Research Article |
Corresponding author: Alejandro Zaldivar-Riveron ( azaldivar@ib.unam.mx ) Academic editor: Michael Sharkey
© 2016 Juan José Martínez, Rubi Nelsi Meza Lázaro, Carlos Pedraza-Lara, Alejandro Zaldivar-Riveron.
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:
Martínez JJ, Lázaro RNM, Pedraza-Lara C, Zaldívar-Riverón A (2016) Sergey gen. n., a new doryctine genus from temperate forests of Mexico and Cuba (Hymenoptera, Braconidae). ZooKeys 589: 143-164. https://doi.org/10.3897/zookeys.589.8291
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The new doryctine genus Sergey gen. n. is described with four new species (S. cubaensis Zaldívar-Riverón & Martínez, sp. n., S. coahuilensis Zaldívar-Riverón & Martínez, sp. n., S. tzeltal Martínez & Zalídivar-Riverón, sp. n., S. tzotzil Martínez & Zalídivar-Riverón, sp. n.) from temperate forests of Mexico and Cuba. Similar to many other doryctine taxa, the new genus has a considerably elongated, petiolate basal sternal plate of the first metasomal tergite, although it can be distinguished from these by having the mesoscutum sharply declivous anteriorly with sharp anterolateral edges. The described species have been characterised molecularly based on two mitochondrial (COI, cyt b) and one nuclear (28S) gene markers. Based on the mitochondrial gene genealogies reconstructed, the evidence suggests the existence of incomplete lineage sorting or hybridization in the populations from Chiapas and Oaxaca assigned to S. tzeltal sp. n.
Doryctinae , Ichneumonoidea , incomplete lineage sorting, new species, taxonomy
The braconid wasp subfamily Doryctinae is a highly diverse, cosmopolitan group that currently comprises 198 genera and about 1,700 species (
One of the main external morphological features that was traditionally used to group genera within the Doryctinae is the relative length of the basal sternal plate of the first metasomal tergite (acrosternite sensu
In a recent molecular phylogenetic study of Notiospathius, various species originally assigned to this genus were nested in two distantly related clades (
In this work, a new doryctine genus, Sergey gen. n., is erected to include the species of the second clade, and four new species are described. Three of these species were collected in cloud forests from México and Cuba, whereas the remaining one was collected in a submontane forest in Coahuila, northeast Mexico. Members of the new genus are morphologically distinct from other doryctine genera with petiolate first metasomal tergite by having the anterolateral corners of mesoscutum sharply pointed and a different pattern of ornamentation in the propodeum, with two divergent carinae that sometimes enclose a more or less distinguishable areola. The phylogenetic relationships within the new genus have been assessed based on separate analyses of one nuclear and two mitochondrial (mt) markers, and provide evidence that suggests the existence of incomplete lineage sorting between two populations of one of the described species.
Specimens were collected in four different localities in Mexico and Cuba, preserved in 100% ethanol, kept at 20 °C until they were processed for DNA sequencing, and subsequently dried, labelled and mounted. The examined specimens are deposited in the Colección Nacional de Insectos,
The morphological terminology follows
Sequences of three gene markers have been examined for specimens belonging to the new genus. These included 34 and 26 sequences that were previously published in
Corrected pairwise genetic distances for the three gene markers were calculated using the K2P model with MEGA version 6 (
Species of this new genus can be distinguished from members of the remaining doryctine genera with long, petiolate first metasomal tergite (e.g. Bolivar, Notiospathius, Pecnobracon Kieffer et Jöergensen, Spathius, Trigonophasmus Enderlein) by having the mesoscutum sharply declivous anteriorly with sharp anterolateral corners. Sergey could be included in the key to dorcytine genera of the New World (
69 (65) | First subdiscal cell of fore wing open at apex, 2cu-a absent, occasionally an infuscate spot or short line present between 2–1A and 2CUa but no distinct vein present (Fig. |
70 |
- | First subdiscal cell closed at apex, 2cu-a present and distinctly meeting 2–1A (Fig. |
75 |
70 (69) | Anterolateral corners of mesoscutum sharply pointed into two flanges, metasoma petiolated | Sergey gen. n. |
- | Anterolateral corners of mesoscutum not sharply pointed anteriorly, the mesoscutum may be sharply raised anteriorly with respect to the pronotum, but flanges are not present, metasoma variable | 70’ |
70’(70) | Most of mesosoma smooth and shining, mesonotum occasionally coriaceous, propodeum occasionally rugose | 71 |
- | Most of mesosoma sculptured, rugose or coriaceous, at most mesopleuron smooth below sternaulus (precoxal sulcus) | 73 |
Head: not depressed. Ocelli arranged in almost equilateral triangle. Frons not distinctly excavated, without a median keel between antennal sockets. Occipital carina complete, fused with hypostomal carina before mandible. Malar suture absent. Clypeus not high, delineated from face by distinct furrow, with fine lower flange. Hypoclypeal depression wide, round. Postgenal bridge narrow. Maxillary palpi 5-segmented, apical segment longer than fourth segment; labial palpi short, 4-segmented, third segment not shortened. Scape of antenna wide and rather short, without flange apically and ventroapical lobe, without basal constriction; ventral margin of scape shorter than dorsal margin in lateral view. First flagellar segment about the same length as second segment, usually several apical or subapical segments whithish. Apical segment more or less pointed apically, without “spine”.
Mesosoma: not depressed. Neck of prothorax short but visible in dorsal view. Pronotum dorsally weakly convex (lateral view), with a transverse carina and a scrobiculate pronotal sulcus. Pronope absent. Propleural dorsoposterior flange rather short. Mesonotum distinctly elevated above pronotum. Anterolateral corners of mesoscutum projected in two flanges (Figs
Wings: veins RS and r-m present, thus first and second submarginal cells entirely closed. Second submarginal cell rather long and narrow. First subdiscal cell open postero-apically, vein 2cu-a absent. Veins 1a and 2a absent. Hind wing with vein C+Sc+R longer than vein SC+R. Vein RS arising from vein R far from vein r-m. Marginal cell more or less distinctly narrowed towards apex, without vein r. Vein cu-a present. Vein M+CU about 0.6-0.7 times as long as 1M; vein m-cu straight. Male hind wing without stigma-like swelling of basal veins.
Legs: Fore tibia on inner surface with several long and slender spines arranged along its anterior margin in almost single vertical line. Hind coxa long and narrow, with basoventral tubercle. Claws simple.
Metasoma: first tergite petiolate, long and narrow, usually striate-coriaceous, with some transverse carinae basally, these carinae sometimes reduced. Basal sternal plate (acrosternite) of first tergite long, 0.6–0.7 times as long as first tergite, extended distinctly beyond level of spiracles. Dorsope of first tergite small and shallow; spiracular tubercles indistinct, situated in basal 0.3 of tergite. Second tergite without distinct furrows and areas. Second suture considerably shallow, complete, almost straight in females and distinctly curved in males. Third tergite without transverse furrow and basal area. Tergites behind second with a single transverse line of sparse long erect setae. Ovipositor distinctly darkened apically, with two distinct subapical nodes. Ovipositor sheaths long, about as long as metasoma or slightly longer.
We are very pleased to name this genus after our dear friend and colleague Dr. Sergey A. Belokobylskij, for his great contribution to the taxonomic knowledge of the braconid subfamily Doryctinae. Gender is to be considered masculine.
Sergey tzeltal sp. n.
1 | Eyes small, their height about as long as malar space (Fig. |
S. coahuilensis sp. n. |
– | Eyes big, their height distinctly longer than malar space (Figs |
2 |
2 | Head and mesoscutum distinctly sculptured, transversally striate (Figs |
S. cubaensis sp. n. |
– | Head and mesoscutum mostly smooth and polished (Figs |
3 |
3 | Antenna with a white apical or subapical band composed of 3-7 (rarely two) flagellomeres in females (Figs |
S. tzeltal sp. n. |
– | Antenna of females with a subapical band only composed of the articulation between the 19th and 20th flagellomeres, five apical flagellomeres brown (Fig. |
S. tzotzil sp. n. |
This is the most distinctive species of the genus. It can be distinguished from the remaining members of Sergey by having the eyes considerably smaller, their height about as long as malar space (distinctly longer than malar space in the remaining species); and the first metasomal tergite broad, 1.5 times longer than its apical width (slender, at least 2.1 times longer than its apical width in the remaining species).
Body length 2.1mm (Fig.
Mesosoma: 2.0 times longer than wide and 1.9 times longer than high (Fig.
Wings: fore wing (Fig.
Legs: fore tibia with a row of spines. Hind coxa slightly striate dorsally, smooth ventrally, with a small but distinct basoventral tubercule.
Metasoma: basal sternal plate/length of first tergum 0.6. First metasomal tergite 2.8 times longer than apically wide (Figs
Male. Unknown.
Known only from a submontane forest at the type locality in Coahuila, Mexico.
Unknown.
The specific epithet refers to Coahuila, the Mexican state where the type locality of this species is located.
Holotype: female (
This distinctive species can be distinguished from the remaining species of Sergey by having: 1) a mostly yellow body colour (brown to black in the remaining species); 2) head and mesoscutum distinctly sculptured, transversally striate (entirely smooth and polished in the remaining species); and 3) fore wing with vein m-cu reaching vein RS+M basally to 2RS, thus vein (RS+M)b present and distinct (m-cu reaching vein RS+M interstitial with respect to vein 2RS, thus vein (RS+M)b absent in the remaining species).
Body length 3.1mm (Fig.
Head: 0.7 times as high as wide in anterior view (Fig.
Mesosoma: about 1.9 times longer than wide and 2.0 times longer than high (Figs
Wings: fore wing length 3.6 mm, length/width ratio 3.7; vein 1cu-a slightly postfurcal to vein 1M, thus vein (RS+M)b present (Fig.
Legs: fore tibia with a row of spines. Hind coxa transversally striate-rugose, with a small but distinct basoventral tubercle..
Metasoma: Basal sternal plate/length of first tergum 0.6. First metasomal tergite 2.5 times longer than apically wide (Fig.
Body length 3.4–4.3 mm. Temple/eye length ratio in dorsal view 0.4–0.5. Antenna with 26–28 flagellomeres. Prescutellar sulcus with four or five carinae. Fore wing length 3.5–3.6 mm, length/width ratio 3.7–3.8 times its maximum width. Ovipositor length 3.5–4.3 mm, 1.8-2.0 times longer than metasoma.
Males. Unknown.
Known only from the type locality in southern Cuba.
Unknown.
This species is named after the Caribbean country where it occurs, Cuba.
Holotype (
This species is similar to S. tzotzil, but it can be distinguished from the latter species by the colour pattern of the white band on the female antenna. In S. tzeltal, the white band is either apical or subapical and is composed of at least two entire whitish flagellomeres, usually more, with at most three apical flagellomeres brown. In S. tzotzil, the white band is subapical and consists only of the lighter color on the articulation between the 19th and 20th flagellomeres, and with the five apical flagellomeres brown.
Body length 3.1 mm (Fig.
Head: 0.8 times as high as wide in anterior view (Fig.
Mesosoma: 2.0 times longer than wide (Fig.
Wings: fore wing (Fig.
Legs: fore tibia with a row of spines. Hind coxa transversally striate dorsally, smooth ventrally, with a distinct basoventral tubercule.
Metasoma: Basal sternal plate/length of first tergum 0.6. First metasomal tergite 2.2 times longer than apically wide (Fig.
Body length 2.6–3.8 mm. Temple/eye length ratio in dorsal view 0.16–0.33. Antenna with 22-27 flagellomeres, white subapical band composed of two to four flagellomeres. In smaller specimens, rugose median area of mesoscutum reduced, though notauli never clearly distinguishable at posterior edge of mesoscutum, obscured among rugosities. Prescutellar sulcus sometimes with para-median carinae reduced, thus only the median carina is clearly distinguishable. Fore wing length 2.2–2.9 mm, length/width ratio 2.9–3.9 times its maximum width. Veins 2RS/2M ratio 0.5–0.55. Basal sternal plate 0.53–0.68 times length of first metasomal tergum. Ovipositor length 2.0–2.1 mm.
Body length 2.5–3.5 mm. Malar space 0.28. Temple/eye length ratio (dorsal view) 0.29–0.39. Flagellomeres 21–26 either entirely brown (Chiapas) (Fig.
This species is known from cloud forests located in the Reserva el Triunfo, Chiapas, and Santiago Comaltepec, Oaxaca, in southeast Mexico.
Unknown.
This species has a considerable variation in the antennal color pattern. We had originally grouped the specimens assigned to this taxon in two morphospecies, each represented by the specimens from Chiapas and Oaxaca, respectively. Females from Oaxaca have a distinct apical white band composed of 6–8 flagellomeres (Fig.
There was no concordance between the corrected COI distances and the geographic provenance and morphological variation for the above specimens. Some of the specimens from Oaxaca had lower COI distances with those from Chiapas than with the remaining specimens from the same locality (0.38–0.76 and 1.7–1.9%, respectively). This incongruence suggests that the existence of incomplete lineage sorting or hybridization between two recently diverged, sympatric species (see below). We have followed a conservative approach and consider the members of the populations from Chiapas and Oaxaca as a single species. One of the specimens from Oaxaca (DNA voucher number CNIN573) has considerably higher COI distances compared with the remaining conspecific specimens (3.6–4.7%). However, it is morphologically undistinguishable and we thus placed it within S. tzeltal.
The name of this species refers to the Tzeltal ethnic group, descendant from the Mayans that inhabits Los Altos, a mountain region located in central Chiapas
Holotype (
See diagnosis of S. tzeltal.
Body length 3.7mm (Fig.
Head: in anterior view 0.9 times as high as wide (Fig.
Mesosoma: 2.1 times longer than wide (Fig.
Wings: Fore wing length 2.9 mm, length/width ratio 3.3; vein 1cu-a slightly postfurcal to vein 1M; veins 2RS/2M ratio 0.5.
Legs: Fore tibia with a row of spines. Hind coxa transversally striate dorsally, smooth ventrally, with a distinct basoventral tubercle.
Metasoma: Basal sternal plate/length of first tergum 0.6. First metasomal tergite 2.1 times longer than apically wide (Fig.
Males. Similar to female, slightly smaller and with antenna uniformly brown.
Known only from the type locality in El Triunfo, Chiapas, Mexico.
Unknown.
This species and S. tzeltal were collected in the same locality in Chiapas.
The name of this species refers to the Tzotzil ethnic group, descendant from the Mayans, who inhabits the Altos, a mountain region located in central Chiapas.
Holotype (
Intraspecific corrected genetic divergences varied from 0 to 2.1 (excluding CNIN573), 0.27 to 3.18 and 0 to 0.33% for COI, cyt b and 28S, respectively. Interspecific distances within Sergey on the other hand ranged from 7.99 to 15.28, 12.64 to 13.6 and 0.17 to 0.5% for COI, cyt b and 28S, respectively.
The Bayesian phylograms derived from the separate COI and cyt b analyses are included in the Figure
The bayesian phylogram derived from the cyt b sequences yielded similar relationships with the COI topology. Again, some of the specimens of S. tzeltal from Comaltepec, Oaxaca were more closely related to the ones from El Triunfo, Chiapas (PP = 0.6) than with the remaining specimens from the same locality. The 28S tree was largely unresolved (phylogram not shown), with the sequenced specimens S. tzeltal and S. tzotzil grouped together (PP = 1.0). The reconstructed mt gene genealogies, together with the geographic provenance and morphological variation found in the specimens of S. tzeltal from Oaxaca and Chiapas suggests that this taxon could consist of two sympatric, recently derived lineages in which there is incomplete lineage sorting or hybridization. Further morphological and genetic studies will help to confirm the taxonomic status of the populations of S. tzeltal from the latter two Mexican regions.
We thank Marysol Trujano for collecting the specimen of S. coahuilensis; Mike Sharkey for providing the two specimens of S. cubaensis, C. Mayorga and G. Ortega for their help with the curation of specimens, Andrea Jiménez and Laura Márquez for their help in the laboratory, S. Guzman for her help taking the digital pictures, and Mario García for his help during AZR’s sabbatical stay at the