Fumariphilus, Nieves-Aldrey, 2022

Nieves-Aldrey, José Luis, 2022, Description of Fumariphilus Nieves-Aldrey, gen. nov., a new genus of herb gall wasps, with a key to genera of the tribe Aulacideini (Hymenoptera: Cynipidae), Zootaxa 5155 (3), pp. 393-413 : 396-400

Fumariphilus Nieves Aldrey, gen. nov.

Figs. 2–8

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Type species: Fumariphilus versicolor ( Nieves-Aldrey, 1985) , designated herein.

Etymology. The name is composed of two words: “ Fumari ” refers to the first part of the name of the host plant subfamily of the two cynipid species and “ philus ”, to the Greek for “love or fondness”. More generally, the name refers to the association of the new genus with plants of the genera Fumaria and Hypecoum in the subfamily Fumarioideae Eaton, 1836 (Papaveraceae) ( Angiosperm Phylogeny Group 2009).

Gender. Masculine

Diagnosis. Fumariphilus gen. nov., differs from the other Aulacideini genera by having the following combination of morphological characters: female antennae with 10 flagellomeres, notauli weakly impressed in the anterior third of the mesoscutum, vertex and mesoscutum weakly coriaceous with scattered piliferous punctures, radial cell open on the anterior margin of the forewing and a R1 vein that reaches the anterior margin. Also, they are gall inducers on Hypecoum and Fumaria ( Fumarioideae , Papaveraceae ).

The new genus morphologically most resembles species of the Isocolus / Aulacidea clade (see Fig. 1 ), however, species of Aulacidea differ from those of Fumariphilus by having antennae with 11 flagellomeres and a closed radial cell. Fumariphilus can be well differentiated from Isocolus by having notauli that are weakly impressed anteriorly, antenna with 10 flagellomeres and a R1 vein that reaches the anterior margin of the forewing. From Neaylax , Fumariphilus can be distinguished by the trapezoidal shape of its clypeus, with the clypeo-pleurostomal lines diverging towards the ventral margin (subparallel or slightly diverging in Neayla x); the longitudinally striated sculpture of its mesopleuron (striate-reticulate in Neaylax ) and the difference in host plant family association, Papaveraceae in Fumariphilus and Lamiaceae in Neaylax .

Fumariphilus gen. nov., is also morphologically similar to Aylax Hartig , and to the other genera of the tribe Aylacini . However, members of this tribe differ from Fumariphilus by having antennae with 12 flagellomeres in females (10 flagellomeres in Fumariphilus ); a pronotum that is short dorso-medially, less than 0.17X as long as the maximum length of the outer lateral margin, and with admedian depressions that are narrowly separated and strongly transverse (in Fumariphilus , the dorso-medial part of the pronotum is one-third as long as the maximum length of the outer lateral margin and the admedian depressions are distinct, spherical to oval and widely separated) and a mesopleuron sculpture that is striate-reticulate or reticulate (longitudinally striate in Fumariphilus ). In addition, Fumariphilus differs from the genera of the tribe Aylacini by its host plant genera association: Aylacini genera only associate with species of Papaver , whereas Fumariphilus associates with plants of the genera Fumaria and Hypecoum .

Description. Body slightly pubescent. Head in anterior view generally oval or more or less triangular ( Figs. 2A , 4C ), weakly alutaceous or coriaceous on front and vertex, with some piliferous punctures; genae not expanded behind eyes. Facial striae radiating from clypeus laterally well marked, reaching ventral margin of compound eyes, faint or absent medially ( Figs. 2A , 4C ). Anterior tentorial pits, epistomal sulcus and, clypeo-pleurostomal lines visible. Clypeus trapezoidal with ventral margin weakly or moderately projecting over mandibles. Malar (subocular) sulcus absent. Head from posterior view moderately pubescent, without occipital carina but with some transversal rugae or ridges ( Figs. 2B , 4B ). Distance between occipital and oral foramina shorter or as long as occipital foramen.

Antenna ( Figs. 2C , 4E ) with 10 flagellomeres in female, 13 in male; flagellum not broadening towards apex. F1 as long as F2.

Pronotum medially long (high), in anterior view ratio of median distance between anterior and posterior margins to lateral distance between these margins about 0.4 ( Fig. 3A ). Submedian pronotal depressions oval transverse, deep, open laterally. Pronotal plate present but inconspicuous; lateral margins of pronotal plate only visible ventrally; pronotal plate medially almost smooth and sparsely pilose. Lateral surface of pronotum without sculpture, more densely pubescent.

Mesoscutum weakly coriaceous, sometimes with weak piliferous puctures ( Figs 2D , 4D ). Notauli narrow and convergent posteriorly, clearly impressed in posterior one half of mesoscutum, weakly impressed to almost absent anteriorly. Median mesoscutal impression absent or visible in posterior one sixth of mesoscutum. Transscutal fissure present. Scutellar foveae from shallow, and smooth ( Fig. 4D ) to oval, with distinct margins and with some rugae ( Fig. 2D ). Meoscutellum with variable sculpture. Posterodorsal and posterior margins of axillula distinct. Mesopleuron entirely longitudinally costulate, the intervals smooth ( Fig. 2E ). Mesopleural triangle distinctly impressed, ventral margin clearly marked.

Metascutellum ( Fig. 3B ) conspicuously constricted medially. Metapleural sulcus meeting anterior margin of metapectal-propodeal complex slightly above mid-height of latter. Lateral propodeal carinae broad, subparallel ( Fig. 3B ). Lateral and median propodeal area smooth, laterally strongly pubescent. Nucha dorsally with some weak longitudinal rugae.

Metatarsal claws simple, without a basal lobe or tooth ( Fig. 3D ).

Forewing hyaline; radial cell open along anterior margin ( Figs. 5C, 5D ); R 1 ending slightly before or just reaching the anterior margin of wing. Areolet either present or absent. Hair fringe along apical present, moderately long.

Metasoma with third abdominal tergum (second gastral) covering less than one third of metasoma, with a conspicuous hair patch present antero-medially ( Fig. 3E ). Projecting part of hypopygial spine short, shorter than basal height of spine; the hypopygial spine bearing some long setae on each side, the apical ones surpassing the apex of the ventral spine. ( Figs. 3E, 3F ).

Terminal-instar larva ( Figs. 7 , 8 and 5E, 5F ): is of type hymenopteriform; body composed of head and 13 segments almost bare and without appendages. Maxillae triangular and with two pairs of maxillary palps and two pairs of maxillary setae present. Labium rhomboidal; salivary opening surrounded by tuberculate sculpture. Labial palps and labial setae present. Mandibles almost symmetrical with three teeth each.

Gall structures: Galls of Fumariphilus species are unilocular and develop in fruits ( Fig. 6 ).

Host plant association: Fumariphilus species associate with plants of the genera Hypecoum and Fumaria ( Fumarioideae , Papaveraceae ).

Biology: Life cycle is univoltine with bisexual reproduction. A single generation emerges in spring, after overwintering in galls that have fallen to the ground from the dried-out host plants.

Distribution: Circum-Mediterranean, extending east to Armenia, Iran and Central Asia ( Nieves-Aldrey 2003).

Key to species of Fumariphilus gen. nov.

1. Head rounded in anterior view; ventral margin of clypeus slightly projecting over mandibles ( Fig. 2A ). Notauli weekly impressed but visible in anterior one third of mesoscutum; median mesoscutal impression visible posteriorly ( Fig. 2D ). Scutellar foveae with some irregular rugae; mesoscutum with coriaceous, dull sculpture; mesoscutellum with rugose sculpture ( Fig. 2D ). Forewing with absent areolet; Rs vein reaching anterior margin of wing. Galls in fruits of species of Fumaria ........................................................................................ F. versicolor (Nieves-Aldrey)

- Head trapezoid in anterior view; ventral margin of clypeus distinctly projecting over mandibles ( Fig. 4C ). Notauli narrow and shallow, clearly visible only in posterior one third of mesoscutum, faint anteriorly; median mesoscutal impression absent ( Fig. 4D ). Scutellar foveae shallow, smooth and shining. Mesoscutum delicately shining coriaceous; mesoscutellum delicately alutaceous medially, with concentric rugae laterally ( Fig. 4D ). Forewing with a visible triangle shaped areolet; Rs vein not quite reaching anterior margin of wing ( Fig. 5C ). Galls in fruits of Hypecoum species. .................... F. hypecoi (Trotter)

Figures

FIGURE 1. Portion of the phylogeny related to herb gall wasps, modified from Fig. 3 in Ronquist et al. (2015). This analysis was based on a combined molecular data set of three genes Numbers on branches indicate posterior clade probabilities (in % units) ± Monte Carlo error (errors less than 1% not shown) across four separate analyses. Only groups with a posterior probability above 70% are shown. The red box indicates the monophyletic clade containing the two species with an incorrect genus identification, evidence that supports the erection of the new genus Fumariphilus to accommodate them. The red arrows point to the phylogenetic positions of the species correctly assigned to Aylax Hartig and Neaylax Nieves-Aldrey.

FIGURE 2. Fumariphilus versicolor, n. comb., SEM photos of a female. (A) Head, anterior view. (B) Head, posterior view. (C) Antenna. (D) Mesosoma, dorsal view. (E) Mesosoma, lateral view.

FIGURE 3. Fumariphilus versicolor, n. comb., SEM photos of a female. (A) Pronotum, antero-dorsal view. (B) Propodeum. (C) Mouth parts, posterior view. (D) Metatarsal claw. (E) Metasoma, lateral view. (F) Metasoma, ventral view.

FIGURE 4. Fumariphilus hypecoi, n. comb., SEM photos of a female. (A) Head, dorsal view. (B) Head, posterior view. (C) Head, anterior view. (E) Mesosoma, dorsal view. (E) Antenna.

FIGURE 5. Habitus, forewing and terminal instar larvae of Fumariphilus hypecoi, n. comb. (A), (C), (E) and Fumariphilus versicolor, n. comb. (B), (D) and (F). All in lateral view.

FIGURE 6. Galls of Fumariphilus versicolor, n. comb on Fumaria officinalis (A, B). (C) Section of a galled fruit showing a larva. (D) Section of a mature fruit with a fully developed adult inside. (E) Galls of Fumariphilus hypecoi, n. comb. on Hypecoum procumbens. (F) Section of a mature fruit with a fully developed adult inside.

FIGURE 7. Fumariphilus versicolor, n. comb., SEM photos of a terminal instar larva. (A) Body, ventral view. (B) Body, lateral view. (C) Head, anterior view. (D) Detail of mouth parts. (E) Left mandible. (F) Right mandible.

FIGURE 8. Fumariphilus hypecoi, n. comb., SEM photos of a terminal instar larva. (A) Body, ventral view. (B) Body, lateral view. (C) Head, anterior view. (D) Detail of mouth parts. (E) Right mandible. (F) Left mandible.