Neotrop Entomol (2013) 42:63–71 DOI 10.1007/s13744-012-0095-z
SYSTEMATICS, MORPHOLOGY AND PHYSIOLOGY
Description of a New Genus and Three New Species of Figitinae (Hymenoptera: Cynipoidea: Figitidae) from Colombia J PUJADE-VILLAR, R PETERSEN-SILVA, J PARETAS-MARTÍNEZ Fac de Biologia, Dept de Biologia Animal, Univ de Barcelona, Barcelona, Spain Fac de Biologia, Dept de Biologia Animal, Univ de Barcelona, Barcelona, Spain
Keywords Ferpereira, Figites, Figitinae, Neotropical, Neralsia Correspondence J Pujade-Villar, Fac de Biologia, Dept de Biologia Animal, Univ de Barcelona, Avda Diagonal, 643, 08028 Barcelona, Spain;
[email protected] Edited by Takumasa Kondo – CORPOICA
Abstract A new genus and three new species of Figitinae (Hymenoptera: Figitidae) are described from Colombia: Ferpereira Pujade-Villar n. gen., Ferpereira fiorellae Pujade-Villar n. sp., Neralsia levis Pujade-Villar & Petersen-Silva n. sp., and Figites colombiensis Pujade-Villar & ParetasMartínez n. sp., which is the first species of Figites Latreille from Colombia and the second from South America. The diagnostic characters of these new taxa are illustrated.
Received 8 May 2012 and accepted 2 November 2012 Published online 23 November 2012 * Sociedade Entomológica do Brasil 2012
Introduction The Figitinae is a cosmopolitan subfamily of Figitidae (Hymenoptera: Cynipoidea) and, when known, they are parasitoids of larvae of cyclorrhaphan Diptera (Ronquist 1999).The Figitinae are circumscribed by the lack of derived characters present in other figitid subfamilies, and Ronquist (1999) described the group as “an obvious classificatory wastebasket.” The analysis of the phylogeny of the Figitidae by Buffington et al (2007) retrieved a monophyletic Figitinae excluding two genera: Lonchidia Thomson, which rendered the Figitinae paraphyletic in the parsimony analysis but was not formally excluded from the subfamily, and Melanips Haliday, which was placed in the Aspicerinae sensu lato. Paretas-Martinez et al (2012) regarded this placement of Melanips as controversial and included this genus in their Figitinae key “for the purpose of completeness.” According to Paretas-Martinez et al (2012), the Figitinae includes 14 valid genera: Amphithectus Hartig, Figites Latreille, Foersterhomorus Pujade-Villar & Petersen-Silva (0 Homorus Förster), Lonchidia Thomson, Nebulovena Pujade-Villar & Paretas-Martinez, Neralsia Cameron,
Paraschiza Weld, Sarothrioides Belizin, Sarothrus Hartig, Seitneria Tavares, Trischiza Förster, Xyalophora Kieffer, Xyalophoroides Jiménez & Pujade-Villar, and Zygosis Förster. The genera with scutellar spine (Neralsia, Xyalophora, and Xyalophoroides) have been revised in depth the last years (Jiménez et al 2004, 2005, 2006, 2008a, b, c, Jiménez & Pujade-Villar 2008, 2009, Petersen-Silva & PujadeVillar 2010, Petersen-Silva et al 2010, Pujade-Villar et al 2008, Pujade-Villar & Caballero-López 2010). Other revised genera include the monospecific genera Foersterhomorus (Pujade-Villar et al 2011), Nebulovena (Paretas-Martinez et al 2012), and Paraschiza (Petersen-Silva et al 2011). The other Figitinae genera have not been treated since Weld (1952) and need to be revised. In this work, we describe a new genus and three new species of Figitinae from Colombia in the Neotropical area; previously, only one genus of Figitinae (Neralsia) had been cited from Colombia, with eight species (Jiménez et al 2008a). In the last years, we have focused on the Figitinae in order to solve the taxonomic chaos of this poorly known, but important group of dipteran parasitoids. Figitinae systematics will help not only to resolve their internal relationships but
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also to establish their affinities with other figitid subfamilies, especially the Aspicerinae.
Material and Methods The Colombian specimens were collected by Malaise trap. The material has been deposited in the “Instituto Alexander von Humboldt” (IAvH; Villa de Leiva, Colombia) and in the Pujade-Villar collection (Universitat de Barcelona (UB), Barcelona, Spain). Scanning electron micrographs (ESEM) were obtained without any coating with the FEI Quanta 200 Environmental SEM at 15 kV or with the Leica Stereoscan S-360 SEM (Cambridge Instruments) at 700–850 V. Radial cells and images in Fig 2 were photographed directly in the stereomicroscope with a digital camera Canon PowerShot SX210 15. Some wing pictures were taken with a digital camera Infinityx-21C built in a compound stereomicroscope Zeiss Discovery V8. The images will be available online from the databank www.morphbank.net. The terminology for morphological structures follows Richards (1977), Ronquist & Nordlander (1989), Ronquist (1995), Ros-Farré et al (2000), and Ros-Farré & PujadeVillar (2007), and the sculpture terminology follows Harris (1979). Measurements and abbreviations in the descriptions include: F1–F12, first and following flagellomeres; T3–T4, third and fourth abdominal tergites; antennal formula is given with the length/width ratio of each segment.
Results Ferpereira Pujade-Villar n. gen. (Figs 1–2) Type species: Ferpereira fiorellae Pujade-Villar n. sp., by present designation and monotypy. Diagnosis Ferpereira resembles Nebulovena having subtriangular head and lacking areolet on the forewing (other figitines have a rounded or sub-quadrangular head and the areolet is present, although partly defined by spectral or nebulous veins). Furthermore, Ferpereira has a scutellar morphology different to other Figitinae: Ferpereira can be distinguished from Neralsia, Xyalophora, and Xyalophoroides by lacking scutellar spine; from Figites, Foersterhomorus, Paraschiza, Sarothrioides, Sarothrus, Trischiza, and Zygosis by having sculptured mesoscutum (smooth in these genera); from Melanips and Seitneria by having circumscutellar carina; from Amphithectus having metasoma as long as mesosoma
(metasoma longer than mesosoma + head in Amphithectus); and from Nebulovena having veins of radial cell hyaline (nebulous in Nebulovena), having very few piliferous points, mesopleuron glabrous below longitudinal carinae, and propodeal carinae poorly defined, curved, and wide. Ferpereira also resembles Melanips, but the presence of the circumscutellar carinae, among other characteristics mentioned before, distinguishes Ferpereira from Melanips. Description See description of type species. Etymology. The generic name Ferpereira is dedicated to my teacher, friend, and colleague, Ferran Pereira López, who unfortunately after be fighting a rare cancer, he died a few months ago. Ferpereira fiorellae Pujade-Villar n. sp. (Figs 1–2)
Description Length. Males: 2.2 mm; females: 2.8 mm. Color. Completely black except antennae and legs dark brown, coxae darker brown; wing venation brown. Head. Subtriangular shaped in anterior view, 1.2 wider than its length; face with micro-sculpture. Center of face less sculptured. Frons shiny, alutaceous with scattered but visible piliferous points; space around compound eyes smooth. Malar space 0.5 times the length of compound eye; malar sulcus absent but defined by a wide stripe with coriaceous sculpture. Clypeus smooth, shiny, well differentiated, and projected above mandibles. The clypeopleurostomal lines well developed. Space between toruli equals toruli diameter, with a marked depression densely pubescent. Distance between toruli and compound eye half the diameter of toruli. Transfacial line equal to eye length. Head in dorsal view 2.2 times wider than its length. Relation POL/OOL/OCO is 9:4:4, maxim diameter of lateral ocelli 3.5. Occiput coriaceous; occipital carinae absent. Genal carinae present. Antenna. Female 13-segmented; antennal formula: 9(4): 5 (4): 14(3): 8(3): 8(3): 8(3): 6(3): 6(3): 6(3.5): 6(4): 5(4): 5(4): 9(4); F1 basally and distally slightly wider; sensillia in all flagellomeres. Male 14-segmenetd; antennal formula: 5(4): 4(3.5): 13(4): 7(2.5): 6.5(2.5): 6(3): 5(3): 5.5(3): 5(3): 5(3): 5.5(3): 4.5(3): 4(3): 7(3); F1 very long, distinctly curved, dorsally flattened, and excavate.
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Fig 1 Ferpereira fiorellae n. sp.: a male, head, and mesosoma in lateral view with detail of pedicel-F2 and radial cell; b female antenna and detail of F1– F2; c head and pronotum in dorsal view; d metasoma in dorsal view.
Mesosoma. Pronotum, mesoscutum, scutellum, mesopleural triangle, metapleuron, and propodeum covered with sparse/dense setae. Pronotum smooth but with some piliferous points, pubescent; pronotal carinae not meeting dorsally, not reaching mesoscutum, not forming an upraised pronotal plate; lateral surface of pronotum with alutaceous sculpture. Mesoscutum shiny but with coriaceous micro-sculpture. Notauli percurrent, wider on posterior part, not well defined on anterior part; medial mesoscutal sulcus superficially marked, lacking any sculpture, reaching tegula level. Antero-admedian lines weak, reaching anterior one third of mesoscutum. Parascutal carina smooth, slightly overlapping tegulae. Mesopleuron smooth with visible, deep median sulcus on lower third; anterior part with some carinae. Scutellum with visible coriaceous hump; circumscutellar carina projected on posterior margin; scutellar foveae superficial, almost confluent, separated by a weak carina more defined in females; lateral pits of foveae deep. Propodeal area pubescent, propodeal carinae sub-parallel, weakly arched, visibly sculptured.
Legs. Internal metatibial spine short, reaching 1/3 length of first tarsomere. Tarsal claws simple. Wings. Pubescent and ciliate; radial cell closed, 2.5 times longer than its width in females, 2.25 longer than wide in males. R2 and Rs almost straight, M and Rs+M veins very thin but visible; areolet absent. Metasoma. Petiole twice as long as wide in males, only little longer than its width in females; petiole dorsally smooth, with some scarcely marked longitudinal carinae laterally; metasomal T2 segment smooth, with two patches of pubescence dorsally; T3 and following without pubescence; postero-dorsal part of the third segment punctuated, weakly in females. T3 longer than T2 dorsally. Ventral spine and hypopygium not visible. Type material: 3♂ and 1♀. HOLOTYPE (♂) with the following labels: “COLOMBIA, Boyacá, SFF Iguaque, Cabaña Mamarramos, 5°25′ N 73°27′ W, 2,855 m, Malaise, 21.xii.2000, 07.i.2001, P. Reina, Leg. M.1072 (white label), “Holotype Ferpereira fiorellae n. gen. & n. sp. desig. JP-V
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Fig 2 Ferpereira fiorellae n. sp.: a head in frontal view; b mesosoma in dorsal view; c lateral view of scutellum; d scutellum in dorsal view; e mesosoma in lateral view; f propodeum.
2011” (red label); “Ferpereira fiorellae ♂ det. JP-V 2011” (white label). PARATYPES. 1♂ & 1♀: “COLOMBIA, Boyacá, SFF Iguaque, Lagunillas, 5°25′ N 73°27′ W, 3,380 m, Malaise, 28.iii.16.iv.2001, P. Reina, Leg. M.1515” (white label), “Paratype Ferpereira fiorellae n. gen. & n. sp.” (red label), “Ferpereira fiorellae n. gen. & n. sp. det. JP-V 2011” (white label); 1♂: “Idem, 16.iv.02.ii.2001, P. Reina, Leg. M.1738” (white label), “Paratype Ferpereira fiorellae n. gen. & n. sp.” (red label); Ferpereira fiorellae n. gen. & n. sp. det. JP-V 2011” (white label). Holotype and two paratypes are deposited in IAvH, one paratype (male-1515) in UB.
Figites colombiensis Pujade-Villar & Paretas-Martínez n. sp. (Fig 3b–e)
Diagnosis
Distribution. Only known from Colombia.
Figites colombiensis n. sp. (Fig 3b–e) morphologically resembles Figites braziliensis Borgmeier (Fig 3a), the only other species of Figites described in South America. Figites colombiensis is distinguished from F. braziliensis having: central area of scutellum smooth (completely rugose in F. braziliensis); oval scutellar foveae (more or less triangular in F. braziliensis); bar separating scutellar foveae completely straight (triangular in F. braziliensis); and discontinuous sculpture on occiput (with a continuous occipital carina in F. braziliensis).
Biology. Unknown. Host unknown.
Length. Females: 2.7 mm; males: unknown.
Etymology. The specific name fiorellae is dedicated to Fiorella Ruiz Pace, Ferran Pereira’s wife.
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Fig 3 a Dorsal view of Figites braziliensis; b–e Figites colombiensis n. sp.: b head, antenna, and mesosoma in dorsal view; c radial cell of forewing; d mesosoma; e metasoma.
Color. Mesosoma, metasoma, and head black; antennae and mandibles brown; legs, amber yellow; proximal and distal areas of metasoma lighter. Head. Scarce bristled pubescence in compound eyes; round shaped in frontal view, in dorsal view 1.6 times longer than its width. Transfacial line longer than eye length (13:11); relation POL/OOL/LOL is 7:3:4, ocellus diameter 2; face with alutaceous sculpture, with two shiny, almost smooth areas; genal sulcus defined, with transversal costulae, visible between coriaceous sculpture; malar sulcus absent; malar space 0.65 times as long as compound eye, with coriaceous sculpture extending to surround all compound eye; occipital carinae present but incomplete dorsally, in dorsal view looking wrinkled not well-defined sculpture. Frons alutaceous, shiny. Antenna. Female antennal formula: 6(4): 3(3): 6(3): 4(3): 4 (3.5): 4.5(3.5): 5(4): 4.5(4): 4(4): 3.5(3.5): 4(3.5): 3.5(3.5): 7 (4); sensilla present but rare in F3, more abundant on F4–F11. Mesosoma. Smooth, shiny, with few long setae except mesopleuron; pronotal carinae meeting dorsally, forming
an upraised pronotal plate dorsally incised; lateral areas of pronotum smooth, with some few, short carinae on dorsal part, next to pronotal plate, with some carinae on ventral part; mesopleural area finely carinate; mesoscutum medial sulcus clearly visible, 1/4 of total length of mesoscutum; notauli complete, percurrent, wide on posterior part; parapsidal sulcus going beyond tegulae; scutellar foveae deep, interior smooth, oval shaped, separated by a carina at same level than foveae; scutellar plate irregularly wrinkled, not well defined on anterior part; circumscutellar carina clearly defined. Wings. Hyaline, light brown venation; closed radial cell, 1.9 times longer than width; setae on margin and disk. Areolet nebulous. Metasoma. Metasomal T2 lacking dense patches of setae, with only few sparse setae; with few, short strigae on basal area, distal part smooth. Following tergites punctuate. Type material: 1♀. HOLOTYPE (♀) with the following labels: “COLOMBIA, Huila, PNN Cueva de Los Guácharos, Cabaña Cedros, 1°37′N 76°6′W, 1,950 m, Malaise, 20.v.5-
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Fig 4 Neralsia laevis n. sp.: a head of female in frontal view; b head of male in frontal view; c female antenna; d male antenna; e head and mesosoma in dorsal view; f pronotal plate; g scutellum in dorsal view; h metasoma in lateral view; i body in lateral view.
vi.2003, C. Cortés Leg. M.3735” (white label), “Holotype Figites colombiensis n. sp. desig. JP-V 2011” (red label); “Figites colombiensis ♀ det. JP-V 2011” (white label). Holotype deposited in IAvH. Etymology. Specific name refers to the country where this species was found. Distribution. Only known from Colombia. Biology. Unknown. Host unknown. Neralsia laevis Pujade-Villar & Petersen-Silva n. sp. (Fig 4)
Diagnosis Neralsia laevis n. sp. belongs to the group of Neralsia species with low interfoveal carinae, not higher than level of scutellar foveae (Fig 4h), and with metasomal T2 smooth. Morphologically resembles Neralsia albipennis (Kieffer), Neralsia gibbosa Jiménez & Pujade-Villar, and Neralsia pseudoneralsia Jiménez & Pujade-Villar, by having the scutellar plate without longitudinal carinae (Fig 4e). Neralsia laevis is distinguished from N. albipennis and N. gibbosa by lacking gibbous scutellum, and from N. pseudoneralsia by having longer scutellar spine, 1/10 of scutellum length in N. pseudoneralsia and 1/4 in N. laevis. Length. Females: 3.1 mm; males: 3.5 mm.
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Color. Black, except antenae dark brown; scape, pedicel and F1 darker brown; legs light brown; coxae and femur black.
V 2011” (white label). Holotype and two paratypes deposited in IAvH, two paratypes (male and female) in UB.
Head. Shiny with scarce pubescence; transfacial line shorter than eye size; round head in frontal view, as long as its width, in dorsal view two times longer than its width; relation POL/ OOL/LOL is 5:2:4, ocellus diameter 3; face strigae from clypeus to compound eyes; genal sulcus defined, with extremely weak and not well-defined transversal costulae; occiput with discontinuous carinae ending behind ocelli.
Etymology. The specific name refers to the smooth and shiny scutellum of this species.
Antenna. Female antennal formula: 7(3): 3(2): 3.5(2): 3 (2.5): 4(2.75): 4(3): 4(3): 3.5(3): 3.5(3): 4(3.5): 4(3.5): 8 (3.5); sensilla starting on F3, weakly impressed. Male antennal formula: 5(3): 2(3): 6(3): 7(3): 9(3): 9(3): 9(3): 9(3): 8 (3): 8(3): 8(3): 8(3): 7(3): 10(3); sensilla on all segments.
Discussion
Mesosoma. Smooth, shiny, with scarce pubescence; mesopleural area without setae; pronotal carinae meeting dorsally, forming an upraised pronotal plate dorsally incised; lateral areas of pronotum with few weak carinae on dorsal area, with some carinae on the ventral part; mesopleural area mostly smooth only with some weak carinae on anterior part; mesoscutum medial sulcus short; notauli complete, percurrent; parapsidal sulcus going beyond tegulae; scutellar foveae separated by a carina at same level than foveae; scutellar foveae posteriorly not limited by a carina; scutellar plate smooth, without carinae; scutellar spine almost 1/4 of scutellum length; circumscutellar carina clearly defined; in males some short carinae close to circumscutellar carina. Wings. Smoked with dark brown venation in males, hyaline with light brown venation in females; open radial cell, two times longer than its width; setae on margin and disk; areolet slightly visible, almost absent; R1 weakly curved, Rs distally curved, both reaching wing margin. Metasoma. Metasomal T2 smooth, lacking setae; big metasomal tergite smooth in females, punctuate in males on posterior part; metasomal tergite VIII straight in females. Type material: 3♂ and 2♀. HOLOTYPE (♀) with the following labels: “COLOMBIA, Boyacá, SFF Iguaque, Cabaña Mamarramos, m1, 5°25′N 73°27′W, 2,855 m, Malaise, 08–25.vi.2000, P. Reina, Leg. M.184” (white label), “Holotype Neralsia laevis n. sp. desig. JP-V 2011” (red label); “Neralsia laevis ♀det. JP-V 2011” (white label). PARATYPES. 1♂ and 1♀ with the same collecting data of Holotype, “Paratype Neralsia laevis n. sp.” (red label), “Neralsia laevis n. sp. det. JP-V 2011” (white label); 1♂ and 1♀: “COLOMBIA, Boyacá, SFF Iguaque, Cabaña Mamarramos, m4, 5°25′N 73°27′W, 2,855 m, Malaise,19.iv.06.v.2000, P. Reina, Leg. M.55” (white label), “Paratype Neralsia laevis n. sp.” (red label), “Neralsia laevis n. sp. det. JP-
Distribution. Only known from Colombia. Biology. Unknown. Host unknown.
Figitidae biodiversity is very poorly known. Nordlander (1984) estimated for this family a global diversity of 24,000 species; currently, only 1,400 figitid species have been described (Buffington et al 2007), most of them from the Holarctic region (Dalla Torre & Kieffer 1910, Weld 1952). Thus, as it happens in many other biological groups, the lack of biodiversity knowledge in the Figitidae is especially critical in the tropical regions. Recent studies have shown that the microcynipoids (Cynipidae and Figitidae) may have a Gondwanan origin (Nieves-Aldrey et al 2009, Buffington et al 2012). If so, describing the figitid (and cynipid) fauna in the Southern Hemisphere is critical not only to increase our knowledge of the biodiversity of this family but also to understand the evolutionary history of this group of parasitoid wasps. In the Neotropical region, several authors have pointed out how the fauna of different figitid subfamilies remains practically undescribed (Fergusson & Hanson 1995, FontalCazalla & Nieves-Aldrey 1999, Paretas-Martínez & PujadeVillar 2007). In the last years, Neotropical Aspicerinae, Plectocynipinae, and Thrasorinae have been studied by Ros-Farré & Pujade-Villar (2006, 2007, 2010). With the exception of Xyalophora giraudi Jiménez & Pujade-Villar and F. braziliensis, our knowledge on the biodiversity of Neotropical Figitinae is almost restricted to the genus Neralsia, with 48 species described (Jiménez et al 2008a; Jiménez & Pujade-Villar 2008, 2009, Petersen-Silva & Pujade-Villar 2010, Pujade-Villar & Caballero-López 2010). The work we present here contributes to a better knowledge of the Neotropical figitine fauna, with the description of a new genus and three new species, including the second species of Figites in this region. Figites is a poorly circumscribed genus that includes more than 60 species; although a revision of Figites is needed to understand its intrageneric variation (and to tackle the systematics of the entire Figitinae), the combination of characters of F. colombiensis n. sp.—absence of scutellar spine, base of metasomal T2 striate and lacking dense patches of setae, bristled pubescence in compound eyes and mesosoma, pronotum and mesopleuron carinate—justifies
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the inclusion of this new species in this genus, following the most updated key to Figitinae (Paretas-Martinez et al 2012). The new genus described here, Ferpereira n. gen., mostly resembles Nebulovena having a subtriangular head and lacking areolet on the forewing, two characters not found in other figitine genera. Although Paretas-Martinez et al (2012) mention that the shape of the head of Nebulovena resemble morphologically the Anacharitinae, the head of anacharitines is perfectly triangular with a small mouth region and mandibles protruding and broadly overlapping (synapomorphy of the Anacharitinae), whereas the head of Nebulovena (we have been able to study more specimens of Nebulovena persa from Iran) and Ferpereira n. gen. is subtriangular, with a wider dorsal margin, a broadened mouth region and broader mandibles not overlapping. These two genera also lack the other synapomorphy of the Anacharitinae, a large upraised pronotal plate, having instead only two lateral carinae that do not meet medially neither form a complete upraised plate. Thus, following the most updated key to figitid subfamilies (Paretas-Martínez et al 2011), Ferpereira n. gen. must be included in the Figitinae. Ferpereira n. gen. and Nebulovena make the Figitinae an even more morphologically heterogeneous group that it has traditionally been. The erection of new higher taxa (tribes/subfamilies) has been suggested to accommodate figitine genera that are morphologically very different, like Lonchidia (Buffington et al 2007). However, further work is needed before any formal division of the Figitinae is proposed. A detailed revision of all genera included in the Figitinae (especially the species-rich Lonchidia and Figites) should be done to fully understand the morphological variation of this subfamily and its affinities with other genera/subfamilies. In this sense, a revision of Melanips, previously included in the Figitinae but now included in the Aspicerinae sensu lato (after Buffington et al 2007), is also needed because it lacks the morphological synapormorphies of the Aspicerinae sensu stricto (Ros-Farré et al 2000); the Aspicerinae sensu lato (Aspicerinae + Melanips) is supported genetically (Buffington et al 2007, 2012), but not morphologically (Buffington et al 2007). These revisions are needed to make a phylogenetic study of the Figitinae + Aspicerinae including all genera of both subfamilies in order to elucidate the placement of genera with very “particular” morphology like Lonchidia, Melanips, Nebulovena, and Ferpereira n. gen. Acknowledgments We are very grateful to Dr. Claudia Medina for the loan of the undetermined material from Colombia in which this study is based. We also want to thank Dr. Notton (NHM), Dr. Villemant (MNHN), Dr. Schmidt (ZSM), and Dr. Brandao (USP) for the loan of type material; Mar Ferrer-Suay (UB) for the optic pictures; and Palmira Ros-Farré (UB) for the SEM pictures that illustrate this study. Finally, we thank Dr. Zarazaga for his help and comments regarding the ICZN.
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