colombia, and the new genus Barucynips from Panama described here, demonstrating that the phylogenetic diversity is high in this region. All of these data support Kinsey's (1936) hypothesis, which postulated America as the center of origin and radiation of the Cynipini, and challenges the more recent hypothesis that postulates that gall wasp lineages diverged in Asia (Stone et al. 2008). New data collected in the Eastern Palearctic and oriental biogeographic region, as well as ongoing phylogenetic and biogeographical studies, will shed more light on this problem. Current data on the richness and diversity of oak gall wasps in the the Neotropic clearly indicate that this fauna reflects the result of the population dynamics along Pleistocene glaciations (circa 2.0 million to 12,000 years before present), with the high peaks of Neotropical mountains playing the role of postglacial refugees for the gall wasp fauna and their host plants.