Zagrammosoma Ashmead

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, Zootaxa 4916 (1), pp. 1-108 : 12-29

publication ID	https://doi.org/ 10.11646/zootaxa.4916.1.1
publication LSID	lsid:zoobank.org:pub:803ED7FD-4A09-44F4-AAE2-6F70A0509E86
DOI	https://doi.org/10.5281/zenodo.4474152
persistent identifier	https://treatment.plazi.org/id/BC53327A-FFB1-6C2A-25CF-F91BD226C58C
treatment provided by	Plazi
scientific name	Zagrammosoma Ashmead
status

Treatment

Zagrammosoma Ashmead View in CoL View at ENA

Hippocephalus Ashmead, 1888: 7 . Type species: Hippocephalus multilineatus Ashmead ; by monotypy.

Zagrammosoma Ashmead, 1904: 354 View in CoL . Replacement name for Hippocephalus Ashmead , preoccupied by Hippocephalus Swainson, 1839 , later recognized as a junior synonym of Percis Scopoli, 1777 View in CoL ( Scorpaeniformes View in CoL : Agonidae View in CoL ).

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 View FIGURE 1 ), notaulus curving to meet the axilla and never reaching or approaching the posterior margin of the mesoscutum ( Fig. 1F View FIGURE 1 ), and median carina on the propodeum present ( Fig. 1F View FIGURE 1 ). 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 View FIGURE 1 ). 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 View FIGURE 1 ). 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 View FIGURE 8 , 23A View FIGURE 23 ) or present ( Fig. 1A View FIGURE 1 ). Occiput yellow with dark, vertical stripes or bands present or absent. Pronotum and mesoscutum dorsally variable: completely yellow ( Fig. 45D, E View FIGURE 45 ), completely dark ( Fig. 21D View FIGURE 21 ), with interrupted median line ( Fig. 1E, F View FIGURE 1 ), or with uninterrupted median line ( Figs 10D View FIGURE 10 , 12D View FIGURE 12 , 23D View FIGURE 23 ). 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 View FIGURE 36 ), yellow with an uninterrupted dark stripe on dorsal margin curving posterolaterally on outside of femur ( Fig. 18B View FIGURE 18 ), yellow with interrupted dark stripe - resulting in a “spot” on outside of femur ( Fig 4B View FIGURE 4 , 30B View FIGURE 30 ), with a wide dark band encircling femur ( Fig. 16B View FIGURE 16 ), or completely dark ( Fig. 28B View FIGURE 28 ). Gaster color variable: all yellow, yellow with varying lateral and longitudinal dark stripes or bands, or all dark brown or black.

Head: malar sulcus (ms) present ( Fig. 1A View FIGURE 1 ). Tentorial pits indistinct and not enlarged. Upper ocular sulcus (uos) variable: present ( Fig. 4A View FIGURE 4 ) or absent ( Fig. 48A View FIGURE 48 ). 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 View FIGURE 1 ); posterior ocelli (po) positioned lower on occiput ( Fig. 1I View FIGURE 1 ) (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 View FIGURE 1 ); 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 View FIGURE 1 ).

Body: pronotum (no 1) shape variable. Pronotal collar carina absent; bristle-like seta laterally. Propleura separated posteriorly, exposing a visible prodiscriminal groove ( Fig. 1I View FIGURE 1 ). Midlobe of mesoscutum (msc) sculpturing evenly reticulate ( Fig. 1F View FIGURE 1 ) or linearly reticulate ( Fig. 48 View FIGURE 48 D–F); one pair of bristle-like setae present posterolaterally, originating posteriorly to anterior margin of axilla ( Fig. 1E View FIGURE 1 ). 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 View FIGURE 1 ). Axilla advanced, more than 75% of longitudinal length, beyond posterior margin of mesoscutum ( Fig. 1F View FIGURE 1 ). Mesoscutellum (sct) with two pairs of bristle-like setae; posterior pair longer than anterior ( Fig. 1F View FIGURE 1 ); additional, unpaired bristle rarely found between anterior and posterior bristles ( Fig. 6D View FIGURE 6 inset); submedian grooves (smg) absent or present. Dorsellum not enlarged. Median carina (mc) on propodeum (ppd) present, sometimes faint, and posteriorly expanded ( Fig. 1F View FIGURE 1 ); plicae and costula absent. Propodeal spiracle (spr) rim complete and exposed ( Fig. 1F View FIGURE 1 ). 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 View FIGURE 1 ). 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 View FIGURE 1 ), 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 View FIGURE 1 ). 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 View FIGURE 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 View in CoL

1. Notaulus absent, present anteriorly, or complete and extending medially to anterior margin of mesoscutellum or posterior half of axilla ( Fig. 1D, G View FIGURE 1 ); if notaulus extends to advanced axilla, vertex not vaulted ( Fig. 1B View FIGURE 1 ) and/or median carina on propodeum absent ( Fig. 1G View FIGURE 1 )......................................................................... most Cirrospilini

– Notaulus complete and curving laterally to meet advanced axilla ( Fig. 1E, F View FIGURE 1 ), vertex vaulted ( Fig. 1A, I View FIGURE 1 ), and median carina on propodeum present ( Fig. 1F View FIGURE 1 ).............................................................. Zagrammosoma View in CoL , 2

2. Female gaster completely dark brown or black with no patterns ( Figs 6D View FIGURE 6 , 16D View FIGURE 16 , 19D View FIGURE 19 ); male gaster all dark or if yellow and black, then mesoscutellum either black with yellow center band ( Fig. 16E View FIGURE 16 ) or yellow with two black stripes converging posteriorly and with wide, black center band ( Fig. 44E View FIGURE 44 )................................................................... 3

– Female and male gaster completely yellow or yellow with varying dark color patterns and mesoscutellum not as above ( Figs 4 View FIGURE 4 , 8 View FIGURE 8 , 10 View FIGURE 10 , 30 View FIGURE 30 , 36 View FIGURE 36 , 40 View FIGURE 40 ).................................................................................... 10

3. Metallic luster present on body ( Fig. 32D View FIGURE 32 ) ….................................... Zagrammosoma metallicum n. sp.

– Metallic luster absent on body …......................................................................... 4

4. Hind wing with infuscation on apical margin ( Fig. 28E View FIGURE 28 ); fore wing with basal, cubital, parastigmal, stigmal, and postmarginal bands darkened and connected; apical band darkened ( Fig. 28C View FIGURE 28 )........................... Zagrammosoma lineaticeps View in CoL

– Hind wing hyaline ( Figs 18B View FIGURE 18 , 26B View FIGURE 26 ); fore wing pattern different than above ( Figs 6C View FIGURE 6 , 15C View FIGURE 15 , 19C View FIGURE 19 , 21C View FIGURE 21 ) …................ 5

5. Mesoscutellum all yellow ( Fig. 6D, E View FIGURE 6 ); known only from the Galapagos Islands ….............. Zagrammosoma buselus View in CoL

– Mesoscutellum all or mostly black ( Figs 19D, E View FIGURE 19 , 21D, E View FIGURE 21 ), black laterally ( Fig. 16D, E View FIGURE 16 ) or yellow with black bands ( Fig. 44E View FIGURE 44 ) ................................................................................................... 6

6. Procoxa and mesocoxa yellow ( Fig. 16B View FIGURE 16 ) …................................................................ 7

– Procoxa and mesocoxa dark ( Figs 19B View FIGURE 19 , 21B View FIGURE 21 ) …............................................................. 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 View FIGURE 16 )........................................................... Zagrammosoma flavolineatum View in CoL

– 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 View FIGURE 44 )......................................................... Zagrammosoma trifurcatum n. sp.

8. Mesoscutum yellow with dark median stripe present ( Fig. 15D, E View FIGURE 15 )....................... Zagrammosoma fisheri n. sp.

– Mesoscutum completely dark ( Fig. 21D, E View FIGURE 21 ) or mostly dark ( Fig. 19D, E View FIGURE 19 ) and lacking a distinct median stripe............ 9

9. Hind basitarsus longer than 2 nd tarsomere; occiput black with median yellow stripe ( Fig. 21D View FIGURE 21 ); propodeum all black ( Fig. 21D, E View FIGURE 21 ).............................................................................. Zagrammosoma hobbesi View in CoL

– Hind basitarsus subequal in length with 2 nd tarsomere; occiput all black, or with 5 stripes radiating from occipital foramen ( Fig. 19D View FIGURE 19 ); propodeum dark with yellow markings laterally ( Fig. 19D, E View FIGURE 19 )................... Zagrammosoma headricki n. sp.

10. Funiculars asymmetrical ( Fig. 48A View FIGURE 48 ); linear-reticulate sculpturing on mesoscutum and mesoscutellum ( Fig. 48E View FIGURE 48 )...................................................................................... Zagrammosoma yanegai n. sp.

– Funiculars symmetrical ( Fig. 1I View FIGURE 1 ); evenly reticulate sculpturing on mesoscutum and mesoscutellum ( Fig. 1E, F View FIGURE 1 )......... 11

11. Mesosoma with one wide, dark band extending from pronotum through mesoscutellum ( Figs 10D View FIGURE 10 , 26D View FIGURE 26 , 38D View FIGURE 38 ).......... 12

– Body pattern not as above - wide band absent ( Fig. 45D View FIGURE 45 ), or with single or multiple narrower stripes ( Figs 4D View FIGURE 4 , 8D View FIGURE 8 , 23D View FIGURE 23 ).. 14

12. Gaster with wide dark band continuing along entire dorsum in female ( Fig. 10D View FIGURE 10 ), brief interruption at base of gaster in males ( Fig. 10E View FIGURE 10 ); fore wing disc with wide dark band running parallel to anterior margin; postmarginal band complete across entire width of fore wing ( Fig. 10C View FIGURE 10 ).................................................. Zagrammosoma centrolineatum View in CoL

– Gaster with dark lateral stripes in females ( Figs 26D View FIGURE 26 , 38D View FIGURE 38 ) and dark spots dorsally in males ( Figs 26E View FIGURE 26 , 38E View FIGURE 38 ); fore wing disc hyaline, postmarginal band absent ( Figs 26C View FIGURE 26 , 38C View FIGURE 38 )......................................................... 13

13. Fore wing with darkening on apical margin ( Fig. 38C View FIGURE 38 ); propodeum mostly yellow, with dark areas medially and laterally ( Fig. 38D, E View FIGURE 38 ); known only from the Neotropic................................................. Zagrammosoma seini View in CoL

– Fore wing with no darkening on apical margin ( Fig. 26C View FIGURE 26 ); propodeum black, with small yellow patches laterally ( Fig. 26D, E View FIGURE 26 ); known only from Australasia and the Indo-Malay.................................... Zagrammosoma latilineatum View in CoL

14. Mesoscutum and mesoscutellum completely yellow with no longitudinal dark stripes ( Fig. 45D, E View FIGURE 45 ) … Zagrammosoma velerii View in CoL

– Mesoscutum with medial longitudinal stripe present, interrupted or uninterrupted; mesoscutellum with dark stripes or bands ( Figs 8D View FIGURE 8 , 34D View FIGURE 34 , 36D View FIGURE 36 )................................................................................. 15

15. Head, pronotum, mesoscutum, and gaster heavily setose ( Figs 8 View FIGURE 8 , 46 View FIGURE 46 )........................................... 16

– Head, pronotum, mesoscutum, and gaster sparsely setose ( Figs 4 View FIGURE 4 , 12 View FIGURE 12 , 23 View FIGURE 23 , 30 View FIGURE 30 , 36 View FIGURE 36 , 40 View FIGURE 40 ).............................. 17

16. Occiput with three dark stripes, sometimes faint ( Fig. 8A View FIGURE 8 ); mesosoma with three darks stripes in lateral view ( Fig. 8 View FIGURE 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 View FIGURE 46 ); known only from the Nearctic........................................................................... Zagrammosoma villosum n. sp.

17. Lower face with transverse stripe present ( Fig. 12A View FIGURE 12 ); female gaster with no median stripe connecting transverse stripes ( Fig. 12D View FIGURE 12 ); male gaster with one transverse, interrupted stripe ( Fig. 12E View FIGURE 12 ); known only from the Afrotropic.................................................................................................... Zagrammosoma crowei View in CoL

– Lower face with transverse stripe absent ( Figs 4A View FIGURE 4 , 23A View FIGURE 23 , 36A View FIGURE 36 , 40A View FIGURE 40 ); gaster with median stripe connecting transverse stripes ( Figs 14D View FIGURE 14 , 23D View FIGURE 23 , 40D View FIGURE 40 ); male as female, or with multiple, interrupted transverse stripes ( Figs 23E View FIGURE 23 , 25E View FIGURE 25 , 30F View FIGURE 30 ); known only from the Nearctic, Neotropic, or Palearctic.................................................................... 18

18. Prepectus and metacoxa completely yellow ( Fig. 36B View FIGURE 36 , 40B View FIGURE 40 ).................................................. 19

– Prepectus yellow with dark dorsal margin ( Figs 18B View FIGURE 18 , 30B View FIGURE 30 ); metacoxa yellow, dark basally.......................... 20

19. Hind basitarsus shorter than second tarsomere; parastigmal band present ( Fig. 36C View FIGURE 36 ); known only from the Nearctic................................................................................ Zagrammosoma occidentale n. sp.

– Hind basitarsus longer than second tarsomere; parastigmal band absent ( Fig. 40C View FIGURE 40 ); known only from the Palearctic........................................................................................ Zagrammosoma talitzkii View in CoL

20. Fore wing with broad medial patch ( Fig. 25C View FIGURE 25 ); female gaster with solid dark area dorsomedially ( Fig. 25D View FIGURE 25 ).......................................................................................... Zagrammosoma intermedium View in CoL

– Fore wing without broad medial patch ( Figs 18C View FIGURE 18 , 30C View FIGURE 30 , 34C View FIGURE 34 , 42C View FIGURE 42 ); female gaster pattern not as above - yellow or yellow with median and transverse stripes ( Figs 4D View FIGURE 4 , 18D View FIGURE 18 , 23D View FIGURE 23 , 34D View FIGURE 34 , 42D View FIGURE 42 )................................................ 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 View FIGURE 30 )..................................................................................... Zagrammosoma melinum View in CoL

– Pronotum, in dorsal view, bell shaped (1.05 <w:l <2.0); basal fold setose with more than two setae dorsally ( Figs 4C View FIGURE 4 , 18C View FIGURE 18 , 34C View FIGURE 34 ).............................................................................................. 22

22. Anterior margin of mesoscutum with dark, transverse stripe or band extending towards notaulus (sometimes hidden under posterior margin of pronotum) ( Fig. 4D View FIGURE 4 ), often appearing as brown spot extending towards notaulus ( Fig. 18D View FIGURE 18 )......... 23

– Anterior margin of mesoscutum with dark, transverse stripe or band absent or not extending towards notaulus ( Figs 14D View FIGURE 14 , 23D View FIGURE 23 ).............................................................................................. 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 View in CoL

– Anterior margin of mesoscutum with dark, transverse stripe not reaching notaulus ( Figs 4D View FIGURE 4 , 18D View FIGURE 18 ); second funicular wider than long …............................................................................................ 24

24. Posterior margin of mesoscutum with dark, transverse stripe present ( Fig. 18D, E View FIGURE 18 ); 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 View FIGURE 4 , 42D View FIGURE 42 ); mesoscutellum with submedian grooves present, often faint; known only from the Nearctic.......................................................... 25

25. Fore wing with triangular stigmal band ( Fig. 42C View FIGURE 42 ); intermediary setae absent; hind basitarsus shorter than 2 nd tarsomere............................................................................ Zagrammosoma triangulum n. sp.

– Fore with elliptical stigmal band ( Fig. 4C View FIGURE 4 ); intermediary setae present; hind basitarsus longer to subequal in length to 2 nd tarsomere......................................................................... Zagrammosoma americanum View in CoL

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 View FIGURE 14 ); 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 View FIGURE 23 ); known only from the Nearctic.............................. Zagrammosoma interlineatum View in CoL stat. rev.

Species of Zagrammosoma View in CoL

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.