Research Article |
Corresponding author: Angélica M. Penteado-Dias ( angelica@ufscar.br ) Academic editor: Jose Fernandez-Triana
© 2021 Flávia R. Joele, Alejandro Zaldívar-Riverón, Angélica M. Penteado-Dias.
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:
Joele FR, Zaldívar-Riverón A, Penteado-Dias AM (2021) Six new species of Allorhogas (Hymenoptera, Braconidae, Doryctinae) from south and southeast Brazil with host-plant record. Journal of Hymenoptera Research 82: 199-220. https://doi.org/10.3897/jhr.82.62345
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Six new Brazilian species of the gall-associated Doryctinae genus Allorhogas are described and illustrated: A. copaiba sp. nov., A. ilexaffinis sp. nov., A. inquilinus sp. nov., A. quarentenus sp. nov., A. vassununga sp. nov. and A. viridis sp. nov. We provide host plant records for five of these species, three and one of which are new host plant genera (Ilex L., Copaifera L. and Eugenia P. Micheli ex L.) and new host plant family (Aquifoliaceae) records, respectively. Allorhogas inquilinus sp. nov., whose biology was previously reported, represents the first confirmed case of phytophagous inquilinism in the genus. An updated key to Brazilian species of Allorhogas is provided.
Gall-association, Ichneumonoidea, inquilinism, Neotropics, phytophagous
Allorhogas
The Doryctinae is mostly represented by parasitoid species. However, Allorhogas belongs to a clade composed of nine genera whose species are associated with members of various vascular plant families (
Recent molecular phylogenetic studies have shown that Allorhogas is polyphyletic (
The type material of the six species described here was obtained after examination of specimens deposited in the Coleção Taxonômica do Departamento de Ecologia e Biologia Evolutiva (
This new species and A. inquilinus sp. nov. (described below) are the only two known species of Allorhogas from Brazil with vertex and frons rugose. However, A. copaiba sp. nov. can be distinguished from the latter and the remaining species of Allorhogas from this country by having the following combination of morphological features: 1) frons and vertex striate-rugose and coriaceous (strongly rugose-coriaceous in A. inquilinus sp. nov., usually coriaceous in the remaining species); 2) first metasomal tergite longitudinally costate-rugose and coriaceous (generally longitudinally costate-coriaceous in the remaining species); and 3) vein 2RS distal with m-cu (interstitial with m-cu in A. inquilinus sp. nov., variable in the remaining species).
Female. Body size 2.9 mm (Fig.
Head: transverse in dorsal view, 2.1 times wider than its median length (dorsal view) (Fig.
Mesosoma: 1.4 times longer than high (Fig.
Wings: forewing 3.0 times longer than wide (Fig.
Legs: hind coxa with a distinct basoventral tooth. Hind femur 3.0 times longer than wide.
Metasoma: first tergite 1.3 times wider than long, longitudinally costate-rugose and coriaceous; anteriorly delimited by an indistinct transverse carina, with two parallel longitudinal carinae running to apex (Fig.
Body size 2.8–3.0 mm. Antenna with 24–27 flagellomeres.
Male. Similar to female. Body size 2.8–3.0 mm. Antenna with 24–27 flagellomeres. Hind femur swollen, 2.6 times longer than wide. Prescutellar furrow with three transverse carinae.
Female (
(
The type specimens of A. copaiba were reared from spherical galls on stems and buds of Copaifera langsdorffii Desf. (Fabaceae) (Fig.
A spherical gall on a stem of Copaifera langsdorffii Desf. (Fabaceae) from where A. copaiba sp. nov. was reared B A. ilexaffinis sp. nov., head, frontal view C A. quarentenus sp. nov., head, frontal view D A. inquilinus sp. nov., head, frontal view E A. vassununga sp. nov., head, dorsal view F A. viridis sp. nov., head, frontal view.
The name of this species refers to its association with the vascular plant genus Copaifera, whose common name in Brazil is Copaíba.
This new species could be distinguished from the remaining species of Allorhogas from Brazil by having the following combination of features: 1) mesosoma brown to dark brown; 2) first and second metasomal tergites costate, smooth between carinae, remaining tergites smooth and polished (first and second tergites costate but with coriaceous sculpture between carinae and third tergite also sculptured in other Brazilian species with dark brown colour).
Female. Body size 2.7 mm (Fig.
Head: transverse in dorsal view, 1.5 times wider than its median length (dorsal view) (Fig.
Mesosoma: 1.6 times longer than high (Fig.
Wings: forewing 2.7 times longer than wide (Fig.
Legs: hind coxa with a distinct basoventral tooth. Hind femur 3.2 times longer than wide.
Metasoma: first tergite 1.3 times wider than long, longitudinally costate, smooth between carinae; with two subparallel longitudinal carinae running to apex, anteriorly delimited by an indistinct transverse carina (Fig.
Body size 2.7–2.9 mm. Head and mesosoma brown to dark brown. Antenna with 25–26 flagellomeres.
Male. Similar to female. Body size 2.6–2.7 mm. Antenna with 28 flagellomeres. Hind femur swollen, 2.6 times longer than wide.
Female (
(
The type specimens of this species were reared from fruits of Ilex affinis Gardner (Aquifoliaceae).
The name of this species refers to the host plant species where the type specimens were reared.
This is a morphologically distinctive species, which could be distinguished from the remaining described members of Allorhogas from Brazil by having: 1) eyes small, malar space 0.8 times eye height (eyes larger, malar space less than 0.8 times eye height in the remaining species); 2) frons and vertex strongly rugose-coriaceous (generally coriaceous in the remaining species; striate-rugose and coriaceous in A. copaiba); 3) face coriaceous with longitudinal v-shape striation (variable but never with this sculpture in the remaining species); 4) propodeum entirely strongly rugose-areolate, with two indistinct diverging carinae (at least partially coriaceous and with distinct diverging carinae in the remaining species); 5) hindwing with vein m-cu almost straight, slightly curved towards wing base (slightly curved towards apex in the remaining species); 6) apical half of third and remaining metasomal tergites punctate-slightly costate (usually smooth in the remaining species).
Female. Body size 3.3 mm (Fig.
Head: transverse in dorsal view, 2.4 times wider than its median length (dorsal view) (Fig.
Mesosoma: 1.6 times longer than high (Fig.
Wings: forewing 3.2 times longer than wide (Fig.
Legs: hind coxa with distinct, pointed basoventral tooth, with large, dense pilosity, ventrally rugose. Hind femur 3.3 times longer than wide.
Metasoma: first tergite 1.7 times wider than long, costate, slightly punctate laterally, with two longitudinal carinae only distinct at base; transverse basal carina distinct (Fig.
Body size 3.0–4.0 mm. Antenna with 28–31 flagellomeres.
Male. Similar to female. Body size 4.0 mm. Antenna with 30 flagellomeres. Hind femur swollen, about 3.0 times longer than wide.
Female (
(DCBU420453-58, 420472; IBUNAM). 1 male, 4 females, 1 specimen without metasoma (
A detailed description of the feeding biology of A. inquilinus was previously reported by
The epithet of this species is a reference to its inquiline biology.
This new species is morphologically similar to the following species that also have been associated with legumes, with the first being described from Costa Rica and the remaining four from Brazil: A. infuscotarsus Marsh, A. brasiliensis Marsh, A. mineiro Zaldívar-Riverón & Martínez, A. spermaphagus Marsh and A. vulgaris Zaldívar-Riverón & Martínez. Allorhogas quarentenus can be distinguished from the latter five species by its dark brown tarsi (only hind tarsus brown in A. infuscotarsus; yellow to honey yellow in the remaining species). Moreover, it can be distinguished from A. infuscotarsus and A. mineiro by having its ovipositor sheaths 0.6 times as long as metasoma (0.33 and 1.1–1.3 in the latter species, respectively); from A. brasiliensis by having 24–27 flagellomeres and forewing m-cu interstitial with 2RS (32–32 flagellomeres and forewing m-cu basal to 2RS in A. brasiliensis); and from A. spermaphagus by having the propodeum basally coriaceous (rugulose-coriaceous in A. spermaphagus).
Female. Body size 3.5 mm (Fig.
Head: transverse in dorsal view, 1.8 times wider than its median length (dorsal view) (Fig.
Mesosoma: 1.8 times longer than high (Fig.
Wings: forewing 2.8 times longer than wide (Fig.
Legs: hind coxa with distinct, pointed basoventral tooth. Hind femur 3.7 times longer than wide.
Metasoma: first tergite 1.6 times wider than long, longitudinally costate-rugulose and coriaceous, with costate carinae partially indistinct basally, smooth medio-apically, with two distinct longitudinal subparallel carinae; basally delimited by a transverse carina (Fig.
Body size 3.1–3.8 mm. Metasoma honey yellow to yellow. Most part of lateral mesoscutal lobes dark brown (four specimens). Fore and middle tarsi honey yellow, brown or dark brown. Antenna with 24–27 flagellomeres. Prescutellar furrow with 2–3 transverse carinae.
Male. Similar to female. Body size 2.7–3.1 mm. Most part of lateral mesoscutal lobes dark brown (one specimen). Antenna with 25–27 flagellomeres. Hind femur swollen, 2.3–2.9 times longer than wide.
Female (
(
The type specimens of A. quarentenus were reared from legumes of an undetermined species of Inga.
The name of this species refers to the COVID-19 pandemics with its subsequent undefined quarantine, which occurred while the authors were describing it.
Specimens of A. quarentenus and A. vulgaris could be either uniformly honey yellow or with dark brown areas. However, in the former species they are only present along the lateral mesoscutal lobes (also present in the basal areas of propodeum and central areas of second to fourth tergites in A. vulgaris).
This new species can be distinguished from the remaining described species of Allorhogas from Brazil by having the following combination of morphological features: 1) first metasomal tergite longitudinally costate-rugose, with two longitudinal carinae only distinct at base (sculpture variable, often with longitudinal carinae running along the entire tergite in the remaining species); 2) frons excavation defined by sharp lateral margins and a median longitudinal carina (frons excavation with or without sharp lateral margins and median longitudinal carina in the remaining species); 3) mesosoma and metasoma mostly dark brown to black, mesopleuron medially and posteriorly and sixth to remaining tergites honey yellow (always with a different colour pattern in the remaining species).
Female. Body size 3.4 mm (Fig.
Head: slightly transverse in dorsal view, 2.0 times wider than its median length (dorsal view) (Figs
Mesosoma: 1.9 times longer than high (Fig.
Wings: forewing 2.7 times longer than wide (Fig.
Legs: hind coxa with distinct, pointed basoventral tooth. Hind femur 4.1 times longer than wide.
Metasoma: first tergite 1.3 times wider than long, longitudinally costate-rugose, with two longitudinal carinae only distinct at base, anteriorly delimited by a transverse carina (Fig.
Body size 3.0–3.4 mm. Antenna with 28–30 flagellomeres.
Male. Similar to female. Body size 3.6 mm. Antenna with 29 flagellomeres. Hind femur swollen, about 3.6 times longer than wide.
Female (
(
Unknown.
The name of this new species refers to the locality where the type specimens were collected, Parque Estadual de Vassununga, in the state of São Paulo, Brazil.
This new species could be distinguished from the remaining described species of Allorhogas by having the following combination of morphological features: 1) body greenish pale yellow (never greenish pale yellow in the remaining species); 2) claws white (dark in the remaining species); 3) anterior half of notauli strongly scrobiculate and distinct, posterior half less sculptured and distinct, not meeting, reaching the end of scutellum in a smooth-rugose area with two subparallel longitudinal carinae (notauli usually entirely distinct, without such longitudinal carinae in the remaining species); 4) vein 2RS distal with m-cu, vein RS+Mb absent (variable in the remaining species).
Female. Body size 2.9 mm (Fig.
Head: transverse in dorsal view, 1.7 times wider than its median length (dorsal view) (Fig.
Mesosoma: 1.7 times longer than high (Fig.
Wings: forewing 3.0 times longer than wide (Fig.
Legs: hind coxa coriaceous, striate ventrally with distinct, pointed basoventral tooth. Hind femur 3.2 times longer than wide.
Metasoma: first tergite 0.9 times as wide as long, longitudinally costate, with two indistinct longitudinal subparallel carinae, with a basal transverse carina (Fig.
Body size 2.5–2.9 mm. Prescutellar furrow with 4–5 transverse carinae.
Male. Similar to female. Body slightly darker in one specimen. Body size 2.8–2.9 mm. Antenna with 22–23 flagellomeres. Hind femur swollen, 2.2–2.6 times longer than wide.
Female (
(
The type specimens of A. viridis were reared from conical galls apparently made by an undetermined cecidomyiid dipteran on leaves of Eugenia rotundifolia Casar. (Myrtaceae). These galls are cylindrical, unilocular, externally brown and internally white (
The epithet of this species derives from the Latin word viridi (green), in reference to its distinctive greenish colour.
1 | Mesosoma compact, 1.2–1.5 times longer than high; pronotal collar short, not visible in dorsal view; propodeum almost entirely oblique in lateral view (associated to Melastomataceae fruits or floral buds) | 2 |
– | Mesosoma longer, at least 1.7 times longer than high; propodeum not oblique in lateral view | 4 |
2 (1) | Mesoscutal lobes coriaceous, without rugose areas surrounding notauli; hind femur and tibia brown | A. granivorus Zaldívar-Riverón & Martínez |
– | Mesoscutal lobes coriaceous, with rugose areas surrounding notauli; hind legs entirely honey yellow | 3 |
3 (2) | Ovipositor length about 0.2 times as long as metasoma; antenna with 20–21 flagellomeres; head, mesosoma and metasoma mostly dark brown, with some black areas | A. uberlandiensis Joele & Zaldívar-Riverón |
– | Ovipositor length about 0.5 times as long as metasoma; antenna with 22–25 flagellomeres; head, mesosoma and metasoma black | A. clidemiae Martínez & Zaldívar-Riverón |
4 (2) | Ovipositor shorter than first metasomal tergite | 5 |
– | Ovipositor longer, at least 0.5 length of metasoma | 6 |
5(4) | Malar space 2/5 eye height | A. dyspistus Marsh |
– | Malar space half as long as eye height | A. muesebecki (Guimarães) |
6(4) | Vertex and frons with marked rugose sculpture | 7 |
– | Vertex and frons mainly coriaceous, without distinct rugose sculpture | 8 |
7(6) | Frons and vertex striate-rugose and coriaceous; vein 2RS distal with m-cu | A. copaiba sp. nov. |
– | Frons and vertex strongly rugose-coriaceous; vein 2RS interstitial with m-cu | A. inquilinus sp. nov. |
8(6) | Body greenish pale yellow, tarsal claws white | A. viridis sp. nov. |
– | Body not greenish pale yellow, tarsal claws dark | 9 |
9(8) | Forewing m-cu arising distal to 2RS | A. heringeri (Guimarães) |
– | Forewing m-cu arising basal or interstitial to 2RS | 10 |
10(9) | Forewing m-cu arising basal to 2RS, thus vein RS +Mb distinct | 11 |
– | Forewing vein m-cu interstitial to 2RS, RS +Mb not distinguishable | 12 |
11(10) | Body length 4.0–4.5 mm, 32–33 flagellomeres | A. brasiliensis (Marsh) |
– | Body length 2.9–3.1 mm, 25–28 flagellomeres | A. vulgaris Zaldívar-Riverón & Martínez |
12(10) | Body mostly dark brown to black, with some areas brown to honey yellow | 13 |
– | Body honey yellow to light brown | 14 |
13(12) | First metasomal tergite with two longitudinal carinae only distinct at base; frons excavation defined by sharp lateral margins and a median longitudinal carina | A. vassununga sp. nov. |
– | First metasomal tergite with two subparallel longitudinal carinae running to apex; frons excavation distinct but not defined by sharp lateral margins, without a median longitudinal carina | A. ilexaffinis sp. nov. |
14(12) | Forewing vein r 0.7 times as long as 3RSa | A. taua Penteado-Dias & Carvalho |
– | Forewing vein r about as long as vein 3RSa | 15 |
15(14) | Tarsi dark brown | A. quarentenus sp. nov. |
– | Tarsi entirely yellow | 16 |
16(15) | Ovipositor sheaths 1.1–1.3 times the length of metasoma | A. mineiro Zaldívar-Riverón & Martínez |
– | Ovipositor sheaths 0.7 times the length of metasoma | A. spermaphagus Marsh |
The conserved external morphology in Allorhogas makes rearing records an important diagnostic feature to distinguish its species, since they appear to be highly specific to their host plants. This study increases to 17 the number of described species of Allorhogas from Brazil, and to 12 the number of host plant families that are associated with this genus. The plant association reported for A. ilexaffinis to fruits of Ilex affinis represents the first record for a member of the plant family Aquifoliaceae. Moreover, Copaifera L. (Fabaceae) and Eugenia P. Micheli ex L. (Myrtaceae), which are associated with A. copaiba and A. viridis, respectively, are new host genus records for Allorhogas.
Of the six species described here, only the feeding biology of A. inquilinus could be confirmed. This represents the first confirmed record of a gregarious inquiline species in Allorhogas, and the only described species that is associated with the plant family Anacardiaceae. The galls where the type specimens of A. copaiba were reared also contained an undetermined lepidopteran moth species, and thus we presume that this could be the actual gall former in the system. Interestingly, A. inquilinus and A. copaiba are morphologically similar, sharing some unique features within the genus (e.g. vertex and frons with distinct rugose sculpture). Allorhogas viridis was, on the other hand, reared from leaf galls made by an undetermined cecidomyiid dipteran, though its feeding biology remains unknown.
Of all the previously proposed morphological synapomorphies of Allorhogas (
We especially thank Gilson R. P. Moreira for donating the type material of A. inquilinus; we also thank Luciana Bueno dos Reis Fernandes for taking the SEM and stereo microscope digital photographs at DEBE/UFSCar; Cristina Mayorga, Guillermina Ortega and Susana Guzmán for their help with the curation of the type specimens at the IBUNAM. This work was funded by a grant given by CNPq, CAPES and FAPESP (project INCT-HYMPAR) to AMPD.