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Research Article
Two new species and distribution records for the genus Bohayella Belokobylskij, 1987 from Costa Rica (Hymenoptera, Braconidae, Cardiochilinae)
expand article infoIlgoo Kang‡, Scott R. Shaw§, Nathan P. Lord‡
‡ Louisiana State University Agricultural Center, Baton Rouge, United States of America
§ University of Wyoming, Laramie, United States of America

Corresponding author: Ilgoo Kang ( ikang1@lsu.edu )

Academic editor: Jose Fernandez-Triana
© 2020 Ilgoo Kang, Scott R. Shaw, Nathan P. Lord.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Kang I, Shaw SR, Lord NP (2020) Two new species and distribution records for the genus Bohayella Belokobylskij, 1987 from Costa Rica (Hymenoptera, Braconidae, Cardiochilinae). ZooKeys 996: 93-105. https://doi.org/10.3897/zookeys.996.59075
ZooBank: urn:lsid:zoobank.org:pub:2550092B-AC75-48F1-AEDF-84E3F30DC4EC
Open Access

Abstract

Two new species of Bohayella Belokobylskij, 1987 from Costa Rica are described: Bohayella geraldinae Kang, sp. nov. and Bohayella hansoni Kang, sp. nov. These are new distribution records for the genus in the Neotropical region. In addition, a key to species of the genus Bohayella of Costa Rica is presented. The current work elevates the number of species included in Bohayella from nine to eleven.

Keywords

Morphology, New World, parasitoid wasp, taxonomy

Introduction

Costa Rica is one of the biodiversity hotspots, and a total estimated hymenopteran fauna in the country is ~ 20,000 species, including ~ 2,000 estimated species of braconid wasps (Gaston et al. 1996). Cardiochilinae is a subfamily of Braconidae, containing 17 genera and 220+ species (Yu et al. 2012; Kang et al. 2020). Bohayella Belokobylskij, 1987 (Belokobylskij 1987) is an unusual genus of the subfamily, with nine previously described species that are only known from the Old World, including Afrotropical, Australasian, Oriental, and southern central Palearctic regions (Dangerfield et al. 1999; Mercado and Wharton 2003; Yu et al. 2012). Among the nine Old World species of Bohayella, two species, B. adina (Wilkinson, 1930) and B. exiguurus (Huddleston & Walker, 1988), have rearing records (Huddleston and Walker 1988). B. adina was reared from larvae of Phazaca theclata (Guenée, 1857) (Lepidoptera: Uraniidae) in India, (Beeson and Chatterjee 1935; Dangerfield 1995; Dangerfield et al. 1999), and B. exiguurus was reared from larvae of the citrus looper Cleora tulbaghata (Felder & Rogenhofer, 1875) (Lepidoptera: Geometridae) South Africa (Dangerfield et al. 1999).

Cardiochiles nigricans Mao, 1949 (Mao 1949) was transferred into Bohayella by Dangerfield et al. (1999) and recorded as the first species of Bohayella in the New World. Mercado and Wharton (2003) transferred the species into Toxoneuron Say, 1836 because the first metasomal tergite (T1) of the species is different from T1 of other members of Bohayella. Subsequently, members of Bohayella have been restricted to the Old World and new species of the genus have not been reported from the New World.

The first author (IK) had the opportunity to examine Costa Rican cardiochiline specimens housed in University of Wyoming Insect Museum (UWIM). Using the key to world genera of the subfamily Cardiochilinae and other diagnostic characters of Bohayella (Dangerfield et al. 1999), nine Bohayella specimens were identified. The characters of New World Bohayella are discussed in detail in diagnosis and discussion sections of this paper. Other Costa Rican cardiochiline specimens borrowed from several institutions were examined, but no more specimens of Bohayella were discovered. As a result, the nine specimens of Bohayella were confirmed as two species based on morphological data. Herein, we describe two new species and present a key to species of the genus Bohayella of Costa Rica. Distribution maps for both species are included.

Materials and methods

Specimens for this project were provided by UWIM (University of Wyoming Insect Museum; 1000 East University Avenue, University of Wyoming, Laramie, Wyoming 82071-3354, USA). We conducted morphological analyses using a Leica MZ75 stereomicroscope. The morphological terms and terms of wings mostly follow Dangerfield (1995) and Dangerfield et al. (1999). Morphological terminology can be checked at the Hymenoptera Ontology website (http://portal.hymao.org/projects/32/public/ontology/) as well. Terms for sculpturing are based on Harris (1979). Color habitus images were taken using a Visionary Digital BK Plus imaging system (Dun, Inc.), equipped with a Canon EOS 5DS DSLR camera. Images were stacked via Zerene Stacker v.1.04 (Zerene Systems LLC.). All images were edited using Adobe Photoshop CS 6 (Adobe Systems, Inc). Body parts of each species were measured via Adobe Photoshop CS 6 (Adobe Systems, Inc). Each number in parentheses in species descriptions indicate 0.01 times the actual length, width, or height of each body part. For example, 42 and 124 in parentheses (42:124) indicate 0.42 mm and 1.24 mm, respectively. Distribution maps of two Bohayella species were produced using QGIS 3.10.0 (QGIS Development Team 2019). Google satellite maps were downloaded using the QuickMapServices plugin. The following abbreviations are used throughout the current paper: POL: distance between posterior ocelli, T1, T2 (second metasomal tergite), T3 (third metasomal tergite), T5 (fifth metasomal tergite), T6 (sixth metasomal tergite), T7 (seventh metasomal tergite), and T8 (eight metasomal tergite). Holotypes and paratypes are deposited in the UWIM.

Results

Bohayella Belokobylskij, 1987

Type species

Bohayella tobiasi Belokobylskij, 1987.

Diagnosis

(based on Dangerfield et al. (1999) with modifications and additions). Diagnostic characters of Bohayella based on Old World members were described in Belokobylskij (1987) (in Russian) and Dangerfield et al. (1999) (in English). The following are re-described or additional characters based on morphological characters of both Old World and New World members.

Members of the genus can be identified by setose compound eyes (length and density variable); ventro-posteriorly moderately extended gena (Fig. 3F).broad clypeus without clypeal tubercles (Figs 3B, 4C); absence of occipital carina; uni- or bi-dentate mandible; 5- or 6-segmented maxillary palpus; 4-segmented labial palpus; short mouthparts (galea and glossa); deep and broad notauli and scutellar sulcus (Figs 3C, F, 4B, D); scutellum with apical cup-like pit (Figs 3C, F, 4B, D); fully developed propodeal areola (Figs 3D, F, 4B, E); moderately to strongly sculptured pronotum and mesopleuron; presence of epicnemial carina (Figs 3A, 4A); well-defined and crenulate precoxal sulcus (Figs 3A, 4A); absence of apical cup-like projection of hind tibia (Figs 3A, 4A); cylindrical or antero-posteriorly slightly expanded hind basitarsus (but never expanded like hind basitarsi found in members of Hartemita Cameron, 1910) (Figs 3A, 4A); pectinate tarsal claw with sharp or obtuse apical tooth; entirely or apically infuscate forewing; absence of 1r vein of forewing; absence of 3r vein of forewing; basally angled or smoothly curved Rs vein; absence of 2-1A vein of hind wing; narrow and elongate T1 (median length of T1 4.0–6.3× longer than its apical width) (Figs 3D, F, 4B, E); short T2; a medio-basal ball-like projection of T2 (Figs 3D, F, 4B, E); short and truncate hypopygium (Figs 3A, 4A); short ovipositor (if protruded, strongly downcurved); short ovipositor sheath (< ~0.2× longer than hind tibia) (Figs 3A, 4A).

Key to species of the genus Bohayella of Costa Rica

1 Median crenula of notauli as long as median crenula of scutellar sulcus (A); scutellar sulcus with one median crenula (A); T3–T8 mostly pale (AA) B. geraldinae sp. nov.
– Median crenula of notauli shorter than median crenula of scutellar sulcus (B); scutellar sulcus with three crenulae (B); T3–T8 mostly melanistic (BB) B. hansoni sp. nov.

Bohayella geraldinae Kang, sp. nov.

http://zoobank.org/FC39B76A-3AC3-41D7-9420-7148410F6D2C
Figure 3

Material examined

Holotype Costa Rica • ♀; female, Heredia, 3 km S. Puerto Viejo OTS, La Selva; 100 m; x.1992; P. Hanson; huertos Malaise trap set by G. Wright. Paratypes Costa Rica • 1 ♀; same data as for holotype; xi.1992 • 1 ♂; male; same collecting data as for preceding; 10°26'N, 84°01'W; 4. iv. 1987; H. A. Hespenheide.

Diagnosis

Bohayella geraldinae sp. nov. can be recognized by the following combination of characters: apical maxillary palpomere as long as fifth maxillary palpomere; median crenula of notauli as long as median crenula of scutellar sulcus; scutellar sulcus with one median crenula; hind basitarsus antero-posteriorly slightly expanded; dorsal metasoma mostly pale.

Description

Female. Body 4.6–4.8 mm. Forewing length: ~ 4.2 mm Antenna length: ~ 4.8 mm. Head. Antenna 34-segmented. Interantennal space with well-developed median carina. POL ~ 1.38× longer than diameter of anterior ocellus (11:8) (Fig. 3F). Eye sparsely setose with short setae; median width of eye 0.75× longer than median width of gena in lateral view (36:48). Width of clypeus ~ 2.07× longer than height (60:29) (Fig. 3B). Malar space ~ 2.62× longer than basal width of mandible (34:13) (Fig. 3B). Mandible bidentate. Maxillary palpus 6-segmented; apical maxillary palpomere as long as fifth maxillary palpomere. Mesosoma. Mesoscutum with sharp margin (Fig. 3C, F). Notauli broadly converging at base, with 11 crenulae; median crenula of notauli as long as median crenula of scutellar sulcus (Fig. 3C, F). Scutellar sulcus with one median crenula (Fig. 3C, F). Postscutellar depression present (Fig. 3C, F). Propodeum rugulose with well-defined median areola; median transverse carina on propodeum reaching lateral margin (Fig. 3D, F). Pronotum dorso-posteriorly crenulate and antero-ventrally smooth. Mesopleuron dorsally and posteriorly with crenulate margin (Fig. 3A). Mesosternal sulcus broad and crenulate. Metapleuron rugulose. Legs. Basal spur on fore tibia ~ 0.86× longer than basitarsus (30:35). Width of hind femur ~ 0.34× longer than its length (42:124). Basal spur on hind tibia ~ 0.76× longer than basitarsus (58:76). Hind tarsal claw pectinate, with four sharp teeth (Fig. 3E). Wings. Forewing second submarginal cell trapezoidal, ~ 0.35× longer than maximum width (30:85); 3r absent (Note: one specimen has basally present 3r vein in particular angle); Rs sharply angled at basal third; stigma ~ 2.67× longer than medial width (80:30). 1CUa short, 0.23× longer than 1CUb (12:52). Hind wing 2-1A absent. Metasoma. T1 with a pair of lateral sutures posteriorly reduced, median length of T1 ~ 5.07× longer than apical width (71:14) (Fig. 3D, F). T2 with a ball-like projection, medially 0.21× longer than T1 (15:71) (Fig. 3D, F). T3 ~ 2.13× longer than T2 medially (32:15) (Fig. 3D, F). Protruded ovipositor sheath ~ 0.13× longer than hind tibia and apically setose (20:154) (Fig. 3A).

Male. Body ~ 5.0 mm. Same as female except for the following characters: antenna 32-segmented, melanistic color does not reach the dorsal margin of foramen magnum.

Color. Body mostly pale; the following areas are melanistic: antenna, vertex, frons, dorsal occiput, maxillary palpus, labial palpus, lateral mesonotal lobe (pale basally), lateral scutellum, margin of metanotum, apical fore femur, fore tibia, apical fore tarsus, apical mid femur, mid tibia, apical mid tarsus, apical hind femur, basal and apical hind tibia, apical hind tarsus, posterior T5 and T6 (weakly), entire T7 and T8, ovipositor sheath. Wings entirely infuscate, stigma entirely melanistic.

Host

Unknown.

Distribution

Bohayella geraldinae sp. nov. is known only from the La Selva Biological Station owned and managed by Organization for Tropical Studies (OTS) in Heredia, Costa Rica at an elevation of 100 m (Figs 1, 2). The station is located in the Caribbean lowlands, at a confluence of the Sarapiquí river and Puerto Viejo (McDade and Hartshorn 1994). According to Holdridge’s life zone system (Holdridge 1967), the station is in the tropical wet forest region (Hartshorn and Peralta 1987), and the average annual precipitation in the area is ~ 4,000 mm (Sanford et al. 1994).

Figure 1. 

Distribution map of the species of Bohayella in Costa Rica. Map data 2020 Google.

Figure 2. 

Distribution map of B. geraldinae sp. nov. in La Selva Biological Station in Costa Rica. Map data 2020 Google.

Figure 3. 

B. geraldinae sp. nov., holotype A lateral habitus B anterior head C dorsal mesonotum D dorsal propodeum and T1–T3 E hind tarsal claw F dorsal habitus.

Etymology

This species is named in honor of Dr Geraldine Wright, a former student of the second author (SRS), Rhodes Scholar, professor in the Department of Zoology in the University of Oxford (United Kingdom), and the person who set the trap that collected the specimens.

Bohayella hansoni Kang, sp. nov.

http://zoobank.org/7749425B-2B7F-4E69-A115-B65ED9CAD0CF
Figure 4

Material examined

Holotype Costa Rica • ♀; female, Puntarenas, San Vito, Estac. Biol., Las Alturas; 1,500 m; vi.1992; Paul Hanson; traps #1 + #2, Malaise. Paratypes Costa Rica • 2 ♀; same data as for holotype • 2 ♀; same collecting data as for preceding • 1 ♀; female; same collecting data as for preceding; 1,700 m; 11.iv.1993.

Diagnosis

Bohayella hansoni sp. nov. can be distinguished from B. geraldinae sp. nov. by the following characters: apical maxillary palpomere slightly longer than fifth maxillary palpomere; median crenula of notauli ~ 0.38× longer than median crenula of scutellar sulcus; scutellar sulcus with three crenulae; hind basitarsus cylindrical; dorsal metasoma mostly melanistic.

Description

Female. Body 3.9–4.1 mm. Forewing length: 3.9–4.1 mm Antenna length: 4.1–4.5 mm. Head. Antenna 32–34-segmented. Interantennal space with well-developed median carina. POL 1.22× longer than diameter of anterior ocellus (11:9) (Fig. 4B). Eye sparsely setose with short eye setae; length of eye ~ 0.86× longer than median width of gena in lateral view (31:36). Width of clypeus 2.00× longer than height (56:28) (Fig. 4C). Malar space 1.80× longer than basal width of mandible (36:20) (Fig. 4C). Mandible bidentate. Maxillary palpus 6-segmented; apical maxillary palpomere 1.31× longer than fifth maxillary palpomere (17:13). Mesosoma. Mesoscutum with sharp margin (Fig. 4B, D). Notauli broadly converging at base, with 11 crenulae; median crenula of notauli ~ 0.38× longer than median crenula of scutellar sulcus (6:16) (Fig. 4B, D). Scutellar sulcus with three crenulae (Fig. 4B, D). Postscutellar depression present (Fig. 4B, D). Propodeum rugulose, with well-defined median areola; median transverse carina on the propodeum reaching lateral margin (Fig. 4B, E). Pronotum dorso-posteriorly crenulate and antero-ventrally smooth. Mesopleuron dorsally and posteriorly with crenulate margin (Fig. 4A). Mesosternal sulcus broad and crenulate. Metapleuron rugulose. Legs. Basal spur on fore tibia ~ 0.87× longer than basitarsus (26:30). Width of hind femur ~ 0.30× longer than its length (33:111). Basal spur on hind tibia ~ 0.81× longer than basitarsus (58:72). Hind tarsal claw pectinate with four acute teeth. Wings. Forewing second submarginal cell trapezoidal, ~ 0.34× longer than its maximum width (26:77); 3r absent; Rs sharply angled at basal third; stigma ~ 2.82× longer than medial width (79:28). 1CUa short, 0.23× longer than 1Cub (11:47) (Fig. 4A). Hind wing 2-1A absent. Metasoma. T1 with a pair of lateral sutures posteriorly reduced, median length of T1 4.00× longer than apical width (56:14) (Fig. 4B, E). T2 with a ball-like projection, medially ~ 0.20× longer than T1 (11:56) (Fig. 4B, E). T3 ~ 2.55× longer than T2 medially (28:11) (Fig. 4B). Protruded ovipositor sheath ~ 0.20× longer than hind tibia and apically setose (26:129) (Fig. 4A).

Figure 4. 

B. hansoni sp. nov., holotype. A lateral habitus B dorsal habitus C anterior head D dorsal mesonotum E dorsal propodeum and anterior metasoma.

Color. Body mostly pale; the following areas melanistic: antenna, vertex, frons, dorsal occiput, maxillary palpus, labial palpus, lateral mesonotal lobe (basally pale), lateral scutellum, margin of metanotum, apical fore femur, fore tibia, apical fore tarsus, apical mid femur, mid tibia, apical mid tarsus, apical hind femur, basal and apical hind tibia, apical hind tarsus, T2–T8, ovipositor sheath. Wings entirely infuscate, stigma entirely melanistic.

Male. Unknown.

Host

Unknown.

Distribution

Bohayella hansoni sp. nov. is known only from the Las Alturas Biological research station owned and operated by Stanford University in Las Alturas, San Vito, Costa Rica at the elevations of 1,500 m and 1,700 m (Figs 1, 5).

Figure 5. 

Distribution map of B. hansoni sp. nov. from Las Alturas Biological Research Station. Map data 2020 Google.

Etymology

This species is named in honor of Dr Paul Hanson, collaborator and professor at the Escuela de Biología, Universidad de Costa Rica. He worked tirelessly for many years collecting and sorting Costa Rican braconids from Malaise samples. SRS is very grateful for his dedication to Hymenoptera studies.

Discussion

Most genus-level diagnostic characters are shared by both Old World and New World members (B. geraldinae sp. nov. and B. hansoni sp. nov.). None of the New World members have a mostly black body, 5-segmented maxillary palpi, or apically infuscate forewings. The following characters are only shared by New World members: angled Rs vein of forewing (Figs 3F, 4A), pectinate hind tarsal claw with sharp apical tooth (Fig. 3E), and antero-posteriorly slightly expanded hind basitarsus (Fig. 3A).

Specimens of B. hansoni sp. nov. collected at altitudes above 1,500 m have more melanistic metasoma than specimens of B. geraldinae sp. nov. collected at a low altitude of 100 m (Figs 3F, 4B). The melanism associated with high elevation was confirmed not only in braconid wasps such as members of the genus Sendaphne Nixon, 1965 (Nixon 1965) (Fernandez-Triana et al. 2014) and Meteorus pulchricornis (Wesmael, 1835) (Abe et al. 2013), but also in other hymenopteran insects such as members of a vespid species, Agelaia pallipes (Olivier, 1792) (de Souza et al. 2020) as well as an undescribed scelionid species of Lapitha Ashmead, 1893 (Mora and Hanson 2019). According to Abe et al. (2013), emerged adults of M. pulchricornis were more melanistic when cocoons were reared at lower temperatures, and the effects of the melanism resulted in increasing body temperatures and improved flight ability of adult M. pulchricornis. Melanism of B. hansoni sp. nov. at high elevations may induce similar outcomes as in M. pulchricornis. Further research is needed when enough live samples are available to confirm this.

The elevation of Costa Rica ranges from sea level to 3,819 m (Hanson and Gauld 1995). If additional sampling is conducted across the country and more species of Bohayella are discovered, altitudinal distribution patterns of members of Costa Rican Bohayella can be investigated in the future (e.g., Aguirre et al. 2018).

Acknowledgements

The first author is grateful to all members of Louisiana State Arthropod Museum and the Department of Entomology as well as LSU Agricultural Center for financial support. We thank Drs Michael Sharkey and James Whitfield for their invaluable help and advice. We also thank Dr Paul Hanson in the Universidad de Costa Rica for the loan of specimens. This study was partially supported by NSF DEB #1841704 to NPL. Research support for SRS was partly provided by National Science Foundation grant DEB 14-42110 (Dimensions of Biodiversity Program). This work was also supported by Wyoming Agricultural Experiment Station funding to SRS provided through the USDA National Institute of Food and Agriculture, McIntire-Stennis project 1021111. Any opinions, findings, and conclusions expressed are those of the authors and do not necessarily reflect the views of the National Science Foundation.

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