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Research Article
Revision of the neotropical genus Sendaphne Nixon (Hymenoptera, Braconidae, Microgastrinae)
expand article infoJose L. Fernandez-Triana, James B. Whitfield, M. Alex Smith§, Winnie Hallwachs|, Daniel H. Janzen|
† Canadian National Collection of Insects, Ottawa, Canada and University of Guelph, Guelph, Canada
‡ University of Illinois, Urbana, United States of America
§ University of Guelph, Guelph, Canada
| University of Pennsylvania, Philadelphia, United States of America
Open Access

Abstract

The Neotropical genus of parasitoid wasps Sendaphne (Hymenoptera, Braconidae, Microgastrinae) is revised and the following six new species are described, all authored by Fernández-Triana and Whitfield: anitae, bennetti, broadi, dianariaspennae, penteadodiasae, and rogerblancoi. The greatest species richness is found in northern South America, but the genus extends north to 23° N in Mexico. Most species have been collected in rainforest below altitudes of 900 m, with only a few species found in cloud forests up to 1900 m. Nothing is known of the host caterpillars for these parasitoid wasps.

Keywords

Sendaphne, Microgastrinae, Neotropics, Area de Conservación Guanacaste, taxonomic revision, parasitoid wasps, DNA barcoding

Introduction

The genus Sendaphne was described by Nixon (1965) to accommodate two Neotropical species with a number of distinctive features: a very long and bilobate glossa, slender body, extensive yellow coloration, mostly smooth mesosoma and metasoma, enlarged hypopygium, and very long and curved ovipositor. Penteado-Dias (1995) described two additional species, and Scatolini and Penteado-Dias (1999) added another one. Four of the five species described so far are from Brazil, with the exception being Sendaphne sulmo Nixon (1965), recorded from southeastern Mexico. No host is known for Sendaphne, although the related genus Promicrogaster Brues & Richardson, 1913, has been reared from hosts living in bracket fungi (Mason 1981), among other concealed hosts.

In spite of the rather unique morphological traits, the validity of Sendaphne as a distinct genus has been questioned even by the same author who described it (Nixon 1965: 204), and it might eventually be determined to be a synonym of the more abundant and diverse Promicrogaster. However, Mason (1981) considered the differences as sufficient to maintain them as two genera. That decision has been followed by all subsequent authors and, without a comprehensive phylogenetic study of Microgastrinae, we agree that is better to keep it that way at present. The morphological and molecular analysis of Whitfield et al. (2002), although not conclusive, also found those two genera to be closely related. An anticipated revision of Promicrogaster may help to elucidate the limits of these two genera in the future.

As part of comprehensive studies on the fauna of Microgastrinae from Area de Conservación Guanacaste (ACG), northwestern Costa Rica (e.g., Fernández-Triana et al. 2014 and references cited there) we found a new species from that area. The Canadian National Collection of Insects (CNC) in Ottawa, Canada and the NSF-funded “Insect Survey of a Hyperdiverse Country: Colombia” projects led by M. J. Sharkey, B. V. Brown and the Humboldt Institute (Colombia) also contained additional undescribed species from Central and South America. All these new species are described below, altogether with a key to all known, previously described species of Sendaphne.

Methods

Among Microgastrinae, Sendaphne is one of the most rarely collected genera, and is poorly represented in collections. This study is based on 73 specimens from four sources: 38 Neotropical specimens deposited in the CNC, 18 specimens from Colombia (Humboldt Institute), 10 specimens from the ACG inventory, four specimens from Unité d’Entomologie fonctionnelle et évolutive, Gembloux Agro-Bio Tech, Université de Liège (FUSAGx) and three specimens from the Natural History Museum, London, England (BMNH).

All species previously described were deposited in the Universidade Federal de São Carlos, São Carlos, São Paulo, Brazil (DCBU), Universidade Federal do Paraná, Curitiba, Paraná, Brazil (UFPR), or in the BMNH. We did not examine the holotypes of those species; however, their original descriptions and illustrations are sufficiently detailed to allow us to describe the new species with confidence.

Morphological terms and measurements of structures are mostly as used by Mason (1981), Huber and Sharkey (1993), Whitfield (1997), Karlsson and Ronquist (2012), and Fernández-Triana et al. (2014). Because the ovipositor in Sendaphne is strongly curved, its length was difficult to measure accurately; the ovipositor length measurements provided for each new species are only intended as an approximation. In any case, the ovipositor and its sheaths are some of the longest observed in any Microgastrinae genera; they are usually two times longer than the metatibia length.

Descriptions of the new species are based on the study of all available female specimens, so as to reflect intraspecific variation, but always include data from the holotype. As an exception, two new species only known from males are described below because they were sufficiently distinct to be distinguished from all others; the males of those two species may be identified by the key, but males of most other species may not be readily identified unless associated with females via rearing or molecular data.

The descriptions include 17 characters that are commonly used in describing Microgastrinae (e.g., body measurements such as length of body and fore wing, ovipositor sheath; and also color of particular body areas). Those descriptions are complemented with extensive color photos of every species. Geographic distribution is also provided in the key as supplementary information to aid the morphological identification of species, though we recognize that with time the current known geographic distribution may eventually become obsolete.

Photos were taken with a Keyence VHX-1000 Digital Microscope, using a lens with a range of 13–130×. Multiple images through the focal plane were taken of a structure and these were combined to produce a single in-focus image, using the software associated with the Keyence System.

Together with morphological studies, we also analyzed DNA barcodes (the 5’ region of the cytochrome c oxidase I (CO1) gene, Hebert et al. 2003) whenever available. DNA barcodes for all ACG inventory Sendaphne specimens were obtained using DNA extracts prepared from single legs using a glass fibre protocol (Ivanova et al. 2006). Briefly, total genomic DNA was re-suspended in 30 μl of dH2O, and a 658-bp region near the 5’ terminus of the CO1 gene was amplified using standard primers (LepF1–LepR1) following established protocols (Smith et al. 2006, 2007, 2008). If the initial 658 bp amplification was unsuccessful, smaller sequences were generated using internal primers. If each amplification worked a composite sequence was generated, however in cases where only one read amplified, this shorter sequence was used. All information for the sequences associated with each individual specimen can be retrieved from the Barcode of Life Data System (BOLD) (Ratnasingham and Hebert 2007) using the following public DOI: https://doi.org/10.5883/DS-SENDAPH. A Neighbor-Joining tree based on Kimura 2-parameter distances of all described species of Sendaphne with DNA barcodes available in BOLD was also generated (Suppl. material 2).

Results

Six new species of Sendaphne are described below, increasing the total known species from five to 11. We are aware of potential additional new species in the CNC collection, from Costa Rica (but not ACG), Ecuador, and Brazil. However, they are not described here because each is only represented by a single specimen and they are not sufficiently distinct to warrant description without further specimens and evidence.

Sendaphne is a Neotropical genus. To date it is most abundant and diverse in South America (eight species), while Central America has three species. It extends from 23° N in central Mexico (Durango) to 27° S in Paraguay (Pirapo) and southern Brazil (Santa Catarina). Most of the species have been collected in rain forests, at altitudes between 100 m and 900 m. However, a few species have been found only in cloud forests between 1,450 m and 1,900 m. The specimens collected at higher altitudes have darker coloration (especially on mesosoma and metasoma) than those found in the lowlands.

An interesting result of our morphological study was the relation between body and fore wing lengths. Body proportions in Microgastrinae have not been explored in detail, but in most genera and species with available data the fore wing length tends to be slightly longer than the body length (usually by 0.1–0.2 mm). In the specimens of Sendaphne described here, the body length was longer than the fore wing length (usually by 0.2–0.4 mm). The main reason is the long and slender body form (rather than exceptionally short wings), and an unusually enlarged and extended hypopygium.

Observations of COI barcodes for Sendaphne

Of 46 specimens sampled (Suppl. material 1), we recovered sequences for only 24 specimens. Only three sequences were over 600 base pairs, while the rest were mostly minibarcodes of 102–390 base pairs each. However, six out of the 11 known species now have some molecular data associated (Suppl. material 2). Many of the initial amplifications from ACG were characterised by unintended amplification of the endosymbiont bacteria Wolbachia that were amplified in the first round LepF1/LepR1. These sequences were not mistaken for the wasp DNA (Smith et al. 2012) and are retained on BOLD in the trace files. The standard strategy of amplifying two smaller and overlapping regions secondarily was then followed (as the first round was considered a ‘failure’). In this case the 5’ amplification (LepF1/C_ANTMR1D) worked and the 3’ amplification (RonMWASPdeg_t1 - LepR1) failed. Thus many of the ACG Sendaphne are ~260–280bp in length. Finally, regarding the CNC specimens, these were, on average, 35 years old. In those cases, greater success with smaller amplifications compared to larger amplifications is to be expected (Hajibabaei et al 2006). In 2010 and 2011 when these specimens were submitted to the Biodiversity Institute of Ontario (Guelph) for DNA barcoding, they only had the primers generating mini-barcode (smaller amplicons) used for PCR. The longer amplicon generating primer pairs were not attempted on these specimens.

Sendaphne

Sendaphne Nixon, 1965: 203.

Diagnosis

Glossa elongate and bilobate (Figs 2, 9, 34, 48, 58, 70). Lateral face of scutellum with polished area (=lunules) occupying most of the lateral face (Figs 5, 12, 19, 28, 44, 49, 56). Propodeum usually smooth and without carina (exceptionally having sparse punctures and few rugae on the nucha) (Figs 7, 13, 19, 28, 33, 44, 49, 60, 74). Metacoxa very long, about the same length as metafemur length and metatibia length (Figs 8, 13, 15, 29, 50, 54, 61, 68, 74). Mediotergite 1 strongly narrowing towards posterior margin (Figs 4, 14, 19, 50, 51). Mediotergite 2 subtriangular, much longer medially than its width at its anterior margin (and usually also longer medially than its width at posterior margin). Ovipositor very long for a microgastrine wasp (two times longer than metatibia length) and strongly curved (Figs 1, 20, 35, 53, 54, 57, 64, 68); apex of ovipositor usually not sinuate (exceptionally with very slight sinuation). Fore wing with very wide first discal cell, and with small areolet (Figs 3, 17, 17, 24, 39, 55, 62, 69) (areolet sometimes not well-defined because veins 3RSa and r-m are spectral, as in Figs 10, 31, 47). Body color often mostly yellow to orange (with a few exceptions from species collected at higher altitudes, which have head, mesosoma and parts of metasoma dark brown to black). Body length longer than the fore wing length, usually by 0.2–0.4 mm. Within Microgastrinae, Sendaphne can only be confused with Promicrogaster, but the later has a more transverse mediotergite 2, apex of ovipositor clearly sinuate, and propodeum usually with more sculpture and carination present.

Figures 1–7.

Sendaphne anitae, female holotype. 1 Habitus, lateral view 2 Head, frontal view 3 Fore wing 4 Metasoma, dorsal view 5 Head and mesosoma (partially) lateral view 6 Metasoma, ventro-lateral view 7 Mesosoma and metasoma (partially) dorsal view.

Figures 8–14.

Sendaphne sulmo, male specimen from Mexico. 8 Habitus, lateral view 9 Head, frontal view 10 Fore wing 11 Meso- and metasoma (partially), lateral view, and hind legs 12 Head and mesosoma, dorsal view 13 Meso- and metasoma (partially), dorsal view 14 Metasoma, dorsal view.

Figures 15–22.

Sendaphne dianariaspennae, female holotype. 15 Habitus, lateral view 16 Head, frontal view (some inclination downwards) 17 Fore wing 18 Propodeum and metasoma, dorsal view 19 Meso- and metasoma (partially), dorsal view. 20–22: Details of the ovipositor, ovipositor sheaths, and hypopygium.

Figures 23–28.

Sendaphne broadi, male holotype. 23 Habitus, lateral view 24 Fore wing 25 Metasoma, dorsal view 26 Metasoma, lateral view 27 Head and mesosoma (partially), dorsal view 28 Mesosoma and T1-T2, dorsal view.

Figures 29–36.

Sendaphne jatai, female specimen from Brazil. 29 Habitus, lateral view 30 Head and mesosoma (partially), dorso-lateral view 31 Fore wing 32 Metasoma, dorsal view 33 Scutellar disc, propodeum and T1 (partially), dorsal view 34 Head, frontal view 35 Metasoma, lateral view 36 Mesosoma and metasoma (partially), dorsal view.

Figures 37–44.

Sendaphne bennetti, male holotype. 37 Habitus, lateral view 38 Habitus, dorsal view 39 Fore wing 40 Head and mesosoma (partially), dorsal view 41 Head, fronto-lateral view 42 Metasoma, dorsal view 43 Metasoma, lateral view 44 Scutellar disc, propodeum and T1-T2 (partially), dorsal view.

Figures 45–53.

Sendaphne olearus, female specimen from Brazil. 45 Habitus, lateral view 46 Head and mesosoma, lateral view 47 Fore wing 48 Head, frontal view 49 Mesosoma and T1, dorsal view 50 Mesosoma and metasoma (partially), dorsal view 51 Metasoma, dorsal view 52 Metasoma, lateral view 53 Details of the ovipositor and ovipositor sheaths.

Figures 54–60.

Sendaphne paranaensis, female specimen from Brazil. 54 Habitus, lateral view 55 Fore wing 56 Head and mesosoma (partially), dorsal view 57 Metasoma and legs, lateral view 58 Head, frontal view 59 Metasoma (partially), dorsal view 60 Scutellar disc, propodeum and T1-T2 (partially), dorsal view.

Figures 61–67.

Sendaphne penteadodiasae, female holotype. 61 Habitus, lateral view 62 Fore wing 63 Metasoma, dorsal view 64 Ovipositor and hypopygium, lateral view 65 Head, frontal view 66 Head and mesosoma (partially), dorsal view 67 Propodeum, T1-T2 (partially), dorsal view.

Figures 68–75.

Sendaphne rogerblancoi, female holotype. 68 Habitus, lateral view 69 Fore wing 70 Head, frontal view 71 Hypopygium and hind legs (partially), lateral view 72 Ovipositor, ovipositor sheaths, and hypopygium, lateral view 73 Head and mesosoma (partially), lateral view 74 Propodeum and T1, dorsal view 75 Propodeum and mesosoma, dorsal view.

Key to Sendaphne species

[This key is intended for female specimens, although two species are only known from males, and in those cases the key accommodates them. Generally, males tend to have darker coloration than the females, especially on the metasoma].

1 Head and mesosoma entirely dark brown to black (Figs 23, 24, 27, 28, 3744, 6875), metasoma entirely dark brown (Figs 69, 74, 75) or at most with mediotergites 1–2 orange-yellow (as in Figs 24, 25, 28, 39, 44) [all specimens collected above 1,460 m, in cloud forests] 2
Color variable but much lighter than previous couplet, if head entirely dark brown to black, then mesosoma entirely or mostly yellow, orange or reddish-yellow, and metasoma at most with brown bands on posterior margins of mediotergites 3+ [all specimens collected below 900 m, in rainforests] 4
2(1) Tegula, metacoxa, and mediotergites 1–2 dark brown to black (Figs 69, 7375); body very slender, metasoma longer than combined length of head and mesosoma (Fig. 68) [Costa Rica] Sendaphne rogerblancoi Fernández-Triana & Whitfield, sp. n.
Tegula, metacoxa and mediotergites 1–2 yellow to reddish-yellow (Figs 2328, 3740, 4244); body less slender, metasoma shorter than combined length of head and mesosoma (Figs 23, 37) [Ecuador, Mexico] 3
3(2) Mediotergite 1 length 6.0 × its width at posterior margin, mediotergite 2 length 1.5 × its width at posterior margin (Figs 24, 25, 28); first discal cell 2.0 × as wide as high (Fig. 24) [Ecuador] Sendaphne broadi Fernández-Triana & Whitfield, sp. n.
Mediotergite 1 length 4.0 × its width at posterior margin, mediotergite 2 length 1.0 × its width at posterior margin (Figs 39, 42, 44); first discal cell 1.3 × as wide as high (Fig. 39) [Mexico] Sendaphne bennetti Fernández-Triana & Whitfield, sp. n.
4(1) Head yellow to reddish-yellow 5
Head dark brown to black 7
5(4) Mesosoma with dark brown areas on anteromesoscutum and mesopleuron (Figs 54, 56, 58); dark brown coloration on posterior margin of mediotergite 3 and most of mediotergites 4–6 (Figs 54, 55, 57, 59) [Brazil, Paraguay] Sendaphne paranaensis Scatolini & Penteado-Dias, 1999
Mesosoma uniformly orange-yellow to reddish-yellow; metasoma either entirely yellow or with brown bands on posterior margin of mediotergites 4–6 6
6(5) Metasoma with brown bands on posterior margin of mediotergites 4–6 (Figs 5053) [Brazil, French Guiana, Peru] Sendaphne olearus Nixon, 1965
Metasoma entirely yellow (Figs 31, 32, 35, 36) [Brazil, Ecuador, French Guiana] Sendaphne jatai Penteado-Dias, 1995
7(4) Female metasoma either with extensive dark brown coloration on tergites 3+ or with some narrow brown bands on posterior margin of mediotergites 5–7 (Figs 63, 64) 8
Female metasoma entirely yellow (Figs 3, 4, 7, 18, 19, 22) 9
8(7) Metasoma with dark brown bands on posterior margin of mediotergites 3–6 and mediotergite 7 entirely dark brown to black; fore wing vein 1Cu-a much shorter than vein 1Cu-b; T1 10.0 × as long as width at posterior margin; T1 2.0 × as long as T2; metacoxa 1.1 × as long as metafemur Sendaphne brasilianus Penteado-Dias, 1995
Metasoma with some narrow brown bands on posterior margin of mediotergites 5–7 (Figs 63, 64); fore wing veins 1Cu-a and 1Cu-b subequal; T1 6.0 × as long as width at posterior margin; T1 1.4 × as long as T2; metacoxa 0.9–1.0 × as long as metafemur Sendaphne penteadodiasae Fernández-Triana & Whitfield, sp. n.
9(7) Fore wing r and 2RS not clearly distinct from each other (Fig. 3); mediotergite 2 length 1.0 × its width at posterior margin (Figs 3, 4, 7); face centrally orange (Fig. 2); ovipositor 1.8–1.9 × as long as metatibia [Ecuador] Sendaphne anitae Fernández-Triana & Whitfield, sp. n.
Fore wing with veins r and 2RS clearly distinct from each other, and meeting at a sharp angle (not clearly visible in Fig. 17); mediotergite 2 length 1.8–2.0 × its width at posterior margin (Figs 18, 19); face entirely dark brown to black (Fig. 16); ovipositor 2.0–2.2 × as long as metatibia [Brazil, Colombia, or Mexico] 10
10(9) Distance between anatomical line tangent to posterior margin of anterior ocellus and anterior margin of posterior ocelli 0.5 × diameter of anterior ocelli (Fig. 77); ocular–ocellar line 2.5 × as long as posterior ocellus diameter; interocellar distance 1.6 × as long as posterior ocellus diameter; T1 relatively narrower medially, T1 width at half length of tergite clearly less than width at anterior margin, and 1.5 × as wide as width at posterior margin (Fig. 78); T2 1.8 × as long as wide [Mexico] Sendaphne sulmo Nixon, 1965
Distance between anatomical line tangent to posterior margin of anterior ocellus and anterior margin of posterior ocelli 0.2–0.3 × diameter of anterior ocelli (partially visible in Fig. 16); ocular–ocellar line 2.1–2.4 × as long as posterior ocellus diameter; interocellar distance 1.2–1.4 × as long as posterior ocellus diameter; T1 relatively wider medially, T1 width at half length of tergite about same as width at anterior margin, and at least 2.0 × as wide as width at posterior margin (Figs 18, 19); T2 2.0 × as long as wide [Brazil, Colombia] Sendaphne dianariaspennae Fernández-Triana & Whitfield, sp. n.
Figures 76–78.

Sendaphne sulmo, female holotype. 76 Habitus, lateral view 77 Head and mesosoma (partially), dorsal view 78 Propodeum and metasoma, dorsal view.

Figures 79–80.

Sendaphne olearus, female specimen from Peru. 79 Habitus, lateral view 80 Detail of the fore wing.

Taxonomic treatment of species, in alphabetical order

Sendaphne anitae Fernández-Triana & Whitfield, sp. n.

Figs 1–7

Holotype

Female, CNC. ECUADOR, Napo, Reventador; 11.iii.1983; coll. L. Huggert. DNA Voucher code: CNCHYM 07027.

Paratypes

4 ♀ (CNC), same locality and collecting date than holotype. Two of the specimens also have DNA Voucher codes: CNCHYM 07026, CNCHYM 07028.

Diagnosis

This species is morphological similar to Sendaphne dianariaspennae and S. sulmo but differs from those species in the shape and angle of junction of veins r and 2RS, shape of mediotergite 2, ovipositor length, and head coloration (orange-yellow on clypeus and face centrally in anitae, head entirely dark brown to black in dianariaspennae and sulmo).

Description

Head color: dark brown to black, except for orange-yellow on clypeus and face centrally. Mesosoma color: orange-yellow. Tegula color: orange-yellow. Metasoma color (dorsally): yellow. Metacoxa color: yellow. Anatomical line tangent to posterior margin of anterior ocellus crossing very slightly (less than 0.01 mm) above anterior margin of posterior ocelli. Ocular–ocellar line: 0.20 mm. Interocellar distance: 0.09 mm. Posterior ocellus diameter: 0.09 mm. Body length: 4.0–4.5 mm. Fore wing length: 3.6–4.1 mm. Ovipositor length: 2.4–2.6 mm. Metacoxa length: 1.3 mm. Metafemur length: 1.2 mm. Metatibia length: 1.3–1.4 mm. T1 length/width at posterior margin: 0.6 mm/0.15 mm. T2 length/width at posterior margin: 0.26–0.30 mm/0.3 mm.

Distribution

Only known from the type locality in Ecuador.

Molecular data

No DNA was recovered from the three specimens sampled.

Etymology

Named after Ana María (Anita) Fernández Galliano, daughter of the senior author, for being such a joyful and wonderful person.

Sendaphne bennetti Fernández-Triana & Whitfield, sp. n.

Figs 37–44

Holotype

Male, CNC. MEXICO, Durango, 39km W of La Ciudad, 2120m; 2.vii.1964; coll. WRM Mason. DNA Voucher code: CNCHYM 07032.

Diagnosis

This species is morphologically similar to Sendaphne broadi from Ecuador, but S. bennetti has a wider mediotergite 1 (length 4.0 × its width at posterior margin vs 6.0 × in broadi), a less transverse first discal cell in the fore wing (1.3 × as wide as high vs 2.0 ×), and a geographical distribution far apart.

Description

Head color: black. Mesosoma color: black. Tegula color: yellow. Metasoma color (dorsally): mediotergites 1, 2 and anterior half of 3 yellowish-red, rest dark brown to black. Metacoxa color: yellow. Anatomical line tangent to posterior margin of anterior ocellus crossing beneath anterior margin of posterior ocelli. Ocular–ocellar line: 0.21 mm. Interocellar distance: 0.14 mm. Posterior ocellus diameter: 0.09 mm. Body length: 4.1 mm. Fore wing length: 4.0 mm. Metacoxa length: 1.1 mm. Metafemur length: 1.2 mm. Metatibia length: 1.4 mm. T1 length/width at posterior margin: 0.60 mm/0.12 mm. T2 length/width at posterior margin: 0.35 mm/0.35 mm.

Distribution

Only known from the type locality in Mexico.

Molecular data

No DNA could be recovered from the specimen sampled.

Comments

Even though only one male specimen is known, it is sufficiently distinctive to warrant description. This species is the northernmost known distribution of the genus Sendaphne.

Etymology

Named after Dr. Andrew Bennett of the Canadian National Collection, Ottawa, and Canadian expert on Ichneumonidae, in appreciation for his support and encouragement to study braconid wasps.

Sendaphne brasilianus Penteado-Dias, 1995: 251.

Holotype

Female, DCBU. BRAZIL. Distrito Federal, Brasília, xii-1981, Malaise trap (not examined).

Diagnosis

This species is morphologically similar to Sendaphne penteadodiasae but it has a slightly different color pattern, fore wing vein 1Cu-a much shorter than vein 1Cu-b (subequal in penteadodiasae), much longer and narrow T1, and slightly longer metacoxa.

Distribution

Only known from the type locality in Brazil.

Molecular data

No specimen is known to have been sampled for DNA.

Comments

We could not study a specimen of this species, but the original description is sufficiently detailed for recognition, including several line drawings of the metasoma, fore wing, tip of antenna and hind leg (Penteado-Dias 1995).

Sendaphne broadi Fernández-Triana & Whitfield, sp. n.

Figs 23–28

Holotype

Male, CNC. ECUADOR, Napo, 5km S of Baeza, 1,700 m; 13.ii.1983; coll. Masner & Sharkey. DNA Voucher code: CNCH3323.

Paratypes

2 ♂ (CNC). One specimen with same locality and collecting date than holotype, the other collected at 1900 m on 9.ii.1983. DNA Voucher codes: CNCH3322 and CNCH3324.

Diagnosis

This species is morphologically similar to Sendaphne bennetti from Mexico, but S. broadi has a narrower mediotergite 1 (length 6.0 × its width at posterior margin vs 4.0 × in bennetti), a more transverse first discal cell in the fore wing (2.0 × as wide as high vs 1.3 ×), and a geographical distribution far apart.

Description

Head color: dark brown. Mesosoma color: dark brown. Tegula color: yellow. Metasoma color (dorsally): mediotergites 1–2 yellowish-brown, rest brown to dark brown. Metacoxa color: yellow. Anatomical line tangent to posterior margin of anterior ocellus crossing very slightly (less than 0.01 mm) above anterior margin of posterior ocelli. Ocular–ocellar line: 0.17 mm. Interocellar distance: 0.09 mm. Posterior ocellus diameter: 0.07 mm. Body length: 3.5–3.7 mm. Fore wing length: 3.3–3.5 mm. Metacoxa length: 1.1 mm. Metafemur length: 1.2 mm. Metatibia length: 1.4 mm. T1 length/width at posterior margin: 0.45 mm/0.06–0.07 mm. T2 length/width at posterior margin: 0.25 mm/0.20 mm.

Distribution

Ecuador.

Molecular data

No DNA could be recovered from the three specimens sampled.

Comments

Even though only male specimens of this species are known, they are sufficiently distinctive to warrant description.

Etymology

Named after Dr. Gavin Broad, of the Natural History Museum, London, England, and British expert on Ichneumonidae, in appreciation for his support over the years, including sharing pictures of and facilitating access to type material deposited in London.

Sendaphne dianariaspennae Fernández-Triana & Whitfield, sp. n.

Figs 15–22

Holotype

Female, CNC. BRAZIL, Rio de Janeiro, Mangaratiba; i.1976; coll. M. Alvarenga. DNA Voucher code: CNCHYM 07025.

Paratypes

3 ♀, 1 ♂ (CNC), Brazil, same locality and date than holotype. 1 ♀ (CNC), Brazil, Nova Teutonia, 27°11'S, 52°23'W, 300–500m, 21.iii.1961, coll. F. Plaumann. 2 ♂ (CNC), Brazil, Guanabara and Represa Rio Grande, i.1969 and i.1972. 1 ♂ (CNC), Brazil, Caruaru, Pernambuco, v.1972. 8 ♀, 10 ♂ (Humboldt Institute), Colombia, Magdalena, PNN Tayrona Zaino, 50m, 11°20'N, 74°2'W, specimens collected between 13.v.2000 and 30.viii.2000, coll. R. Henriquez.

Diagnosis

This species is morphological similar to Sendaphne sulmo but differs in the smaller, less separated ocelli (which form a lower triangle, compared to higher triangle in sulmo), T1 relatively wider medially, and T2 relatively slender than sulmo. The geographical distribution of the two species is more than 2,000 km apart.

Description

Head color: dark brown to black. Mesosoma color: orange-yellow. Tegula color: orange-yellow. Metasoma color (dorsally): yellow. Metacoxa color: yellow. Anatomical line tangent to posterior margin of anterior ocellus crossing slightly (0.01–0.02 mm) above anterior margin of posterior ocelli. Ocular–ocellar line: 0.17 mm. Interocellar distance: 0.10 mm. Posterior ocellus diameter: 0.07–0.08 mm. Body length: 3.2–3.5 mm. Fore wing length: 3.0–3.2 mm. Ovipositor length: 2.3–3.0 mm. Metacoxa length: 0.90–0.95 mm. Metafemur length: 0.90 mm. Metatibia length: 1.12–1.35 mm. T1 length/width at posterior margin: 0.45–0.50 mm/0.05–0.06 mm. T2 length/width at posterior margin: 0.35 mm/0.18–0.20 mm.

Distribution

Brazil and Colombia.

Molecular data

Four of the paratypes from Brazil (DNA Voucher codes: CNCHYM 07023, CNCHYM 07024 and CNCHYM 07040) as well as the holotype were sampled for DNA. Only one of the paratypes (CNCHYM 07040) rendered a minibarcode of 103 base pairs.

Comments

The Brazilian specimens of S. dianariaspennae were collected between January and March, while the Colombian specimens were collected between May and August.

Etymology

Named after Diana Carolina Arias-Penna (Colombia), in recognition of her blossoming career studying neotropical Braconidae, especially species from South America.

Sendaphne jatai Penteado-Dias, 1995: 252.

Figs 29–36

Holotype

Female, DCBU. BRAZIL. Sao Paulo, Reserva Ecológica do Jataí, gallery forest of Mogi River; 24.v.1991; sweeping (not examined).

Specimens examined

1 ♀ (CNC), Brazil, Mato Grosso, Sinop, x.1974, coll. M. Alvarenga, DNA Voucher code: CNCH3320. 1 ♀, 1 ♂ (CNC), Ecuador, Pichincha, 47 km S of Santo Domingo, Rio Palenque, 200m, 18–30.v.1975 and 22-31.vii.1976, coll. S. & J. Peck, DNA Voucher codes: CNCHYM 07034 and CNCHYM 07035. 1 ♀ (FUSAGx), French Guiana, Saul, Crique popote, Mont Belvédere, 3°36'N, 53°10'W, ii.2001, coll. J. Tarin.

Diagnosis

This is the only species where female specimens have the body entirely yellow.

Distribution

Brazil, Ecuador, French Guiana.

Molecular data

The two specimens from Ecuador rendered minibarcodes of 102 base pairs (CNCHYM 07034) and 164 base pairs (CNCHYM 07035).

Comments

The male specimen from Ecuador has some dark bands on mediotergites 5–7, but otherwise is similar to the original description.

Sendaphne olearus Nixon, 1965: 204.

Figs 45–53, 79–80

Holotype

Female, BMNH. BRAZIL, Nova Teutonia, 2.iii.1937 (not examined).

Specimens examined

1 ♀ (CNC), Brazil, Nova Teutonia, 27°11'S, 52°23'W, 300-500m, 2.iv.1966, DNA Voucher code: CNCHYM 07019. 3 ♂ (FUSAGx), French Guiana, Kaw Mountain, Patawa, 4°32'42.20"N, 52°09'09.19"W, v.1999 and viii.1999, Malaise trap. 1 ♀ (BMNH) Peru, Loreto, Estacion Jenaro Herrera, 4°53'55.0"S, 73°39'00.4"W, 121m, 13–23.i.2011, D. Karlsson & N. Dale-Skey, BMNH(E) 2011-72, BMNH(E)#1249978.

Diagnosis

This species is similar to Sendaphne jatai, both having the lightest (i.e., more yellowish) coloration among all known species within the genus. S. olearus has brown bands on posterior margin of mediotergites 4–6, and its body coloration is generally more yellow-reddish (jatai body color is entirely yellow).

Distribution

Brazil, French Guiana, Peru.

Molecular data

The CNC specimen from the type locality in Brazil (CNCHYM 07019) rendered a minibarcode of 164 base pairs.

Comments

We could not see the holotype of this species, but the original description is adequately detailed for discrimination. We examined one female from the type locality (collected 30 years after the holotype female), the only known male specimens (from French Guiana), which were discussed in Braet (2006), and the first known specimen from Peru.

Sendaphne paranaensis Scatolini & Penteado-Dias, 1999: 53.

Figs 54–60

Holotype

Female, UFPR. BRAZIL, Reserva Biologica Samuel Klabin, Malaise trap (not examined).

Specimens examined

1 ♀ (CNC), Brazil, Rio de Janeiro, Guanabara, Silva Jardim, iii.1974, DNA Voucher code: CNCHYM 07022. 1 ♂ (CNC), Brazil, Espirito Santo, Castello, xi.1976, DNA Voucher code: CNCHYM 07039. 2 ♂ (CNC), Paraguay, Pirapo, 1–3.i.1972, DNA Voucher codes: CNCHYM 07037, and CNCHYM 07038.

Other material mentioned in the original description

Almost 70 specimens (females and males) from Brazil, Paraná, Telêmaco Borba, collecting dates between viii.1986 and iii.1987.

Diagnosis

S. paranaensis is one of only three Sendaphne species with head entirely yellow (or yellow-orange). It differs from the other two species (S. jatai and S. olearus) in having dark brown areas on the anteromesoscutum and mesopleuron, and mediotergites 4+ entirely black.

Distribution

Brazil, Paraguay.

Molecular data

Of the four specimens in the CNC sampled for DNA, only one (CNCHYM 07037) rendered a minibarcode of 164 base pairs.

Comments

The male specimens from Paraguay are much darker in coloration, but similar variation is mentioned in the original description for the male paratypes from Brazil (Scatolini and Penteado-Dias 1999: 53).

Sendaphne penteadodiasae Fernández-Triana & Whitfield, sp. n.

Figs 61–67

Holotype

Female, CNC. BRAZIL, Campina Grande, near Curitiba, 15.ii.1966, coll. H. & M. Townes. DNA Voucher code: CNCHYM 07020.

Paratype

1 ♂, same locality as holotype, 10.ii.1966. DNA Voucher code: CNCHYM 07021.

Diagnosis

This species is morphologically similar to Sendaphne brasilianus but it has a slightly different color pattern, fore wing vein 1Cu-a subequal to vein 1Cu-b (much shorter in brasilianus), shorter and wider T1, and slightly shorter metacoxa.

Description

Head color: dark brown to black. Mesosoma color: orange-yellow. Tegula color: orange-yellow. Metasoma color (dorsally): mostly orange-yellow, with some narrow brown bands on posterior margin of mediotergites 5–7. Metacoxa color: yellow. Anatomical line tangent to posterior margin of anterior ocellus crossing very slightly (less than 0.01 mm) above anterior margin of posterior ocelli. Ocular–ocellar line: 0.12 mm. Interocellar distance: 0.10 mm. Posterior ocellus diameter: 0.08 mm. Body length: 4.2 mm. Fore wing length: 3.9 mm. Ovipositor length: 2.5 mm. Metacoxa length: 1.20 mm. Metafemur length: 1.25 mm. Metatibia length: 1.11 mm. T1 length/width at posterior margin: 0.35 mm/0.06 mm. T2 length/width at posterior margin: 0.25 mm/0.12 mm.

Distribution

Only known from the type locality in Brazil.

Molecular data

No DNA could be recovered from the two specimens sampled.

Comments

The specimens of this species (housed in the CNC) were previously identified by W.R.M. Mason as “Sendaphne sulmo”. However, the morphological differences from the original description of Sendaphne sulmo (see key and diagnosis above), and the disparate geographical distribution allows us to consider the Brazilian and Mexican specimens as separate species.

Etymology

Named after Dr. Angélica Maria Penteado Martins-Dias (Brazil), in recognition of her career studying Braconidae, and also for her work describing most of the previously known species of Sendaphne.

Sendaphne rogerblancoi Fernández-Triana & Whitfield, sp. n.

Figs 68–75

Holotype

Female, CNC. COSTA RICA, Guanacaste, Area de Conservación Guanacaste, Sector Cacao, Sendero Cima, 1,460 m, 10.93328, -85.45729; 18.xii.2008; coll. D.H. Janzen & W. Hallwachs. DNA Voucher code: DHJPAR0031465.

Paratypes

4 ♀, 5♀ (BMNH, CNC, INBio, NMNH), same locality as holotype. Collecting dates are mostly on December 2008, with one record each in November 2008, March 2009, and April 2009. All specimens are from one Malaise trap at the very peak of Volcán Cacao.

Diagnosis

This is the most distinctive species of Sendaphne based on coloration (head, mesosoma, metasoma, and metacoxa black), shape of first discal cell, and narrow mediotergite 1.

Description

Head color: black. Mesosoma color: black. Tegula color: dark brown. Metasoma color (dorsally): black. Metacoxa color: dark brown to black. Anatomical line tangent to posterior margin of anterior ocellus crossing very slightly (less than 0.01 mm) above anterior margin of posterior ocelli. Ocular–ocellar line: 0.17–0.18 mm. Interocellar distance: 0.09 mm. Posterior ocellus diameter: 0.08 mm. Body length: 4.3–4.6 mm. Fore wing length: 4.2–4.3 mm. Ovipositor length: 3.8–4.2 mm. Metacoxa length: 1.10–1.11 mm. Metafemur length: 0.85–0.90 mm. Metatibia length: 1.5–1.6 mm. T1 length/width at posterior margin: 0.80–0.85 mm/0.15–0.16 mm. T2 length/width at posterior margin: 0.45 mm/0.25 mm.

Distribution

Only the summit cloud forest at 1,450 m on Volcán Cacao, northwestern Costa Rica.

Molecular data

In BOLD there are data for 16 specimens of this species (the holotype, the paratypes and other specimens that we could not examine) which rendered partial barcodes, most of them from 260 to 390 base pairs. Only the holotype (DNA Voucher code: DHJPAR0031465) had a longer barcode (633 base pairs).

Etymology

This unique species, the only Sendaphne known from Costa Rica so far, is named after Sr. Roger Blanco Segura, of Area de Conservación Guanacaste (ACG), northwestern Costa Rica, in recognition of his 3+ decades of intense care and management of ACG in an enormous variety of circumstances and for a very large array of purposes.

Sendaphne sulmo Nixon, 1965: 204.

Figs 8–14, 76–78

Holotype

Female, BMNH. MEXICO, Tabasco, Teapa (not examined).

Specimens examined

3 ♂ (CNC). Mexico, Chiapas, 16°58'N, 91°47'W, 560 m; (collecting dates: 6.ix.1978, 28.x.1978, 8–11.xi.1978); coll. J. Rawlins. DNA Voucher codes: CNCHYM 07030 CNCHYM 07031 and CNCHYM 07033.

Diagnosis

This is the only known species of Sendaphne with a higher ocellar triangle (i.e., anatomical line tangent to posterior margin of anterior ocellus crosses far above anterior margin of posterior ocelli). The distance between anatomical line tangent to posterior margin of anterior ocellus and anterior margin of posterior ocelli is 0.5 × the diameter of anterior ocelli (Fig. 77), while for all other known species of Sendaphne it is usually 0.1–0.3 ×.

Distribution

Only known from lowlands (100–560 m) of Mexico.

Comments

We could only study some photos of the holotype (Figs 7678) and the original description which included a line drawing of the metasoma (Nixon 1965: 207, Figure 255). The drawing shows a T1 slightly narrower medially than the photos of the actual holotype reveal, but the rest of the original description is in agreement with the photos we examined. The males mentioned above are, however, different from the female holotype in having a darker anteromesoscutum, scutellar disc, and metasoma (Figs 8–14), and also the ocellar triangle is not as elevated. Lacking more specimens to examine (especially females), we have refrained from considering those male specimens as a different species because the two localities are not too far apart and males are known to be darker in other species of Sendaphne. If more material becomes available for study in the future, the status of those specimens may be clarified.

Molecular data

The three male specimens sampled for DNA rendered minibarcodes of 103 base pairs each.

Acknowledgements

We emphatically and gratefully acknowledge the support of the ACG parataxonomist team in finding and rearing these caterpillars, their parasitoids and their hyperparasitoids, and Area de Conservacion Guanacaste (ACG) for preserving the forests in which they live, and the Guanacaste Dry Forest Conservation Fund, the Wege Foundation, the International Conservation Fund of Canada, the JRS Biodiversity Foundation, Jessie Hill, Steve Stroud, Permian Global, and the University of Pennsylvania for funding portions of the research. Several Colombian specimens were collected during a project supported by NSF grant DEB 0205982 awarded to M. J. Sharkey and B. V Brown. This study was also supported by NSF DEB 0515699 to DHJ and by a Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant to MAS. Laboratory analyses of these sequences were funded by the Government of Canada through Genome Canada and the Ontario Genomics Institute (2008-0GI-ICI-03), and by BOLD/iBOL of the Biodiversity Institute of Ontario and University of Guelph. JFT thanks Yves Braet (Belgium) for making specimens from French Guiana available for this study and Gavin Broad (BMNH) for sending pictures of the specimens housed in London. The suggestions from two anonymous reviewers and the editor considerably improved the final version of this paper.

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