Zagrammosoma Ashmead
Creators: name: Perry, Ryan K.
name: Heraty, John M.
Publisher: name: Zenodo
Year: 2021
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Hymenoptera
Eulophidae
Zagrammosoma

description: Zagrammosoma Ashmead
Hippocephalus Ashmead, 1888: 7. Type species: Hippocephalus multilineatus Ashmead; by monotypy.
Zagrammosoma Ashmead, 1904: 354. Replacement name for Hippocephalus Ashmead, preoccupied by Hippocephalus Swainson, 1839, later recognized as a junior synonym of Percis Scopoli, 1777 (Scorpaeniformes: Agonidae).
Zagrammatosoma Schulz, 1906: 142. Unjustified emendation. Corrected by Bouček, 1959: 172.
Mirzagrammosoma Girault, 1915: 279. Type species Mirzagrammosoma lineaticeps Girault; by monotypy. Synonymy by La-Salle, 1989: 232.
Diagnosis. Based on the present analysis, Zagrammosoma is characterized by a combination of 3 characters: vaulted vertex that extends above the compound eyes (Fig. 1A), notaulus curving to meet the axilla and never reaching or approaching the posterior margin of the mesoscutum (Fig. 1F), and median carina on the propodeum present (Fig. 1F). Species that are mostly yellow in color generally have dark lines between sclerites, especially along the notaulus, axilla, mesoscutellum, and prepectus.
Within Cirrospilini, Zagrammosoma is most similar, in terms of morphology and color patterns, to Cirrospilus and Burkseus; however, these 2 genera both possess a notaulus that is complete and extends to the posterior margin of the mesoscutum (Fig. 1D, G). Burkseus and some species of Cirrospilus (C. ambiguus, C. margiscutellum, C. sp. 4154, C. sp. D4977) also lack a median carina on the propodeum (Fig. 1G). Pseudiglyphus Girault have a notaulus that curves to meet the axilla, but lack a vaulted vertex, and are drab brown in coloration, with hyaline wings.
Description. Body length: female 0.75–2.9 mm; male 0.63–1.62 mm. Overall body coloration variable: mainly yellow heads and bodies, with varying degrees of dark lines and bands present, or mainly dark heads and bodies, with yellow markings either on the head or dorsal side of mesosoma. Metallic body coloration absent, except in Zagrammosoma metallicum n. sp. Lower face (lof) with dark, median stripe absent (Figs 8A, 23A) or present (Fig. 1A). Occiput yellow with dark, vertical stripes or bands present or absent. Pronotum and mesoscutum dorsally variable: completely yellow (Fig. 45D, E), completely dark (Fig. 21D), with interrupted median line (Fig. 1E, F), or with uninterrupted median line (Figs 10D, 12D, 23D). Fore wing with or without dark patterns on wing disc. Dark coloration at junctions between parastigmal vein and marginal vein, and between marginal vein and postmarginal vein always present. Hind wing hyaline, rarely with infuscation along apical margin. Procoxa, mesocoxa, and metacoxa color variable: all yellow, basally dark, or completely dark. Profemur yellow. Mesofemur usually all yellow, rarely dark basally. Metafemur coloration variable: all yellow (Fig. 36B), yellow with an uninterrupted dark stripe on dorsal margin curving posterolaterally on outside of femur (Fig. 18B), yellow with interrupted dark stripe - resulting in a “spot” on outside of femur (Fig 4B, 30B), with a wide dark band encircling femur (Fig. 16B), or completely dark (Fig. 28B). Gaster color variable: all yellow, yellow with varying lateral and longitudinal dark stripes or bands, or all dark brown or black.
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Head: malar sulcus (ms) present (Fig. 1A). Tentorial pits indistinct and not enlarged. Upper ocular sulcus (uos) variable: present (Fig. 4A) or absent (Fig. 48A). Vertex (vtx) vaulted, forming angular apex of head, creating distinct demarcation of frons and occiput. Anterior ocellus (ao) positioned on occiput adjacent to apex of vertex (Fig. 1I); posterior ocelli (po) positioned lower on occiput (Fig. 1I) (in dried specimens, the vertex will sometimes collapse, making the anterior ocellus appear at the vertex, or on frons). Vertex with scattered, short setae present (Fig. 1A); paired large setae absent. Occipital carina absent.
Antenna: dorsal margin of torulus above lower margin of compound eye. Scape and pedicel setose; scape not extending to vertex; pedicel length variable, longer than wide or subequal. Pedicel lacking carinae. Funiculars with linear sensilla. Two funiculars (fu), always cylindrical, with variable width to length ratio. Clava (clv) longer than wide, consisting of three clavomeres (Fig. 1I).
Body: pronotum (no 1) shape variable. Pronotal collar carina absent; bristle-like seta laterally. Propleura separated posteriorly, exposing a visible prodiscriminal groove (Fig. 1I). Midlobe of mesoscutum (msc) sculpturing evenly reticulate (Fig. 1F) or linearly reticulate (Fig. 48 D–F); one pair of bristle-like setae present posterolaterally, originating posteriorly to anterior margin of axilla (Fig. 1E). Lateral lobe of mesoscutum with one bristle-like seta and small, scattered setae. Notaulus (not) complete and curved to meet anterior margin of axilla (axl) (Fig. 1F). Axilla advanced, more than 75% of longitudinal length, beyond posterior margin of mesoscutum (Fig. 1F). Mesoscutellum (sct) with two pairs of bristle-like setae; posterior pair longer than anterior (Fig. 1F); additional, unpaired bristle rarely found between anterior and posterior bristles (Fig. 6D inset); submedian grooves (smg) absent or present. Dorsellum not enlarged. Median carina (mc) on propodeum (ppd) present, sometimes faint, and posteriorly expanded (Fig. 1F); plicae and costula absent. Propodeal spiracle (spr) rim complete and exposed (Fig. 1F). Paraspiracular sulcus and carina absent. Gaster sessile.
Legs: Profemur and mesofemur with preapical bristle on anterior surface present. Metafemur with preapical bristle on anterior surface absent. Hind tibial spur (tbs) shorter than or equal to basitarsus (bst) in length (Fig. 1C). Hind basitarsus length variable: shorter than 2 nd tarsomere, subequal, or longer than 2 nd tarsomere.
Fore wing: Submarginal vein smoothly joining parastigmal; 3–9 setae dorsally. Costal cell setose. Admarginal setae (ams) present in one (Fig. 1H), or, rarely, two rows. Uncus (unc) position variable: originating on stigma by less than, more than, or its own length from stigmal apex (Fig. 1H). Postmarginal vein present. Basal fold setose, sometimes with only one or two setae. Cubital fold setose, with setal line complete to basal vein or absent anteriorly.
Hosts. Zagrammosoma parasitizes leaf miners and leaf rollers in Diptera, Coleoptera, Hymenoptera, and Lepidoptera (Table 3). Gordh (1978) states that Zagrammosoma has been recovered as a hyperparasite of hymenopteran leaf-miner parasitoids; however, he was referring to Z. nigrolineatum as the hyperparasitic species, which is now a synonym of Burkseus vittatus (Perry & Heraty, 2019). Nystrom & Evans (1989) and Digweed (1998) record Z. multilineatum and Z. sp., respectively, emerging from hymenopteran Birch leafminers (Tenthredinidae). While it is difficult to determine the actual species recovered without direct examination of the specimens, it is accurate to record Zagrammosoma as a parasitoid of hymenopteran leaf miners. One examined specimen of Z. multilineatum was reared from the sawfly Metallus sp. (Tentredinidae) (see species description).
There are several individual host records of Z. americanum emerging from Hemiptera (Table 3), but more evidence is needed to prove this concretely. This could be Zagrammosoma emerging in bulk samples of hemipterans, which may have contained concealed or unnoticed leaf miners in the collected plant material. If this is true, these individuals may be acting as hyperparasitoids, as Zagrammosoma has not previously been recorded parasitizing non-holometabolous insects.
Zagrammosoma has been reared from a large number of agricultural crops, including alfalfa, apple, citrus, coffee, cotton, corn, jojoba, muscadine, potato, and tomato, as well as plants grown for ornamental and timber purposes, including aspen, oak, poplar, pine, and willow. Table 3 presents a full list of host and host plants by species based on the material examined and publications with unambiguously identifiable species of Zagrammosoma.
Distribution. The majority of Zagrammosoma species are found in the Nearctic, with only a few species found in each of the remaining ecozones: Afrotropical (1), Australasian (1), Indo-Malay (2), Nearctic (16), Neotropical (9), Oceania (1), and Palearctic (1) (number of species in each ecozone noted in parentheses). Five of the 9 species found in the Neotropical region (Z. lineaticeps, Z. multilineatum, Z. occidentale, Z. trifurcatum, Z. villosum) are also found in the Nearctic, while Z. flavolineatum is found in both Oceania (Hawaii) and the Nearctic. Coupled with the results from the ML analysis (Fig. 2), this may suggest origins of Zagrammosoma outside the Nearctic or Neotropic, with rapid diversification and speciation occurring after introduction into the New World.
Etymology. The original designation for the genus, Hippocephalus, means “ horse head ” and accurately describes the shape of the head in lateral view. Ashmead (1904) did not specify why he named Zagrammosoma as he did, but it translates from Greek to mean “body with a lot of writing.” This is appropriate to describe a group of “tattooed wasps,” the official common name for Zagrammosoma, approved by the Entomological Society of America, April 2019 (https://www.entsoc.org/sites/default/files/files/common_name.pdf). As Zagrammosoma is neuter, adjectival species names must also be neuter in gender.
Key to species of Zagrammosoma
1. Notaulus absent, present anteriorly, or complete and extending medially to anterior margin of mesoscutellum or posterior half of axilla (Fig. 1D, G); if notaulus extends to advanced axilla, vertex not vaulted (Fig. 1B) and/or median carina on propodeum absent (Fig. 1G)......................................................................... most Cirrospilini
– Notaulus complete and curving laterally to meet advanced axilla (Fig. 1E, F), vertex vaulted (Fig. 1A, I), and median carina on propodeum present (Fig. 1F).............................................................. Zagrammosoma, 2
2. Female gaster completely dark brown or black with no patterns (Figs 6D, 16D, 19D); male gaster all dark or if yellow and black, then mesoscutellum either black with yellow center band (Fig. 16E) or yellow with two black stripes converging posteriorly and with wide, black center band (Fig. 44E)................................................................... 3
– Female and male gaster completely yellow or yellow with varying dark color patterns and mesoscutellum not as above (Figs 4, 8, 10, 30, 36, 40).................................................................................... 10
3. Metallic luster present on body (Fig. 32D) ….................................... Zagrammosoma metallicum n. sp.
– Metallic luster absent on body …......................................................................... 4
4. Hind wing with infuscation on apical margin (Fig. 28E); fore wing with basal, cubital, parastigmal, stigmal, and postmarginal bands darkened and connected; apical band darkened (Fig. 28C)........................... Zagrammosoma lineaticeps
– Hind wing hyaline (Figs 18B, 26B); fore wing pattern different than above (Figs 6C, 15C, 19C, 21C) …................ 5
5. Mesoscutellum all yellow (Fig. 6D, E); known only from the Galapagos Islands ….............. Zagrammosoma buselus
– Mesoscutellum all or mostly black (Figs 19D, E, 21D, E), black laterally (Fig. 16D, E) or yellow with black bands (Fig. 44E) ................................................................................................... 6
6. Procoxa and mesocoxa yellow (Fig. 16B) …................................................................ 7
– Procoxa and mesocoxa dark (Figs 19B, 21B) …............................................................. 8
7. Hind basitarsus shorter than 2 nd tarsomere; mesoscutellum black with yellow center band; male gaster with 50% of dorsal side darkened (Fig. 16E)........................................................... Zagrammosoma flavolineatum
– Hind basitarsus longer than 2 nd tarsomere; mesoscutellum black with yellow lateral margin; male gaster with median band from Gt 2 –Gt 6 (Fig. 44E)......................................................... Zagrammosoma trifurcatum n. sp.
8. Mesoscutum yellow with dark median stripe present (Fig. 15D, E)....................... Zagrammosoma fisheri n. sp.
– Mesoscutum completely dark (Fig. 21D, E) or mostly dark (Fig. 19D, E) and lacking a distinct median stripe............ 9
9. Hind basitarsus longer than 2 nd tarsomere; occiput black with median yellow stripe (Fig. 21D); propodeum all black (Fig. 21D, E).............................................................................. Zagrammosoma hobbesi
– Hind basitarsus subequal in length with 2 nd tarsomere; occiput all black, or with 5 stripes radiating from occipital foramen (Fig. 19D); propodeum dark with yellow markings laterally (Fig. 19D, E)................... Zagrammosoma headricki n. sp.
10. Funiculars asymmetrical (Fig. 48A); linear-reticulate sculpturing on mesoscutum and mesoscutellum (Fig. 48E)...................................................................................... Zagrammosoma yanegai n. sp.
– Funiculars symmetrical (Fig. 1I); evenly reticulate sculpturing on mesoscutum and mesoscutellum (Fig. 1E, F)......... 11
11. Mesosoma with one wide, dark band extending from pronotum through mesoscutellum (Figs 10D, 26D, 38D).......... 12
– Body pattern not as above - wide band absent (Fig. 45D), or with single or multiple narrower stripes (Figs 4D, 8D, 23D).. 14
12. Gaster with wide dark band continuing along entire dorsum in female (Fig. 10D), brief interruption at base of gaster in males (Fig. 10E); fore wing disc with wide dark band running parallel to anterior margin; postmarginal band complete across entire width of fore wing (Fig. 10C).................................................. Zagrammosoma centrolineatum
– Gaster with dark lateral stripes in females (Figs 26D, 38D) and dark spots dorsally in males (Figs 26E, 38E); fore wing disc hyaline, postmarginal band absent (Figs 26C, 38C)......................................................... 13
13. Fore wing with darkening on apical margin (Fig. 38C); propodeum mostly yellow, with dark areas medially and laterally (Fig. 38D, E); known only from the Neotropic................................................. Zagrammosoma seini
– Fore wing with no darkening on apical margin (Fig. 26C); propodeum black, with small yellow patches laterally (Fig. 26D, E); known only from Australasia and the Indo-Malay.................................... Zagrammosoma latilineatum
14. Mesoscutum and mesoscutellum completely yellow with no longitudinal dark stripes (Fig. 45D, E) … Zagrammosoma velerii
– Mesoscutum with medial longitudinal stripe present, interrupted or uninterrupted; mesoscutellum with dark stripes or bands (Figs 8D, 34D, 36D)................................................................................. 15
15. Head, pronotum, mesoscutum, and gaster heavily setose (Figs 8, 46)........................................... 16
– Head, pronotum, mesoscutum, and gaster sparsely setose (Figs 4, 12, 23, 30, 36, 40).............................. 17
16. Occiput with three dark stripes, sometimes faint (Fig. 8A); mesosoma with three darks stripes in lateral view (Fig. 8); known only from South America.................................................................. Z. calvini n. sp.
– Occiput with two dark stripes; mesosoma with two dark stripes in lateral view (Fig. 46D, E); known only from the Nearctic........................................................................... Zagrammosoma villosum n. sp.
17. Lower face with transverse stripe present (Fig. 12A); female gaster with no median stripe connecting transverse stripes (Fig. 12D); male gaster with one transverse, interrupted stripe (Fig. 12E); known only from the Afrotropic.................................................................................................... Zagrammosoma crowei
– Lower face with transverse stripe absent (Figs 4A, 23A, 36A, 40A); gaster with median stripe connecting transverse stripes (Figs 14D, 23D, 40D); male as female, or with multiple, interrupted transverse stripes (Figs 23E, 25E, 30F); known only from the Nearctic, Neotropic, or Palearctic.................................................................... 18
18. Prepectus and metacoxa completely yellow (Fig. 36B, 40B).................................................. 19
– Prepectus yellow with dark dorsal margin (Figs 18B, 30B); metacoxa yellow, dark basally.......................... 20
19. Hind basitarsus shorter than second tarsomere; parastigmal band present (Fig. 36C); known only from the Nearctic................................................................................ Zagrammosoma occidentale n. sp.
– Hind basitarsus longer than second tarsomere; parastigmal band absent (Fig. 40C); known only from the Palearctic........................................................................................ Zagrammosoma talitzkii
20. Fore wing with broad medial patch (Fig. 25C); female gaster with solid dark area dorsomedially (Fig. 25D).......................................................................................... Zagrammosoma intermedium
– Fore wing without broad medial patch (Figs 18C, 30C, 34C, 42C); female gaster pattern not as above - yellow or yellow with median and transverse stripes (Figs 4D, 18D, 23D, 34D, 42D)................................................ 21
21. Pronotum, in dorsal view, subequal (w:l = 0.95–1.05); basal fold asetose or with one or two setae dorsally (Fig. 30C)..................................................................................... Zagrammosoma melinum
– Pronotum, in dorsal view, bell shaped (1.05 <w:l <2.0); basal fold setose with more than two setae dorsally (Figs 4C, 18C, 34C).............................................................................................. 22
22. Anterior margin of mesoscutum with dark, transverse stripe or band extending towards notaulus (sometimes hidden under posterior margin of pronotum) (Fig. 4D), often appearing as brown spot extending towards notaulus (Fig. 18D)......... 23
– Anterior margin of mesoscutum with dark, transverse stripe or band absent or not extending towards notaulus (Figs 14D, 23D).............................................................................................. 26
23. Anterior margin of mesoscutum with dark, transverse stripe complete to notaulus (sometimes hidden under posterior margin of pronotum); second funicular longer than wide...................................... Zagrammosoma multilineatum
– Anterior margin of mesoscutum with dark, transverse stripe not reaching notaulus (Figs 4D, 18D); second funicular wider than long …............................................................................................ 24
24. Posterior margin of mesoscutum with dark, transverse stripe present (Fig. 18D, E); mesoscutellum with submedian grooves absent; known only from the Galapagos ...................................... Zagrammosoma galapagoense n. sp.
– Posterior margin of mesoscutum with dark, transverse stripe absent (Figs 4D, E, 42D); mesoscutellum with submedian grooves present, often faint; known only from the Nearctic.......................................................... 25
25. Fore wing with triangular stigmal band (Fig. 42C); intermediary setae absent; hind basitarsus shorter than 2 nd tarsomere............................................................................ Zagrammosoma triangulum n. sp.
– Fore with elliptical stigmal band (Fig. 4C); intermediary setae present; hind basitarsus longer to subequal in length to 2 nd tarsomere......................................................................... Zagrammosoma americanum
26. Second funicular subequal in width: length; parastigmal band extending well past apex of stigmal vein, curving posteriorly and almost connecting with stigmal band (Fig. 14C); known only from the Neotropic............ Zagrammosoma deliae n. sp.
– Second funicular longer than wide; parastigmal band extending just past apex of stigmal vein, not closely approaching stigmal band (Fig. 23C); known only from the Nearctic.............................. Zagrammosoma interlineatum stat. rev.
Species of Zagrammosoma
A list of all described species of Zagrammosoma is found in Table 4.
1. Zagrammosoma americanum Girault
2. Zagrammosoma buselus (Walker)
3. Zagrammosoma calvini Perry n. sp.
4. Zagrammosoma centrolineatum Crawford
5. Zagrammosoma crowei (Kerrich)
6. Zagrammosoma deliae Perry n. sp.
7. Zagrammosoma fisheri Perry n. sp.
8. Zagrammosoma flavolineatum Crawford
9. Zagrammosoma galapagoense Perry n. sp.
10. Zagrammosoma headricki Perry n. sp.
11. Zagrammosoma hobbesi LaSalle
12. Zagrammosoma interlineatum Girault stat. rev.
13. Zagrammosoma intermedium Gordh
14. Zagrammosoma latilineatum Ubaidillah
15. Zagrammosoma lineaticeps (Girault)
16. Zagrammosoma melinum Gordh
17. Zagrammosoma metallicum Perry n. sp.
18. Zagrammosoma multilineatum (Ashmead)
19. Zagrammosoma occidentalum Perry n. sp.
20. Zagrammosoma seini Wolcott
21. Zagrammosoma talitzkii (Bouček)
22. Zagrammosoma triangulum Perry n. sp.
23. Zagrammosoma trifurcatum Perry n. sp.
24. Zagrammosoma velerii Yefremova
25. Zagrammosoma villosum Perry n. sp.
26. Zagrammosoma yanegai Perry n. sp.
description: Published as part of Perry, Ryan K. & Heraty, John M., 2021, Read between the lineata: A revision of the tattooed wasps, Zagrammosoma Ashmead (Hymenoptera: Eulophidae), with descriptions of eleven new species, pp. 1-108 in Zootaxa 4916 (1) on pages 12-29, DOI: 10.11646/zootaxa.4916.1.1, http://zenodo.org/record/4471803