Citation: Fernández-Triana JL, Whitfield JB, Smith MA, Braet Y, Hallwachs W, Janzen DH (2014) Review of the Neotropical genus Prasmodon (Hymenoptera, Braconidae, Microgastrinae), with emphasis on species from Area de Conservación Guanacaste, northwestern Costa Rica. Journal of Hymenoptera Research 37: 1–52. doi: 10.3897/JHR.37.6748
The Microgastrinae genus Prasmodon is revised and the following 16 new species are described: erenadupontae and verhoogdenokus (authored by Braet and Fernández-Triana), and almasolisae, aureus, bobpoolei, bobrobbinsi, dondavisi, johnbrowni, masoni, mikepoguei, nixoni, paulgoldsteini, scottmilleri, silvatlanticus, subfuscus, and tijucaensis (authored by Fernández-Triana and Whitfield). The greatest species richness is found in the Amazon basin, but the genus extends throughout the rain forests of Central and South America. Leaf-rolling and webbing caterpillars of Crambidae and Elachistidae are the only known hosts for these parasitoids.
Microgastrinae, Prasmodon, parasitoid biology, Neotropics, Area de Conservación Guanacaste, Costa Rica, species descriptions, caterpillars
The subfamily Microgastrinae (Hymenoptera: Braconidae) is a species-rich and taxonomically difficult lineage of parasitoid wasps that attack caterpillars (Lepidoptera), and also one of the main taxa used in the biological control of caterpillar pests in forestry and agriculture (
The last two decades have seen an increase in the study of microgastrine wasps in the Neotropics – as a result of the publication of a key to New World genera (
This paper, a continuation of those studies, deals with Prasmodon. This genus was described by
Since Nixon’s original description of Prasmodon, little progress has been made in determining its relationships with other microgastrine genera.
Most of the specimens used in this study came from ACG. They were reared (usually, see http://janzen.sas.upenn.edu for details for any particular specimen) or Malaise-trapped, and host caterpillars and other ecological information are considered along with DNA barcodes (COI gene) when available (Fig. 105). Additionally, a collection of Prasmodon specimens from French Guiana was available (personal collection of YB, collected during several field expeditions and also obtained through collaborations with various people living there). We also studied the South American specimens of Prasmodon in the Canadian National Collection of Insects (CNC) in Ottawa, Canada. The non-ACG material did not have associated host or molecular data, with the exception of a few, mostly short DNA barcodes (usually 160 bp) from some CNC specimens. Altogether 140+ specimens were studied.
The specimens of the new species are deposited in the CNC, Natural History Museum, London, United Kingdom (BMNH), the Illinois Natural History Survey, Champaign, Illinois, United States (INHS), the Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica (INBio), the National Museum of Natural History, the Smithsonian Institution, Washington DC, United States (NMNH), Naturalis Biodiversity Center Leiden, the Netherlands, (RMNH), Institut Royal des Sciences Naturelles de Belgique Brussels, Belgium (IRSNB), and Museum National d’Histoire Naturelle, Paris, France (MNHN).
Morphological terms and measurements of structures are mostly as used by
The new species descriptions are based on the holotype female, with other specimens studied (when available) for intraspecific variation.
Lucid 3.5.4 (http://www.lucidcentral.com/) software was used to generate automatic descriptions of the species and to prepare Lucid identification keys. A dataset of 22 characters and 90 character-states was used to provide uniform descriptions for all species treated. The description format includes one sentence per character, with the character mentioned first and the character-state following after a colon, e.g., “Scape color: entirely yellow”. Whenever a species scored more than one character-state, the description included all of the pertaining character-states separated by “or”, e.g., “Scape color: entirely yellow or partially brown to black”. Whenever a character-state was coded as uncertain due to poor condition of the specimen(s), the description includes the details of the character-state as best assessed, followed by a question mark, e.g., “Scape color: entirely yellow (?)”. Sometimes a character could not be coded due to missing body parts in the available specimens; in such instances the feature was left out of the description for that particular species.
In most cases we used a simplified convention to code color, considering it as either pale (white, light yellow, orange-yellow, light brown-yellow) or dark (dark brown, black). For details on the exact color patterns on the body, we provide extensive illustrations for every species.
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.
A map with the distribution of all species was generated using SimpleMappr (
DNA barcodes for all ACG inventory Microgastrinae were obtained using DNA extracts prepared from single legs using a glass fibre protocol (
In the taxonomic treatment of species, “Specimens Examined” presents the specimen’s information in the following format: “Number of females/males examined, acronym of the storing institution between parenthesis, COUNTRY: State/Province, Region, city, other locality details, latitude/longitude coordinates (in decimal degrees), collection date, collector name, biological information on host (starting with “ex”), ACG database codes (in the format “yy-SRNP-xxxxxx” for the host caterpillar or parasitoid lot reared from it, or “DHJPARxxxxxxx” for an individual parasitoid specimen).
We describe here 16 new species from seven countries (Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Peru and Suriname); increasing the number of Prasmodon species from two to 18. There are at least three additional species (one each from Brazil, Costa Rica and Ecuador) in the CNC collection, only represented by 1-2 males each, with no molecular or biological data associated. Those species will remain undescribed until more specimens, especially females, become available. Males that cannot be associated with females have usually proven to be almost useless in Microgastrinae taxonomy (e.g.,
The distribution of species by country and by major bioregion shows some interesting patterns (Fig. 104). We found 10 species of Prasmodon in Costa Rica, out of roughly 140 rearings and 8 Malaise-trapped specimens from ACG. Seven species were recorded from Brazil, out of some 30 Malaise-trapped specimens. Ecuador (6 specimens), French Guiana (20 specimens), and Peru (3 specimens) had three species each. Colombia (3 specimens) and Suriname (1 specimen) had one species each.
Altogether, Central America (only known specimens are from ACG) had 10 of the total species found (55.6%), while South America had nine species (50 %). Prasmodon eminens was the only species shared between Central and South America (Fig. 104), but even that might be an artifact because that species is thought to actually comprise a complex of morphologically cryptic species (e.g.
As for major bioregions, ACG had 10 species, sharing one of them with the Amazonia bioregion (for simplicity, we consider here ACG as one region altogether, even though it is a mosaic of ecosystems, just as the other so-called “bioregions”). Amazonia had six species, one shared with ACG, and three shared with the Guyana lowland rainforests (which has no endemic species on its own). The Atlantic Forest of Brazil had three species, all endemic to that region (Fig. 104).
The ACG Prasmodon specimens have been collected in relatively low-elevation areas (95–980 m), in intact rain forest or secondary vegetation where there used to be intact rain forest 0–100 years earlier, except for Prasmodon paulgoldsteini, which has been found (6 Malaise-trapped specimens) only in ACG dry forest at about 300 m elevation. Based on the data currently available, Prasmodon is largely a genus of Neotropical rain forests. Although the most species-rich area at present is Central America (ACG), we speculate that further collecting will reveal the highest diversity to be actually found in South America.
In a first analysis of the biology of the genus,
This is one of the most recognizable genera of Microgastrinae (
[Throughout the key the acronyms T1, T2, etc., are used for morphological terms mediotergite 1, 2, etc.; and F refers to flagellomere number, e.g. F12]
| 1 | Mesosoma and metasoma entirely yellow-orange dorsally (Figs 5, 6, 11, 12, 17, 18, 27, 28, 32, 34, 38, 39, 44, 45, 56, 58, 92, 93), except for very small black spot on axillar complex (as in Fig. 5) | 2 |
| – | Mesosoma and/or metasoma with some brown coloration dorsally (Figs 23, 50, 51, 63, 69, 75, 80, 81, 86, 87, 94, 95), in addition to very small black spot on axillar complex | 10 |
| 2(1) | Apical flagellomeres yellow or yellowish-white, contrasting with rest brown to black (Figs 3, 13, 30, 54, 90) | 3 |
| – | All flagellomeres brown to black (as in Figs 11, 12, 24, 40) | 7 |
| 3(2) | Antenna mostly dark brown to black; with small, apical area (F15–16, and occasionally apical half of F14) which is yellow-brown, lighter in colour than remaining flagellomeres (Figs 3, 13) | 4 |
| – | Antenna with yellow area much larger (at least including F12–15, and usually apical half of F11 and basal half of F16), much lighter compared to remaining flagellomeres (Figs 30, 54) | 5 |
| 4(3) | Metatarsus almost entirely yellow-white (Fig. 13); posterior 0.1–0.2 of mesotibia dark brown to black; body length 4.2–4.3 mm (X=4.3 mm), rarely 4.5 mm; fore wing length 4.5–4.6 mm (X=4.6 mm), rarely 4.8 mm; T1 5.6 × as long as width at posterior margin [Hosts: Elachistidae, mostly Anteotricha feeding on Bignoniaceae] | Prasmodon bobpoolei Fernández-Triana & Whitfield, sp. n. |
| – | Metatarsus dark brown to black (except for anterior 0.7 of first metatarsomere) (Fig. 1); mesotibia entirely yellow; body length 4.5–4.9 mm (X=4.8 mm); fore wing length 5.0–5.3 mm (X=5.1 mm); T1 4.0 × as long as width at posterior margin [Hosts: Elachistidae, mostly Antaeotricha feeding on Meliaceae] | Prasmodon almasolisae Fernández-Triana & Whitfield, sp. n. |
| 5(3) | Fore wing with a pale area centrally (Figs 30, 33) which gives the wing a banded appearance; anterior 0.4–0.5 of vein 3M (and usually part or all of veins 2M and r-m) yellow-orange, contrasting with all other veins which are dark brown (Fig. 33); scutoscutellar sulcus with 4 impressions (Fig. 32); T1 distinctly narrowing at around 0.5 its length, with both anterior and posterior margins clearly wider than its median width (Fig. 34) | Prasmodon erenadupontae Braet & Fernández-Triana, sp. n. |
| – | Fore wing either mostly hyaline, or entirely infumate (except for small hyaline area near veins (RS+M)b and 2M); all veins dark brown (Figs 53, 89); scutoscutellar sulcus usually with more than 5 impressions (Figs 56, 92); T1 distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Figs 58, 89, 93) | 6 |
| 6(5) | Hypostomal carina not raised (e.g. Figs 16, 57); hind wing subbasal cell mostly setose (Fig. 44); T1 less than 5.0 × as long as width at posterior margin; T2 5.0 × as wide at posterior margin as median length | Prasmodon nixoni Fernández-Triana & Whitfield, sp. n. |
| – | Hypostomal carina highly raised (Figs 9, 25, 31); hind wing subbasal cell mostly without setae (Fig. 39); T1 6.0 × as long as width at posterior margin; T2 less than 3.0 × as wide at posterior margin as median length | Prasmodon verhoogdenokus Braet & Fernández-Triana, sp. n. |
| 7(2) | Ovipositor sheaths relatively short, 0.3 × as long as metatibia length (Fig. 43); all legs entirely yellow (Fig. 40) | Prasmodon masoni Fernández-Triana & Whitfield, sp. n. |
| – | Ovipositor much longer, 0.5–0.6 × as long as metatibia length (Figs 7, 24, 35); at least posterior 0.1–0.2 of metatibia and most of metatarsus (and sometimes parts of middle leg) dark brown to black (Figs 7, 26, 35, 37) | 8 |
| 8(7) | Scape entirely yellow (Fig. 9); fore wing with most veins golden yellow, pterostigma mostly yellow, with central part brown (Figs 8, 10) [Ecuador] | Prasmodon aureus Fernández-Triana & Whitfield, sp. n. |
| – | Scape partially dark brown to black (Figs 25, 35, 38); fore wing with most veins and pterostigma entirely dark brown (Figs 24, 36) [Costa Rica, ACG] | 9 |
| 9(8) | Posterior 0.1–0.2 of mesotibia, apical segment of mesotarsus, posterior 0.2–0.3 of metatibia, and metatibia spurs, dark brown to black (Figs 35, 37); scutoscutellar sulcus with 5–6 impressions (Fig. 38); hind wing with subbasal cell mostly without setae (Fig. 39); T1 evenly narrowing towards posterior margin, its length 6.3 × as width at posterior margin (Fig. 39) [Hosts: Crambidae, Asturodes fimbriauralis DHJ02, Eulepte alialis, Piletosoma thialis, Phostria Janzen05, feeding on many plant families] | Prasmodon johnbrowni Fernández-Triana & Whitfield, sp. n. |
| – | Mesotibia, mesotarsus, and metatibia spurs entirely yellow-orange (Figs 24, 26); posterior 0.1–0.2 of metatibia dark brown to black; scutoscutellar sulcus with 4 impressions (Fig. 27); hind wing with subbasal cell mostly setose (although it may be a small area without setae near lower margin of wing, as in Fig. 28); T1 more or less parallel-side until half its length, then slightly narrowing towards posterior margin (Figs 27, 28), its length 3.3–3.7 × as width at posterior margin [Hosts: Crambidae, Prenesta feeding on Apocynaceae: Prenesta Janzen196, Prenesta Janzen195, Prenesta scyllalis DHJ03] | Prasmodon dondavisi Fernández-Triana & Whitfield, sp. n. |
| 10(1) | Only metasoma with some brown coloration on mediotergites; mesosoma completely yellow-orange (Figs 22, 23, 62, 63, 68, 69, 73, 75, 95) | 11 |
| – | Both mesosoma and metasoma with brown to black areas (Figs 50, 51, 80, 81, 86, 87, 94) | 15 |
| 11(10) | Metasoma almost completely yellow-orange (Fig. 95), with only small brown areas centrally on mediotergites 4–7 (usually only on mediotergires 5–6); larger species, body length 5.0–5.3 mm and fore wing length 5.2–5.6 mm | Prasmodon eminens Nixon, 1965 |
| – | Metasoma with brown to black areas covering most of mediotergites 4–7, and also part of mediotergites 3 and 8 (Figs 23, 63, 69, 75); smaller species, body length 3.8–4.0 mm and fore wing length 4.0–4.6 mm | 12 |
| 12(11) | All coxae, pronotum, mesopleuron, metapleuron, and propodeum white to very light yellow, contrasting with rest of mesosoma yellow to orange-yellow (Figs 64, 68, 69); T1 3.7 × as long as width at posterior margin. [Hosts: Crambidae, Omiodes, Triuncidia, feeding on many plant families] | Prasmodon scottmilleri Fernández-Triana & Whitfield, sp. n. |
| – | Metacoxa, pronotum, most of mesopleuron, metapleuron and at least anterior 0.5 of propodeum yellow-orange, same color than most of mesosoma (Figs 19, 22, 23, 59, 61–63, 70, 72, 75); T1 5.0–6.0 × as long as width at posterior margin | 13 |
| 13(12) | Humeral complex, part of axillar complex, and scutoscutellar sulcus dark brown to black; notauli very deeply impressed and with darker coloration than rest of anteromesoscutum (Fig. 73); hind wing subbasal cell mostly without setae (Figs 71, 75) [Brazil] | Prasmodon silvatlanticus Fernández-Triana & Whitfield, sp. n. |
| – | Humeral complex, axillar complex, and scutoscutellar sulcus orange-yellow, same colour than rest of mesosoma (Figs 22, 62); notauli not as strongly impressed as above, and same coloration than rest of anteromesoscutum; hind wing subbasal cell usually mostly setose (as in Fig. 63) [Costa Rica, ACG] | 14 |
| 14(13) | Propodeum with median longitudinal and transverse carinae clearly marked, without additional carination pattern (Fig. 23); T3–T7 mostly brown (Figs 19, 23) [Host: Crambidae, Palpita jairusalis DHJ01 feeding on Apocynaceae] | Prasmodon bobrobbinsi Fernández-Triana & Whitfield, sp. n. |
| – | Propodeum with irregular carinae in addition to median longitudinal and transverse carinae (Fig. 63); T3 yellow-orange; T4–T7 mostly orange with brown coloration restricted to central part of tergites (Fig. 63) [Host: unknown, Malaise-trapped] | Prasmodon paulgoldsteini Fernández-Triana & Whitfield, sp. n. |
| 15(10) | Mesosoma with extensive brown coloration: anteromesoscutum mid-frontal and lateral areas, lateral face of scutellum, axillar complex, and metascutellum (Figs 76, 77, 80, 81); T1 with posterior 0.2–0.3 brown (Figs 77, 81); metacoxa with small brown spot on posterior 0.1 (Fig. 76); metatrochanter and metatrochantellus brown | Prasmodon subfuscus Fernández-Triana & Whitfield, sp. n. |
| – | Mesosoma at most with anteromesoscutum mid-frontal and lateral areas brown (Figs 46, 50, 82, 86, 94); T1, metacoxa, metatrochanter and metatrochantellus fully yellow-white or yellow-orange (Figs 51, 82, 87, 94) | 16 |
| 16(15) | Metatibia and metatarsus entirely yellow (Figs 82, 85); wings hyaline, at most with very faint infumation on anterior 0.5 (Fig. 83); anteromesoscutum with only lateral areas brown, rest orange-yellow (Fig. 86) [Brazil] | Prasmodon tijucaensis Fernández-Triana & Whitfield, sp. n. |
| – | Posterior 0.1–0.2 of metatibia and most of metatarsus dark brown to black (Figs 46, 49, 94); wings infumate (Fig. 47); anteromesoscutum usually with mid-frontal and lateral areas brown, rest orange-yellow (Fig. 50) [Costa Rica, ACG] | 17 |
| 17(16) | Metasoma with only thin brown areas centrally on T4–T8 (Fig. 94); hind wing subbasal cell mostly without setae (Fig. 94) [Hosts: Crambidae, Ategumia, Desmia, Diacme, Herpetogramma, Phostria feeding on five plant families] | Prasmodon zlotnicki Valerio & Rodriguez, 2005 |
| – | Metasoma with most of T3–T8 dark brown (Figs 46, 51); hind wing subbasal cell mostly setose (Figs 48, 50) [Hosts: Elachistidae feeding on three plant families] | Prasmodon mikepoguei Fernández-Triana & Whitfield, sp. n. |
http://zoobank.org/3C6F2705-DC55-4192-89E1-66FF9BC2F570
http://species-id.net/wiki/Prasmodon_almasolisae
Figs 1–6, 101COSTA RICA, ACG, Alajuela Province, Sector Rincon Rain Forest, Sendero Anonas, 405m, 10.90528, -85.27882.
♀ in CNC. Specimen labels: 1. DHJPAR0038908. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 10-SRNP-40859.
2 ♀, 2 ♂ (CNC, NMNH). COSTA RICA, ACG database codes: DHJPAR0038170, DHJPAR0038913, DHJPAR0040001, DHJPAR0040518.
Female. Body length 4.9–5.0 mm, rarely 4.5–4.6 mm. Fore wing length 4.9–5.0 mm, 5.1–5.2 mm, rarely 5.3–5.4 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 5, 6). Scape color: partially dark brown to black (Fig. 5). Flagellomeres color: most flagellomeres brown to black, except for small apical area (F15–16, and occasionally apical half of F14) which is yellow-brown (Fig. 3). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partialy pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 4). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere) (Fig. 1). Fore wing color pattern: hyaline. Fore wing veins color: all veins dark brown (Fig. 2). Pterostigma color: entirely dark brown. Hypostomal carina: not raised. Scutoscutellar sulcus: with 6 impressions (Fig. 5). Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly without setae (Fig. 5). Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 6). Mediotergite 1 length÷width at posterior margin 3.6–4.0 ×. Mediotergite 2 width at posterior margin÷length: 3.1–3.5 ×. Ovipositor sheaths length: 0.6 × as long as metatibia, rarely 0.5 × as long as metatibia.
Male. As female but with flagellomeres fully dark brown to black.
Prasmodon almasolisae 1 Habitus 2 Fore wing 3 Antenna (partially) 4 Metatibia 5 Head and mesosoma, dorsal view 6 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 11, barcode compliant sequences: 11 (Fig. 105).
Hosts: Elachistidae, Antaeotricha radicalis, Antaeotricha ribbei, Antaeotricha Janzen07, Antaeotricha thapsinopa, elachJanzen01 Janzen131, Gonionota Janzen116, Stenoma Janzen129.
Costa Rica, ACG rain forest.
This species is named in honour of Alma Solis of the SEL/USDA laboratory in the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of her decades of taxonomic knowledge and support contributing to understanding the species-level and higher taxonomy of the Crambidae, Pyralidae and Thyrididae of ACG, INBio and Costa Rica, as well as other parts of the world.
http://zoobank.org/306A4FAB-DCD8-42C0-981F-C6B8D97EDC64
http://species-id.net/wiki/Prasmodon_aureus
Figs 7–12ECUADOR, Napo, Limoncocha, 250m.
♀ in CNC. Specimen labels: 1. ECUADOR, Napo, Limoncocha, 250m, 15–28.VI.1976, S. & J. Peck. 2. DNA Voucher, CNCHYM 01955.
Female. Body length 5.5–5.6 mm. Fore wing length 5.9–6.0 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 7, 11, 12). Scape color: entirely yellow (Fig. 11). Flagellomeres color: all flagellomeres brown to black (Figs 11, 12). Tegula and humeral complex color: both pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black. Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: most veins golden yellow (Figs 8, 10). Pterostigma color: mostly yellow, with central part brown (Fig. 10). Hypostomal carina: highly raised (Fig. 9). Scutoscutellar sulcus: with 4 impressions (Fig. 11). Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly with setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width. Mediotergite 1 length÷width at posterior margin 6.1–6.5 ×. Mediotergite 2 width at posterior margin÷length: 3.1–3.5 ×. Ovipositor sheaths length: 0.5 × as long as metatibia.
Male. Unknown.
Prasmodon aureus. 7 Habitus 8 Fore wing 9 Head and mesosoma (partially), lateral view 10 Detail of fore wing 11 Head and mesosoma, dorsal view 12 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 0.
Unknown.
Ecuador, Napo.
From Latin “aureus”= golden, referring to the yellow coloration of the fore wing veins, a unique feature among all known species of Prasmodon.
http://zoobank.org/00CA5006-4B13-4D9C-A7A7-58A6841F7192
http://species-id.net/wiki/Prasmodon_bobpoolei
Figs 13–18, 99COSTA RICA, ACG, Alajuela, Province, Sector Rincon Rain Forest, Montanya Figueres, 460m, 10.88367, -85.29081.
♀ in CNC. Specimen labels: 1. DHJPAR0045302. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 11-SRNP-42719.
4 ♀, 2 ♂ (CNC, NMNH). COSTA RICA, ACG database codes: DHJPAR0038211, DHJPAR0038246, DHJPAR0039997, DHJPAR0040003, DHJPAR0040005, DHJPAR0045165.
Female. Body length 4.1–4.2 mm, 4.3–4.4 mm, rarely 4.5–4.6 mm. Fore wing length 4.5–4.6 mm, rarely 4.7–4.8 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex). Scape color: partially dark brown to black (Figs 17, 18). Flagellomeres color: most flagellomeres brown to black, except for small apical area (F15–16, and occasionally apical half of F14) which is yellow-brown (Fig. 13). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partialy pale. Mesotibia color: posterior 0.1–0.2 dark brown to black. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 15). Metatibia spurs color: yellow-orange (Fig. 15). Metatarsus color: entirely yellow-white. Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 14). Pterostigma color: entirely dark brown. Hypostomal carina: not raised (Fig. 16). Scutoscutellar sulcus: with 5 impressions (Fig. 17). Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 18). Mediotergite 1 length÷width at posterior margin 5.6–6.0 ×. Mediotergite 2 width at posterior margin÷length: 2.6–3.0 ×. Ovipositor sheaths length: 0.6 × as long as metatibia, rarely 0.5 × as long as metatibia.
Male. As female but with flagellomeres fully dark brown to black.
Sequences in BOLD: 15, barcode compliant sequences: 15 (Fig. 105).
Prasmodon bobpoolei. 13 Habitus 14 Fore wing 15 Metatibia 16 Head and mesosoma (partially), lateral view 17 Head and mesosoma, dorsal view 18 Propodeum and metasoma, dorsal view.
Host: Elachistidae, Antaeotricha BioLep38, Antaeotricha Janzen110, Antaeotricha Janzen126, Gonioterma Janzen212, Stenoma Janzen129.
Costa Rica, ACG rain forest.
This species is named in honour of Bob Poole of the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of his decades of taxonomic knowledge and support contributing to understanding the species-level and higher taxonomy of the Noctuoidea and Geometroidea of ACG, INBio and Costa Rica, as well as other parts of the world.
http://zoobank.org/4C754CF8-825E-42AE-99C0-D59F17DC178A
http://species-id.net/wiki/Prasmodon_bobrobbinsi
Figs 19–23, 100COSTA RICA, ACG, Alajuela Province, Sector Rincon Rain Forest, San Lucas, 320m, 10.91847, -85.30338.
♀ in CNC. Specimen labels: 1. DHJPAR0038915. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 10-SRNP-40484.
1 ♀, 1♂ (CNC). COSTA RICA, ACG database codes: DHJPAR0040012, 09-SRNP-3496.
Female. Body length 3.7–3.8 mm or 3.9–4.0 mm. Fore wing length 3.9–4.0 mm or 4.1–4.2 mm. Body color: mesosoma entirely yellow-orange (with the exception of a very small black spot on axillar complex), metasoma with brown to black areas covering most of mediotergites 4–7 (and also part of mediotergites 3 and 8) (Figs 19, 22, 23). Scape color: partially dark brown to black. Flagellomeres color: all flagellomeres brown to black (Figs 19, 21). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partialy pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black. Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 21). Pterostigma color: entirely dark brown. Hypostomal carina: highly raised (Fig. 20). Scutoscutellar sulcus: with 6 impressions (Fig. 22). Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly with setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 23). Mediotergite 1 length÷width at posterior margin 4.1–4.5 × or 4.6–5.0 ×. Mediotergite 2 width at posterior margin÷length: 2.6–3.0 ×. Ovipositor sheaths length: 0.5 × as long as metatibia.
Male. Morphologically similar to females.
Prasmodon bobrobbinsi. 19 Habitus 20 Head and mesosoma (partially), lateral view 21 Fore wing and antenna (partialy) 22 Head and mesosoma, dorsal view 23 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 2, barcode compliant sequences: 2 (Fig. 105).
Host: Crambidae, Palpita jairusalis DHJ01.
Costa Rica, ACG rain forest.
This species is named in honour of Bob Robbins of the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of his decades of taxonomic knowledge and support, contributing to understanding the species-level and higher taxonomy of the Lycaenidae of ACG, INBio and Costa Rica, as well as other parts of the world.
http://zoobank.org/31AF25BD-0151-46EF-A980-1B9BF3BB339E
http://species-id.net/wiki/Prasmodon_dondavisi
Figs 24–28COSTA RICA, ACG, Alajuela Province, Sector Rincón Rain Forest, Jacobo, 461m, 10.94076, -85.3177.
♀ in CNC. Specimen labels: 1. DHJPAR0040002. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 09-SRNP-69586.
1 ♀, 9 ♂ (CNC, INHS, NMNH, ). COSTA RICA, ACG database code: DHJPAR003990, DHJPAR0039992, DHJPAR0039994, DHJPAR0039995, DHJPAR0039998, DHJPAR004000, DHJPAR0040004, DHJPAR0040006, DHJPAR0040007, DHJPAR0040010.
Female. Body length 5.3–5.4 mm or 5.5–5.6 mm. Fore wing length 5.5–5.6 mm or 5.7–5.8 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 24, 27, 28). Scape color: partially dark brown to black. Flagellomeres color: all flagellomeres brown to black (Fig. 24). Tegula and humeral complex color: both pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 26). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown. Pterostigma color: entirely dark brown. Hypostomal carina: not raised (Fig. 25). Scutoscutellar sulcus: with 4 impressions (Fig. 27). Areolet height÷vein r length (fore wing): 0.3 ×. Hind wing subbasal cell: mostly with setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing at around 0.5 its length, with both anterior and posterior margins clearly wider than its median width (Fig. 28). Mediotergite 1 length÷width at posterior margin 3.1–3.5 × or 3.6–4.0 ×. Mediotergite 2 width at posterior margin÷length: 3.1–3.5 ×. Ovipositor sheaths length: 0.5 × as long as metatibia.
Male. Morphologically similar to females.
Prasmodon dondavisi. 24 Habitus 25 Head and mesosoma (partially), lateral view 26 Metatibia 27 Head and mesosoma, dorsal view 28 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 17, barcode compliant sequences: 15 (Fig. 105).
Hosts: Crambidae feeding on Apocynaceae: Prenesta Janzen196, Prenesta Janzen195, Prenesta scyllalis DHJ03.
Costa Rica, ACG rain forest.
This species is named in honour of Don Davis of the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of his decades of taxonomic knowledge and support, contributing to understanding the species-level and higher taxonomy of the Tineoidea and other small impossible moths of ACG, INBio and Costa Rica, as well as other parts of the world.
http://species-id.net/wiki/Prasmodon_eminens
Figs 95, 97PERU: Chanchamayo.
♂, NHM (not examined).
8 ♀, 18 ♂ (BMNH, CNC, INBio, INHS, NMNH), Costa Rica, ACG; 2 ♂ (CNC), Ecuador, Pichincha, Rio Palenque, 160–200m.
Female. Body length 4.9–5.0 mm, 5.1–5.2 mm, rarely 5.3–5.4 mm. Fore wing length 5.1–5.2 mm, 5.3–5.4 mm, rarely 5.5–5.6 mm. Body color: mesosoma entirely yellow-orange (with the exception of a very small black spot on axillar complex), metasoma with only small brown areas centrally on mediotergites 4–7 (usually only on mediotergires 5–6) (Fig. 95). Scape color: partially dark brown to black. Flagellomeres color: most flagellomeres brown to black, except for small apical area (F15–16, and occasionally apical half of F14) which is yellow-brown. Tegula and humeral complex color: both pale. Mesotibia color: posterior 0.1–0.2 dark brown to black. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 95). Metatibia spurs color: dark brown to black. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere) (Fig. 95). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 95). Pterostigma color: entirely dark brown. Hypostomal carina: not raised. Scutoscutellar sulcus: with 5 impressions or with 6 impressions. Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 95). Mediotergite 1 length÷width at posterior margin 4.1–4.5 ×. Mediotergite 2 width at posterior margin÷length: 3.1–3.5 x, rarely 2.6–3.0 ×. Ovipositor sheaths length: 0.5 × as long as metatibia or 0.6 × as long as metatibia.
Male. As female but with flagellomeres fully dark brown to black.
Sequences in BOLD: 67, barcode compliant sequences: 61 (Fig. 105).
Hosts: Crambidae, Anarmodia nebulosalis, Asturodes fimbriauralis DHJ02, Bocchoris marucalis, Ceratocilia sixolalis, Desmia Janzen07, Desmia Solis19, Eulepte concordalis, Eulepte Janzen12, Eulepte Solis15, Mimophobetron pyropsalis, Omiodes fulvicauda, Omiodes humeralis, Pantographa suffusalis, Parastenia retractalis, Phostria mapetalis, Piletosoma thialis, Portentomorpha xanthialis, and Syllepte amando DHJ02.
Widely distributed in rain forest in northern South America and Central America, including Brazil (
This species was described by
http://zoobank.org/D163BA78-BDE4-4F8E-B549-909751C9785A
http://species-id.net/wiki/Prasmodon_erenadupontae
Figs 29–34FRENCH GUIANA, Saul.
♀ in MNHN. Specimen labels: 1. Guyane Francaise, Saul, 29.IX.2010, SEAG rec 2010.
2 ♀, 3 ♂ (CNC, IRSNB). French Guiana, same locality than holotype, 17–26.ix.2010 (1 ♀, 1 ♂); Montagne de Kaw, Relais Patawa, ix.1999 (1 ♀); Nouragues Reserve, Inselberg, 16.x.2010 (1 ♂); Brazil, Mato Grosso, Sinop, xi.1975 (1 ♂).
Female. Body length 5.3–5.4 mm, 5.5–5.6 mm or 5.7–5.8 mm. Fore wing length 5.9–6.0 mm or 6.1–6.2 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 29, 32, 34). Scape color: partially dark brown to black. Flagellomeres color: with relatively extense yellow area (at least including F13–15, and usually apical half of F11 and basal half of F16) (Fig. 30). Tegula and humeral complex color: both pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black. Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere) (Fig. 30). Fore wing color pattern: mostly infumate, but with pale area centrally which gives the wing a banded appearance (Figs 30, 33). Fore wing veins color: anterior 0.4–0.5 of vein 3M (and usually part or all of veins 2M and r-m) yellow-orange, contrasting with all other veins which are dark brown (Fig. 33). Pterostigma color: entirely dark brown (Fig. 30). Hypostomal carina: highly raised (Fig. 31). Scutoscutellar sulcus: with 4 impressions (Fig. 32). Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing at around 0.5 its length, with both anterior and posterior margins clearly wider than its median width (Fig. 34). Mediotergite 1 length÷width at posterior margin 2.9–3.0 ×. Mediotergite 2 width at posterior margin÷length: 3.6–4.0 ×. Ovipositor sheaths length: 0.5 × as long as metatibia.
Male. Morphologically similar to females, but with antenna only yellow on flagellomeres 15–16 (females with flagellomeres 12–16 yellow).
Prasmodon erenadupontae. 29 Habitus 30 Body, dorso-lateral view 31 Head and mesosoma (partially), lateral view 32 Head and mesosoma, dorsal view 33 Fore wing 34 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 1.
Malaise trapped.
Brazil, French Guiana.
Named after Erena Dupont (the partner of YB) who shares his passion in entomology and for her strong support.
The only specimen with associated molecular data is the male paratype from Brazil.
http://zoobank.org/F131E6A1-F5FF-414D-803F-7719C0E2A533
http://species-id.net/wiki/Prasmodon_johnbrowni
Figs 35–39, 103COSTA RICA, ACG, Alajuela Province, Sector Rincon Rain Forest, Estación Llanura, 135m, 10.93332, -85.25331.
♀ in CNC. Specimen labels: 1. DHJPAR0035300. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 09-SRNP-44366.
5 ♀, 2 ♂ (CNC, NMNH). COSTA RICA, ACG database code: DHJPAR0038172, DHJPAR0038174, DHJPAR0038175, DHJPAR0038177, DHJPAR0038922, DHJPAR0039993.
Female. Body length 4.5–4.6 mm, 4.7–4.8 mm, 4.9–5.0 mm, rarely 5.1–5.2 mm. Fore wing length 4.9–5.0 mm, 5.1–5.2 mm, rarely 5.3–5.4 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 35, 38, 39). Scape color: partially dark brown to black. Flagellomeres color: all flagellomeres brown to black (Fig. 35). Tegula and humeral complex color: tegula pale, humeral complex dark. Mesotibia color: posterior 0.1–0.2 dark brown to black. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 37). Metatibia spurs color: dark brown to black (Figs 35, 37). Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 36). Pterostigma color: entirely dark brown (Fig. 38). Hypostomal carina: not raised. Scutoscutellar sulcus: with 6 impressions, rarely with 5 impressions. Areolet height÷vein r length (fore wing): 0.3 ×. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 39). Mediotergite 1 length÷width at posterior margin 6.1–6.5 ×. Mediotergite 2 width at posterior margin÷length: 2.6–3.0 ×. Ovipositor sheaths length: 0.5 × as long as metatibia or 0.6 × as long as metatibia.
Male. Morphologically similar to females.
Prasmodon johnbrowni. 35 Habitus 36 Fore wing 37 Metatibia 38 Head and mesosoma, dorsal view 39 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 12, barcode compliant sequences: 12 (Fig. 105).
Hosts: Crambidae, Asturodes fimbriauralis DHJ02, Asturodes fimbriauralis, Eulepte alialis, Piletosoma thialis, Phostria Janzen05.
Costa Rica, ACG rain forest.
This species is named in honour of John Brown of the SEL/USDA laboratory in the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of his decades of taxonomic knowledge and support contributing to understanding the species-level and higher taxonomy of the Tortricoidea of ACG, INBio and Costa Rica, as well as other parts of the world.
http://zoobank.org/39CD8542-C231-4CDB-A67B-57782406696B
http://species-id.net/wiki/Prasmodon_masoni
Figs 40–45BRAZIL, Mato Grosso, Sinop.
♀ in CNC. Specimen labels: 1. BRAZIL, Mato Grosso, Sinop, X.1974., M. Alvarenga, Mal. Trap. 2. DNA Voucher, CNCHYM 01962.
1 ♀, 6 # M (CNC), Brazil, same locality than holtype, collecting dates: x.1974, x.1975, ii.1976.
1 ♀ (CNC) from Brazil, Amazonas, Estirar de Ecuador, Rio Javari, ix.1979.
Female. Body length 3.5–3.6 mm, rarely 3.9–4.0 mm. Fore wing length 3.7–3.8 mm, rarely 4.1–4.2 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 40, 44, 45). Scape color: partially dark brown to black (Fig. 44). Flagellomeres color: all flagellomeres brown to black (Figs 40–42). Tegula and humeral complex color: both pale. Mesotibia color: entirely yellow. Metatibia color: entirely yellow (Fig. 43). Metatibia spurs color: yellow-orange. Metatarsus color: entirely yellow-white (Fig. 43). Fore wing color pattern: hyaline. Fore wing veins color: all veins dark brown. Pterostigma color: entirely dark brown. Hypostomal carina: not raised (Fig. 42). Scutoscutellar sulcus: with 4 impressions (Fig. 44). Areolet height÷vein r length (fore wing): 0.2 x, rarely 0.3 ×. Hind wing subbasal cell: mostly with setae (Figs 44, 45). Hind tarsal claws: with pectination (teeth) very irregular in spacing and length. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 45). Mediotergite 1 length÷width at posterior margin 4.1–4.5 × or 4.6–5.0 ×. Mediotergite 2 width at posterior margin÷length: 2.6–3.0 ×. Ovipositor sheath length: 0.3 × as long as metatibia.
Male. Morphologically similar to female.
Prasmodon masoni. 40 Habitus 41 Fore wing 42 Head and mesosoma (partially), lateral view 43 Hypopygium, ovipositor, ovipositor sheaths, and hind leg (partially), lateral view 44 Head and mesosoma, dorsal view 45 Propodeum and metasoma, dorsal view.
No DNA barcodes were obtained for this species.
Malaise-trapped.
Brazil, Mato Grosso.
Named after William R. M. Mason, the greatest Canadian expert on Microgastrinae. The holotype and paratype specimen had a label written by Mason, who considered them to represent a new species (Mason, 1981), which remained undescribed until now.
We have included here one female from a locality (Rio Javari, Brazil) far from the type locality (Sinop, where all other specimens of the species were collected). The female from Rio Javari resembles the female holotype and another female paratype, but was not included as part of the paratype series in case it is eventually found to represent a different species. We also studied another 15 CNC male specimens of Prasmodon that were collected in Mato Grosso. They seem to represent 2–3 additional species. However, no molecular or host information is available for those specimens, and the morphological data are inadequate to properly describe them at present.
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http://species-id.net/wiki/Prasmodon_mikepoguei
Figs 46–51, 96COSTA RICA, ACG, Guanacaste Province, Sector Pitilla, Leonel, 510m, 10.99637, -85.40195.
♂ in CNC. Specimen labels: 1. DHJPAR0038222. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 09-SRNP-72161.
Male. Body length 3.7–3.8 mm. Fore wing length 4.1–4.2 mm. Body color: meso- and metasoma with brown to black areas, metasoma with most of tergites 2–8 dark brown (Figs 46, 50, 51). Scape color: partially dark brown to black. Flagellomeres color: all flagellomeres brown to black (Fig. 47). Tegula and humeral complex color: tegula pale, humeral complex dark. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 49). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown. Pterostigma color: entirely dark brown. Hypostomal carina: not raised. Scutoscutellar sulcus: with 4 impressions (Fig. 50). Hind wing subbasal cell: mostly with setae (Figs 48, 50, 51). Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 51).
Female. Unknown.
Prasmodon mikepoguei. 46 Habitus 47 Fore wing 48 Detail of the hind wing 49 Metatibia 50 Head and mesosoma, dorsal view 51 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 4, barcode compliant sequences: 4 (Fig. 105).
Hosts: Elachistidae, Chlamydastis tryphon, Antaeotricha BioLep42, Antaeotricha Janzen07.
Costa Rica, ACG rain forest.
This species is named in honour of Mike Pogue of the SEL/USDA laboratory in the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of his decades of taxonomic knowledge and support contributing to understanding the species-level and higher taxonomy of the Notodontidae and Noctuidae of ACG, INBio and Costa Rica, as well as other parts of the world.
http://zoobank.org/989FD927-8A4A-4582-A69D-DE20AFA9841D
http://species-id.net/wiki/Prasmodon_nixoni
Figs 52–58PERU, Loreto, Iquitos, Barillal.
♀ in CNC. Specimen labels: 1. PERU: Loreto, Iquitos, Barillal, 10.ii.1984, L. Huggert. 2. DNA Voucher, CNCHYM 01957.
1 ♀ (IRSNB). French Guiana, Montagne de Kaw, Relais Patawa, v.2001.
Female. Body length 4.3–4.4 mm or 4.5–4.6 mm. Fore wing length 4.5–4.6 mm or 4.7–4.8 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 52, 56, 58). Scape color: partially dark brown to black. Flagellomeres color: with relatively extense yellow area (at least including F13–15, and usually apical half of F11 and basal half of F16) (Figs 52–54). Tegula and humeral complex color: both pale. Mesotibia color: entirely yellow. Metatibia color: entirely yellow (Fig. 55). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: hyaline. Fore wing veins color: all veins dark brown (Fig. 53). Pterostigma color: entirely dark brown. Hypostomal carina: not raised (Fig. 57). Scutoscutellar sulcus: with 6 impressions (Fig. 56). Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly with setae. Hind tarsal claws: with 1 or 2 teeth on anterior 0.3 of claw. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 58). Mediotergite 1 length÷width at posterior margin 4.6–5.0 ×. Mediotergite 2 width at posterior margin÷length: 3.6–4.0 ×. Ovipositor sheaths length: 0.5 × as long as metatibia.
Male. Unknown.
Prasmodon nixoni. 52 Habitus 53 Fore wing 54 Antenna (partially) 55 Metatibia 56 Head and mesosoma, dorsal view 57 Head and mesosoma (partially), lateral view 58 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 0.
Malaise-trapped.
French Guiana, Peru.
Named to honour Gilbert E. J. Nixon, one of the great British experts on Microgastrinae and the first person to describe a species of Prasmodon.
http://zoobank.org/3AD73865-F1F1-415F-9DDB-41BFAA5D0E1A
http://species-id.net/wiki/Prasmodon_paulgoldsteini
Figs 59–63COSTA RICA, Guanacaste, ACG, Sector Santa Rosa, Bosque San Emilio, 300m, 10.84389, -85.61384.
♂ in CNC. Specimen labels: 1. DHJPAR0013126.
4 ♂ (CNC, NMNH). COSTA RICA, ACG database codes: DHJPAR0013131, DHJPAR0013124, DHJPAR0013125, DHJPAR0013132.
Male. Body length 4.1–4.2 mm. Fore wing length 4.3–4.4 mm. Body color: mesosoma entirely yellow-orange (with the exception of a very small black spot on axillar complex), metasoma with only small brown areas centrally on mediotergites 4–7 (usually only on mediotergires 5–6) (Figs 62, 63). Scape color: partially dark brown to black. Flagellomeres color: all flagellomeres brown to black (Fig. 59). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partially pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 59). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 60). Pterostigma color: entirely dark brown. Hypostomal carina: highly raised. Scutoscutellar sulcus: with 5 impressions or with 6 impressions. Hind wing subbasal cell: mostly with setae. Hind tarsal claws: with pectination (teeth) very irregular in spacing and length. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 63).
Female. Unknown.
Prasmodon paulgoldsteini. 59 Habitus 60 Fore wing 61 Head and mesosoma (partially), lateral view 62 Head and mesosoma, dorsal view 63 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 6, barcode compliant sequences: 6 (Fig. 105).
Malaise trapped.
Costa Rica, ACG.
At present this species is only known from males, but it was described because of clear morphological and molecular differences with the other species within the genus.
This species is named in honour of Paul Goldstein working in the SEL/USDA laboratory in the National Museum of Natural History, Smithsonian Institution, Washington, D.C, and with the Department of Entomology of the University of Maryland, in recognition of his intense effort to solve the Desmia (Crambidae, Spilomelinae) many-species taxonomic puzzle for ACG.
http://zoobank.org/8BCFA3EE-53D0-4582-AE6B-A31D544DE9B1
http://species-id.net/wiki/Prasmodon_scottmilleri
Figs 64–69, 98COSTA RICA, ACG, Alajuela Province, Sector Rincon Rain Forest, Camino Albergue Oscar, 560m, 10.87741, -85.32363.
♀ in CNC. Specimen labels: 1. DHJPAR0043031. 2. Voucher: D.H.Janzen & W.Hallwachs, DB: http://janzen.sas.upenn.edu, Area de Conservación Guanacaste, COSTA RICA, 11-SRNP-610.
1 ♂ (CNC). COSTA RICA, ACG database code: DHJPAR0038914.
Female. Body length 3.9–4.0 mm. Fore wing length 4.5–4.6 mm. Body color: mesosoma entirely yellow-orange (with the exception of a very small black spot on axillar complex), metasoma with brown to black areas covering most of mediotergites 4–7 (and also part of mediotergites 3 and 8). Scape color: partially dark brown to black. Flagellomeres color: most flagellomeres brown to black, except for small subapical area (F14–15) which is yellow-brown (Figs 64, 67). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partialy pale. Mesotibia color: posterior 0.1–0.2 dark brown to black. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 64). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere) (Fig. 64). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 65). Pterostigma color: entirely dark brown. Hypostomal carina: highly raised. Scutoscutellar sulcus: with 4 impressions. Areolet height÷vein r length (fore wing): 0.15 × or less. Hind wing subbasal cell: mostly without setae (Figs 65, 69). Hind tarsal claws: with pectination (teeth) very irregular in spacing and length. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 69). Mediotergite 1 length÷width at posterior margin 3.6–4.0 ×. Mediotergite 2 width at posterior margin÷length: 3.6–4.0 ×. Ovipositor sheaths length: 0.6 × as long as metatibia.
Male. As female but with flagellomeres fully dark brown to black.
Prasmodon scottmilleri. 64 Habitus 65 Fore wing 66 Detail of the hind wing 67 Antenna (partially) 68 Head and mesosoma, dorsal view 69 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 6, barcode compliant sequences: 5 (Fig. 105).
Hosts: Crambidae, four species of Omiodes and Triuncidia eupalusalis DHJ02.
Costa Rica, ACG rain forest.
This species is named in honour of Scott Miller of the National Museum of Natural History, Smithsonian Institution, Washington, D.C, in recognition of his decades of taxonomic knowledge and support contributing to understanding the species-level and higher taxonomy of the Dalceridae, Lasiocampidae, and other fat fuzzy moths of ACG, INBio and Costa Rica, as well as other parts of the world, and administrating Lepidoptera biodiversity development at the Smithsonian Institution.
http://zoobank.org/A9F2150A-1D0C-402E-9945-FC21A1663857
http://species-id.net/wiki/Prasmodon_silvatlanticus
Figs 70–75BRAZIL, Est. Rio de Janeiro, Silva Jardim.
♀ in CNC. Specimen labels: 1. BRAZIL, Est. Rio de Janeiro, Silva Jardim, viii.1974, F. M. Oliveira. 2. DNA Voucher CNCHYM 01960.
Female. Body length 4.1–4.2 mm. Fore wing length 4.5–4.6 mm. Body color: mesosoma entirely yellow-orange (with the exception of a very small black spot on axillar complex), metasoma with brown to black areas covering most of mediotergites 4–7 (and also part of mediotergites 3 and 8). Scape color: partially dark brown to black. Flagellomeres color: most flagellomeres brown to black, except for small subapical area (F14 and apical half of F13) which is yellow-brown (Figs 70, 74). Tegula and humeral complex color: tegula pale, humeral complex dark. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black (Figs 70, 72). Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere) (Fig. 70). Fore wing color pattern: hyaline. Fore wing veins color: all veins dark brown (Fig. 71). Pterostigma color: entirely dark brown. Hypostomal carina: highly raised (Fig. 72). Scutoscutellar sulcus: with 6 impressions. Areolet height÷vein r length (fore wing): 0.15 × or less. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination uniform, teeth thick and relatively evenly spaced. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 75). Mediotergite 1 length÷width at posterior margin 5.6–6.0 ×. Mediotergite 2 width at posterior margin÷length: 2.6–3.0 ×. Ovipositor sheaths length: 0.6 × as long as metatibia.
Male. Unknown.
Prasmodon silvatlanticus. 70 Habitus 71 Fore wing 72 Head and mesosoma (partially), lateral view 73 Head and mesosoma, dorsal view 74 Antenna 75 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 0.
Unknown.
Brazil, Rio de Janeiro.
From Latin “silva”=”forest” and “atlanticus”=Atlantic, referring to the Atlantic Forest biome, where this and other species of Prasmodon have been found. The first part of the name also refers to the type locality (Silva Jardim).
The only known specimen was considered by
http://zoobank.org/8BD6E416-FF8F-4FDC-8581-9C97B061BA2A
http://species-id.net/wiki/Prasmodon_subfuscus
Figs 76–81BRAZIL, Rio de Janeiro, Silva Jardim.
#1 in CNC. Specimen labels: 1. BRAZIL, Est. Rio de Janeiro, Silva Jardim, viii.1974, F. M. Oliveira. 2. DNA Voucher CNCHYM 01961.
Male. Body length 4.5–4.6 mm. Fore wing length 5.1–5.2 mm. Body color: meso- and metasoma with brown to black areas, metasoma with most of tergites 2–8 dark brown (Figs 77, 80, 81). Scape color: partially dark brown to black (Fig. 79). Flagellomeres color: all flagellomeres brown to black (Fig. 76). Tegula and humeral complex color: tegula pale, humeral complex dark. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 78). Metatibia spurs color: yellow-orange (Fig. 78). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown (Fig. 77). Pterostigma color: entirely dark brown. Hypostomal carina: highly raised (Fig. 79). Scutoscutellar sulcus: with 5 impressions. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination (teeth) very irregular in spacing and length. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 81).
Female. Unknown.
Prasmodon subfuscus. 76 Habitus 77 Fore wing 78 Metatibia 79 Head and mesosoma (partially), lateral view 80 Head and mesosoma, dorsal view 81 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 0.
Unknown.
Brazil, Rio de Janeiro.
From Latin “subfuscus”=brownish, referring to the extensive brown coloration of the species, by far the least yellow of all known species of Prasmodon.
This species, only known from a single male, was collected in the same locality and on the same date as Prasmodon silvatlanticus, which is known only from a single female. Both specimens are morphologically distinct, with different coloration of mesosoma, mediotergite 1, metacoxa, and trochanter (Figs 70, 73, 75 vs Figs 76, 80, 81), as well as shape of mediotergite 2. We have not found sexual dimorphism among any of the known species of Prasmodon (other than slight variations in the color of apical flagellomeres), and thus we do not think that the differences observed represent sexual dimorphism within a single species. Further evidence to consider those two specimens as separate species is provided by DNA barcoding; even though only short sequences (164 bp) could be recovered from both of them, they differed by 15 bp (9.5%).
The exact collection locality of Prasmodon subfuscus is not clear from the original labels. “Silva Jardim” is a municipality of 938 km2 in the Brazilian state of Rio de Janeiro. Within the municipality is located the “Reserva Biologica de Poço das Antas”, the largest lowland Atlantic Forest reserve in the state of Rio de Janeiro, containing 50 km2 of mostly secondary, evergreen forest. The area is very diverse (e.g., it contains at least 365 plant and 77 mammal species,
http://zoobank.org/0E2B527E-FEDB-478C-B739-08AE323E559D
http://species-id.net/wiki/Prasmodon_tijucaensis
Figs 82–87BRAZIL, Rio de Janeiro, Guanabara, Floresta da Tijuca (Tijuca Forest).
♀ in CNC. Specimen labels: 1. BRAZIL, Guanabara, Floresta da Tijuca, ii.1974, M. Alvarenga. 2. DNA Voucher CNCHYM 01959.
Female. Body length 3.3–3.4 mm. Fore wing length 3.7–3.8 mm. Body color: meso- and metasoma with brown to black areas, metasoma with most of tergites 2–8 dark brown (Fig 86, 87). Scape color: partially dark brown to black (Fig. 86). Flagellomeres color: all flagellomeres brown to black (Fig. 82). Tegula and humeral complex color: both dark. Mesotibia color: entirely yellow. Metatibia color: entirely yellow (Fig. 85). Metatibia spurs color: yellow-orange. Metatarsus color: entirely yellow-white. Fore wing color pattern: hyaline (Fig. 83). Fore wing veins color: all veins dark brown (Fig. 83). Pterostigma color: entirely dark brown. Hypostomal carina: not raised (Fig. 84). Scutoscutellar sulcus: with 7 impressions (Fig. 86). Areolet height÷vein r length (fore wing): 0.3 ×. Hind wing subbasal cell: mostly with setae. Hind tarsal claws: with 1 or 2 teeth on anterior 0.3 of claw. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 87). Mediotergite 1 length÷width at posterior margin 6.1–6.5 ×. Mediotergite 2 width at posterior margin÷length: 2.1–2.5 ×. Ovipositor sheaths length: 0.4 × as long as metatibia.
Male. Unknown.
Prasmodon tijucaensis. 82 Habitus 83 Fore wing 84 Head and mesosoma (partially), lateral view 85 Metatibia 86 Head and mesosoma, dorsal view 87 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 0.
Unknown.
Brazil, Rio de Janeiro.
The name honours the type locality, Floresta da Tijuca, which is among the world’s largest urban forests.
The only known specimen was considered by
http://zoobank.org/7456681F-AAEC-4B62-ABC3-B1C9EB3387F2
http://species-id.net/wiki/Prasmodon_verhoogdenokus
Figs 88–93BRAZIL, Mato Grosso, Vila Vera, 500m, -12.7667, -55.5.
♀ in CNC. Specimen labels: 1. BRAZIL, Vila Vera, M. Grosso, 500m, 12°46'S, 55°30'W, X.1973, M. Alvarenga. 2. DNA Voucher, CNCHYM 01971.
6 ♀ (CNC, RMNH, IRSNB, MNHN). Ecuador, Napo, Tena, 400m, 16–17.ii.1987 (1 ♀); French Guiana, Saul, collecting dates: 2.ix.2010, 17.ix.2010, 26.x.2010 (3 ♀), Saul, Crique popote, Mont Belvedère, xii.2000 (1 ♀), Kourou, Piste Soumourou, 13.vi-20.vii.2002 (1 ♀).
3 ♂ from Colombia, Amazonas, PNN Amacayacu, San Martin, 150m (INHS); 2 ♂ from Ecuador, Napo, P. Misahualli, 350m (CNC); 98 ♂ from French Guiana, Sinamary, Barrage de Petit-Saut; Inselberg Itoupé; RNR Trinité, Zone Aya; Nourages Reserve; Montagne de Kaw, Relais Patawa; and Saul (CNC and IRSNB); 2 ♂ from Peru, Madre de Dios, Rio Tampobata, Sachavacayoc center; and Loreto, Matse’s Reserved Zone, site Jenaro Herrera, Rio Ucayali (CNC); 1 ♂ from Suriname, Paramaribo Zoo (RMNH).
Female. Body length 4.1–4.2 mm, 4.3–4.4 mm, 4.5–4.6 mm, rarely 3.9–4.0 mm or 4.7–4.8 mm. Fore wing length 4.3–4.4 mm, 4.5–4.6 mm, rarely 4.9–5.0 mm. Body color: meso- and metasoma entirely yellow-orange (with the exception of a very small black spot on axillar complex) (Figs 92, 93). Scape color: partially dark brown to black (Fig. 92). Flagellomeres color: with relatively extense yellow area (at least including F13–15, and usually apical half of F11 and basal half of F16) (Fig. 88, 91). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partialy pale. Mesotibia color: entirely yellow. Metatibia color: posterior 0.1–0.3 dark brown to black. Metatibia spurs color: yellow-orange. Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: mostly infumate, but with pale area centrally which gives the wing a banded appearance. Fore wing veins color: all veins dark brown (Fig. 89). Pterostigma color: entirely dark brown. Hypostomal carina: highly raised. Scutoscutellar sulcus: with 6 impressions, rarely with 5 impressions. Areolet height÷vein r length (fore wing): 0.15 × or less. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination (teeth) very irregular in spacing and length. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 93). Mediotergite 1 length÷width at posterior margin 5.6–6.0 ×. Mediotergite 2 width at posterior margin÷length: 2.6–3.0 ×. Ovipositor sheaths length: 0.6 × as long as metatibia, rarely 0.5 × as long as metatibia.
Male. Morphologically similar to females, except for antenna.
Prasmodon verhoogdenokus. 88 Habitus 89 Fore wing 90 Antenna (partially) 91 Metatibia 92 Head and mesosoma, dorsal view 93 Propodeum and metasoma, dorsal view.
Sequences in BOLD: 1, barcode compliant sequences: 0. Two additional short sequences are available from two male specimens, but they were not included in the paratype series (see Comments below).
Light trapped and Malaise trapped.
Brazil, Colombia, Ecuador, French Guiana, Peru, Suriname.
From Dutch “verhoogde nok”= raised ridge, referring to the raised hypostomal carina.
The wide distribution of Prasmodon verhoogdenokus throughout South America, suggests it may represent a complex of morphologically cryptic species. At present there is no host known for the species, and the molecular data are meagre. Two male specimens from Ecuador that appear to be of this species rendered partial DNA sequences that differed by approximately 5% bp from the partial barcode of the female holotype (but the COI sequences for those three specimens only overlap for 240 bp). Although males included under this species look morphologically similar to the female holotype and the paratypes, they have been left out of the paratype series.
http://species-id.net/wiki/Prasmodon_zlotnicki
Figs 94, 102COSTA RICA: ACG, Alajuela Province, Sector San Cristobal, Sendero Corredor, 620 m, 10.87868, -85.38963. In the holotype description, the province was erroneously given as Guanacaste Province.
♀, INBio (examined).
8 ♀, 1 ♂ (CNC, INBio, NMNH).
Female. Body length 4.5–4.6 mm. Fore wing length 5.1–5.2 mm or 5.3–5.4 mm. Body color: meso- and metasoma with brown to black areas, those on metasoma only thin brown areas centrally on tergites 4–8 (Fig. 94). Scape color: partially dark brown to black. Flagellomeres color: most flagellomeres brown to black, except for small apical area (F15–16, and occasionally apical half of F14) which is yellow-brown (Fig. 94). Tegula and humeral complex color: tegula pale, humeral complex partially dark/partialy pale. Mesotibia color: posterior 0.1–0.2 dark brown to black. Metatibia color: posterior 0.1–0.3 dark brown to black (Fig. 94). Metatibia spurs color: dark brown to black (Fig. 94). Metatarsus color: dark brown to black (except for anterior 0.7 or less of first metatarsomere). Fore wing color pattern: uniformly and entirely infumate (except for small hyaline area near veins (RS+M)b and 2M). Fore wing veins color: all veins dark brown. Pterostigma color: entirely dark brown. Hypostomal carina: highly raised. Scutoscutellar sulcus: with 6 impressions. Areolet height÷vein r length (fore wing): 0.2 ×. Hind wing subbasal cell: mostly without setae. Hind tarsal claws: with pectination (teeth) very irregular in spacing and length. Shape of mediotergite 1: distinctly narrowing posteriorly, width at posterior margin clearly less than width at anterior margin and median width (Fig. 94). Mediotergite 1 length÷width at posterior margin 4.6–5.0 x, 5.1–5.5 x, rarely 3.6–4.0 ×. Mediotergite 2 width at posterior margin÷length: 3.1–3.5 × or 3.6–4.0 ×. Ovipositor sheaths length: 0.5 × as long as metatibia.
Male. One male, the first found for the species. It is morphologically similar to females.
94 Prasmodon zlotnicki. Body, dorsal view 95 Prasmodon eminens. Body, dorsal view.
96 Cocoon of Prasmodon mikepoguei 97 Cocoon of Prasmodon eminens.
98 Cocoon of Prasmodon scottmilleri 99 Cocoon of Prasmodon bobpoolei.
100 Cocoon of Prasmodon bobrobbinsi 101 Cocoon of Prasmodon almasolisae.
102 Cocoon of Prasmodon zlotnicki 103 Cocoon of Prasmodon johnbrowni.
Distribution map of Prasmodon species in Central and South America.
Neighbor-Joining tree (
Sequences in BOLD: 11, barcode compliant sequences: 11 (Fig. 105).
Hosts: Crambidae, Ategumia lotanalis DHJ07, Desmia ploralisDHJ01, Desmia Janzen07, Desmia Solis19 Diacme biolep02, Herpetogramma Solis10, Herpetogramma Solis11, Phostria Janzen05.
Costa Rica, ACG rain forest (
JFT is greatly indebted to John Huber and Caroline Boudreault (CNC) for helping with the preparation of many specimens used in this study. Thanks also go to Odette Morvan and Jean Cerda for their help for their long term collections in the “Montagne de Kaw”, the members of the “Société Entomologique Antilles Guyane” for their support in logistic and their collecting in several stations in French Guiana, and all other people who have kindly housed or supported the work of YB. The study has been supported by U.S. National Science Foundation grants BSR 9024770 and DEB 9306296, 9400829, 9705072, 0072730, 0515699 to DHJ, and grants from the Wege Foundation, International Conservation Fund of Canada, Jessie B. Cox Charitable Trust, Blue Moon Fund, Guanacaste Dry Forest Conservation Fund (GDFCF), Area de Conservación Guanacaste, Permian Global, and University of Pennsylvania (DHJ), and by the intense efforts of the ACG/GDFCF parataxonomist team that found and reared the parasitoids described here, to say nothing of the international taxasphere that labors to obtain and develop the caterpillar and parasitoid taxonomy essential to this effort in science and conservation. The reviews of Gavin Broad and two anonymous reviewers greatly improved the manuscript.
Supplementary information for studied specimens of Prasmodon (Hymenoptera: Braconidae, Microgastrinae) from ACG, Costa Rica
Authors: Jose L. Fernandez-Triana, James B. Whitfield, Alex M. Smith, Winnie Hallwachs, Daniel H. Janzen
Data type: Collection details, taxonomic information, DNA sequence, accession numbers
Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.