12. Genus Tettigidea Scudder, 1862

Tettigidea is the biggest group of Batrachideinae, currently with forty species and some subspecies described (Cigliano et al., 2021). Tettigidea has a wide distribution, from North America (northwards to the southern Canada) to South America (southwards to northern Argentina) (Rehn & Grant 1961, Cigliano et al., 2021). The morphology aspect of these species also varies widely as the distribution of this group, since the species of Tettigidea may present brachypronotal or macropronotal forms in a same species, neotenic condition (Rehn & Grant, 1961) and a huge sexual dimorphism.

The most part of known species, with few exceptions for the North American ones, was sampled only few times, the morphological characteristics of these species are poorly understood (Grant, 1962). Tettigidea was never been reviewed and Grant (1962), possibly for the reasons presented here, in his PhD study considered only 12 valid species. Likewise, in our analysis, the number of valid species must also be reduced since, for some species the type specimen was not found in collections and there is no sufficient information in the literature.

Here we divide 19 well defined species of this genus into five species groups, while nine poorly known species are placed in Tettigidea nomina dubia, namely T. chichimeca (Saussure, 1861); T. exigua Kirby, 1910; T. guatemalteca Bolívar, 1887; T. neoaustralis Otte, 1997 (originally described as T. chichimeca australis Bruner, 1910); T. planovertex Hancock, 1913; T. plagiata Morse, 1900; T. imperfecta Bruner, 1906; T. trinitatis Bruner, 1906, and T. gracilis (Heer, 1865) and about the situation of each one:

(i) Tettigidea chichimeca (Saussure, 1861): there are no specimens found in the MHNG collection and the whereabouts of the type material is unknown (Hollier, 2013). Also, there are no detailed images available of this species and the morphological information of description used for identifying this species, as the color of the body, hind femur and tibia (see Hancock, 1902) not allowed distinguished this species from the other Tettigidea;

(ii) Tettigidea exigua Kirby, 1910: only one female was described by Bruner (1900) as Tettigidea gracilis Bruner, 1900 and was changed to T. exigua Kirby, 1910 by Kirby (1910). Neither of the authors designated a storage location of the type and the set of characteristics used in the description are shared with other species (also from other genera besides Tettigidea) (see Bruner, 1900);

(iii) Tettigidea guatemalteca Bolívar, 1887: this species also share characteristics used in the description with other species and, according to the original publication, the type series of this species would belong to the Brunner collection in the NHMW (Vienna) but no specimens have been located of the type series in this collection (Paris, 1994);

(iv) Tettigidea neoaustralis Otte, 1997: according to Grant (1957), this specimen was in poor condition, lacking antennae and right limbs. The holotype was not found at ANSP and the original description is unspecific;

(v) Tettigidea planovertex Hancock, 1913: only one female was described with unspecific and limited information that not allow the delimitation of this species. Also, there is no information about the type specimen;

(vi) Tettigidea plagiata Morse, 1900: only one male was described and there is no diagnostic morphological character and, also the holotype was not found at ANSP;

(vii) Tettigidea imperfecta Bruner, 1906: only one female and is a possible male juvenile of the same species were described by Bruner (1906). The holotype was not found at ANSP and the original description is unspecific;

(viii) Tettigidea trinitatis Bruner, 1906: Orthoptera type specimens deposited at CMNH Pittsburgh were exchanged for Lepidoptera with ANSP (Cadena-Castañeda & Cortés-Torres, 2013). The type specimen of this species is not found in either of the two collections, and the old description does not provide enough information to efficiently delimit it from the other species of the genus. On the other hand, the record of this species cited by Yong & Perez-Gelabert (2014) for Cuba is uncertain, so it is suggested to review the specimens studied by the authors again, since it could be a different taxon;

(iv) † Tettigidea gracilis (Heer, 1865): the whereabouts of the type specimen is unknown (Heads et al. 2014). The brief description and state of conservation of the fossil mentioned by the author of the species do not allow concluding if it belongs to the genus Tettigidea at least.

Thus, most of the specimens comprised here have older descriptions that do not contain exclusive morphological information (diagnosis) that allows recognizing the specimens even in situations where the type is unavailable. For this reason, these species were placed in the nomina dubia category. A key to the new species groups as well as the defined species is given below.

Key to the species groups and well-defined species of the genus Tettigidea

1. Middle of the anterior margin the pronotum armed with a spine with different degrees of development.................. 7

1’. Middle of the anterior margin of the pronotum at most angulate, but in nowise provided with a spine ( lateralis species group) ................................................................................................... 2

2. Tegminal sinus and tegmina absence...................................... T. empedonepia Hubbell, 1937 (Fig. 41)

2’. Tegmina present (occasionally only partly exposed), with a well-indicated tegminal sinus........................... 3

3. Head in lateral view elongated; in dorsal view, narrowing anteriorly, the anterior border angulate; tegmina elongated and narrow.............................................................................................. 4

3’. Head in lateral view robust and less elongated: in dorsal view not at all acuminate; tegmina mostly short or elongated, but broad, never narrow......................................................................................... 5

4. Moderately striated pronotal disc; tegmina with small yellowish spots, with a yellow spot near the tip and/or evident costal pale strip.................................................................. T. prorsa Scudder, 1877 (Figs. 42, 44)

4’. Strongly striated pronotal disc; tegmina with a small transverse stripe at the apex, without strips on the costal border............................................................................ T. spicatoides Hebard, 1932 (Fig. 45)

5. Vertex scarcely truncate, fascial carinae protruding; tegmina with an oblique strip near the apex; first and third valves of ovipositor with hook-like tip............................................................................ 6

5’. Vertex truncate, fascial carinae protruding; tegmina with a round or ovoid spot near the apex; first and third valves of ovipositor without hook-like tip........................................................ T. bruneri Morse, 1900 (Fig. 46)

6. Tegmina with a transverse stripe near the apex, this can vary and be an ovoid spot; winged and macropronotal form, with the posterior edge of the pronotum and hind wings exceeding or not the apex of the abdomen................................................................................................... T. lateralis (Say, 1824) (Figs. 47, 48)

6’. Tegmina marked with a minute light oblique line near apex; middle femur with an apical spine present; winged and macropronotal form, with the posterior edge of the pronotum and hind wings slightly exceeding the apex of the abdomen...................................................................................... T. australis Hancock, 1900 (Fig. 49)

7. Fascial carinae, surpassing the fastigium and vertex in lateral view; fastigium forming an almost right angle (90º); fascial carinae, frontal costa and fastigium produced, almost as a horn-like ( armata species group).......................... 8

7’. Fascial carinae not protruding, not or just slightly surpassing the fastigium and vertex in lateral view; fastigium not forming an almost right angle (90º), but more rounded with the vertex. Fascial carinae, frontal costa and fastigium slightly produced, but not as a horn-like..................................................................................... 9

8. Median carina of pronotum usually, in dorsal outline, without an emargination over the tegmina................................................................................................... T. acuta Morse, 1895 (Fig. 50)

8’. Median carina of pronotum usually, in dorsal outline, with an emargination over the tegmina....... T. armata Morse, 1895

9. Male and/or female body with strongly arched pronotum, for almost the entire length of the pronotum and median carina wellmarked ( paratecta species group)....................................................................... 10

9’. Male and/or female body without strongly arched pronotum, but slightly arched or straight for almost the entire length of the pronotum and median carina usually not well-marked....................................................... 12

10. Anterior margin of pronotum with a small spiniform projection, thin and elongated tegmina................................................................................................... T. paratecta Rehn, 1913 (Fig. 51)

10’. Anterior margin of pronotum completely hook-like, not thin or elongated tegmina................................. 11

11. Arched pronotum almost all its length, wings absent............. T. skejoi Cadena-Castañeda & Cardona, 2015 (Fig. 52)

11’. Arched pronotum in part of it length, wings surpassing the hind femurs............ T. arcuata Bruner, 1910 (Figs. 53, 54)

12. Body robust; facial carine and frontal costa notoriously thickened; spine of anterior edge of pronotum sharp and prominent; all known brachypronotal species; hind wings undeveloped ( cuspidata species group)................................ 13

12’. Body slender; facial carine and frontal slim; spine of anterior edge of pronotum slightly sharp and thin; most known macropronotal species; hind wings developed (except wingless form of T. intermedia) ( scudderi species group)..................... 16

13. Apex of pronotum not reaching the tip of hind femur; pronotal disc with many continuous straight lines of roughness in the dorsal view......................................................................................... 14

13’. Apex of pronotum only reaching the tip of hind femur; pronotal disc without continuous wavy lines of roughness in the dorsal view.................................................................... T. glabrata Bruner, 1920 (Fig. 55)

14. General body coloration uniform, light brown; eyes of medium size, occupying a quarter of the cephalic capsule, in lateral view not covering the vertex; midline of the pronotum elevated; only females of the species are known......................................................................................... T. cuspidata Scudder, 1875 (Figs. 56,58)

14’. Differentiated body color, yellowish brown pronotal disc, the rest of the body brown or black. Conspicuous eyes, occupying a third of the cephalic capsule, in lateral view, partially covering the vertex. Midline of the pronotum moderately elevated.... .................................................................................................. 15

15. Spine of the anterior edge of the pronotum small, without covering the vertex. Apical internal spurs of the hind tibia welldeveloped; male Terminalia normally developed...................... T. angustihumeralis Podgornaya, 1999 (Fig. 59)

15’. Spine of the anterior edge of the pronotum longer than in the previous species, almost completely covering the vertex; apical internal spurs of the hind tibia moderately developed; male with a bulging and prominent terminalia....................................................................... T. parradae Cadena-Castañeda & Cardona, 2015 (Fig. 60)

16. Apex of pronotum not quite reaching the tip of the hind femora................................................ 17

16’. Apex of pronotum surpassing notoriously the tip of the hind femora............................................ 18

17. Pronotum without lateral carinae............................................... T. pulchella Rehn, 1904 (Fig. 61)

17’. Pronotum with lateral carinae parallel.................................... T. intermedia Bruner, 1910 (Figs. 62, 65)

18. Pronotal disc with several conspicuous and parallel lateral carinae; tegmina without a spot or conspicuous strip close to the apex, occasionally with very faintly paler apically............................. T. scudderi Bolívar, 1887 (Figs. 66, 67)

18’. Pronotal disc with slightly conspicuous lateral carinae and not entirely parallel; tegmina with a small, subapical macula...................................................................... T. chapadensis Bruner, 1910 (Figs. 68-71)