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Subfamily Romaleinae - Lubber Grasshoppers

Romalea guttata - Romalea microptera - female Lubber Grasshopper - Romalea microptera - female Robust Toad Lubber - Phrynotettix robustus - female Spanistic Desert Grasshopper - Spaniacris deserticola - female Dragon Lubber - 4th Instar - Dracotettix monstrosus - female Chihuahua Lubber - Phrynotettix tshivavensis - female Phrynotettix tshivavensis - female Horse Lubber (Taeniopoda eques)  - Taeniopoda eques - male - female
Classification
Kingdom Animalia (Animals)
Phylum Arthropoda (Arthropods)
Subphylum Hexapoda (Hexapods)
Class Insecta (Insects)
Order Orthoptera (Grasshoppers, Crickets, Katydids)
Suborder Caelifera (Grasshoppers)
Family Acrididae (Short-horned Grasshoppers)
Subfamily Romaleinae (Lubber Grasshoppers)
Synonyms and other taxonomic changes
Names listed here are only relevant to North American genera - there are many other related names involving genera found further south.

Acrididae Romaliae Brunner von Wattenwyl, 1893 [rank not given, but between tribe and genus - basically a "genus group"]. Based on genus Romalea Serville, 1831
Acrididae Taeniopodae Brunner von Wattenwyl, 1893 [rank not given, but between tribe and genus - basically a "genus group"]. Based on genus Taeniopoda Stål, 1873
Acrididae Locustinae Romali Blatchley, 1920 [as tribe]
Romaleidae (Brunner von Wattenwyl) H.R. Roberts, 1941 [as family]
Acrididae Pamphaginae Phrynotettigini Uvarov, 1943 [as tribe]. Based on genus Phrynotettix Glover, 1872
Acrididae Romaleinae Brachystolini Rehn & Grant, 1959 [as tribe]. Based on genus Brachystola S.H. Scudder, 1876
Acrididae Romaleinae Dracotettigini Rehn & Grant, 1959 [as tribe]. Based on genus Dracotettix L. Bruner, 1889
Acrididae Romaleinae Phrynotettigini (Uvarov) Rehn & Grant, 1959 [as tribe]
Acrididae Romaleinae Spaniacrini Rehn & Grant, 1961 [as tribe]. Based on genus Spaniacris Hebard, 1937

Many described tribes of the Romaleidae [there are many more than listed above] are based on only one or few genera, and the subfamily is probably "over-split". Conversely Orthoptera Species Files currently lists all of our genera (and most others) as members of the single tribe Romaleini, although this is likely an unnaturally inclusive grouping. Perhaps Brachystola, Phrynotettix, Spaniacris, Tytthotyle, and Dracotettix should indeed be excluded and placed in other tribes.
Explanation of Names
New Latin (Serville, 1831), properly Rhomalea, from Greek ρωμαλεοσ, strong of body, from ρωμη, bodily strength (1). English Lubber means clumsy and stout (2).
Romaleinae has been treated both as a full family, and as a subfamily of the Acrididae. Orthoptera Species File Online lists as family Romaleidae Brunner von Wattenwyl, 1893. However, there is no individual morphological character that separates this as uniquely distinct from all other groups within the family Acrididae, and all recent molecular DNA studies place this group well within the Acrididae. In these studies, splitting Romaleinae out as a family causes the remainder of family Acrididae to become "paraphylletic". There is also some indication that the group is not entirely natural and may include some genera that are not particularly closely related to one another. These are areas that need further study.
Pamphiginae is another distinctive (Old World) subfamily of Acrididae that has been treated by some authors at family ranking. Certain "Toad-like" American genera, including several Romaleinae, have in past been placed in the Pamphiginae due to their close physical resemblance, but later study has shown these placements to be in error.
Numbers
There are 7 genera and 9 species in North America north of Mexico. Approximately 100 additional genera occur further south.
Orthoptera Species File treats Romaleidae as a family; lists two subfamilies, with our genera all included within the Romaleinae (Brunner von Wattenwyl, 1893).
Rehn and Hebard (1959 & 1961) treated our genera as belonging to five different tribes within subfamily Romaleinae of family Acrididae (Romalea & Taeniopoda together; Phrynotettix & Tytthotyle together; and Dracotettix, Spaniacris, and Brachystola each in separate tribes.
Size
18-145 mm
The largest Grasshopper species known belong to this subfamily, and occur in the American tropics. The largest recorded are females of Tropidacris cristata; not only long, but with spread wings reaching 25 cm from wing tip to wing tip. These very large species are not found in North America north of Mexico. Our largest species of Lubber Grasshoppers will probably not exceed 6 cm. in length - but are still quite large.
Identification
Many species are so distinctive as to be recognizable on sight. Some southwestern species look remarkably like Band-wing Grasshoppers (Oedipodinae), often with roughened surfaces to the body, sometimes with colored wings, and with cryptic coloration that blends with their surroundings. Even these species are relatively easy to identify, if one remembers that all colored-winged, and/or cryptic ground mimicing grasshoppers are not in the Oedipodinae.
No single character is unique to the subfamily and applies to all members of the group (except perhaps certain traits of genitalia); however, the following are shared by many or most species:
Most species are heavy-bodied, often with shortened wings, often brightly marked. However, Spaniacris is very slender and almost Spider-like; Spaniacris, Tytthotyle, and sometimes Taeniopoda have long fully developed wings.
Head typically broadly rounded and face nearly vertical. Dracotettix is an exception with a prominent protrusion between the eyes, and Spaniacris deserticola has a narrow, somewhat angular head.
Often (i.e. Dracotettix, Romalea and Taeniopoda, and most tropical species) with pronounced "prosternal spine" between the front legs (charater shared with "Spurthroat" subfamilies Cyrtacanthacridinae, Melanoplinae, Leptysminae, Copiocerinae, etc.), but not with "Slant-face" and "Band-wing" subfamilies (Gomphocerinae, Hyaloteryginae, Oedipodinae).
Two rows of immovable spine on hind tibiae relatively large and stout; both the inner and outer immovable spine present at tip of the hind tibia (North American exception is Spaniacris deserticola with only the inner spine at tip). Other subfamilies in North America have only the inner, not the outer. Don't confuse with longer pair of movable spurs or "calcaria" at tip of femur below the immovable spines.
Two basal lobes of the hind femur approximately the same length--compare other Acrididae, where dorsal lobe is usually larger and longer.(Orthoptera of the Northern Great Plains).
Hind femora generally slender, often smallish and looking rather undersized. However, among North American genera not a consistant trait, being noticeably short only in Phrynotettix (though a little under-sized in Romalea & Taeniopoda); not particularly slender in Dracotettix & Phrynotettix; and, noticeably large and thickened in Brachystola.
Forewings often with a course mesh-like appearance due to prominent veins. Sometimes forewings (notably in Brachystola) may be brightly colored
Hind wings (when developed) with an area modified for sound production, involving areas of modified venation and an area that is raised along one of the anterior main veins (does not lay flat when wings are spread). This apparently forms a resonating chamber when the wings are folded. It is poorly developed but present in Spaniacris & Tytthotyle, and well developed and obvious in Romalea & Taeniopoda. Hindwings most often red bluish, or greenish, and usually blackish toward outer margin or tip [nearly clear in Spaniacris].
Range
An American, mostly tropical group, several species in southwestern United States, one in Great Plains (Brachystola magna), and one in southeast (Romalea microptera).
Remarks
Most species tend to be less agile on the ground than a majority of other grasshoppers, sometimes downright clumsy, but those capable of flight can be quite adept at making an escape, and not all are lubberly and clumsy (i.e. Spaniacris & Tytthotyle).
This subfamily (or family) is highly diverse, and perhaps not monophylletic. Recent molecular studies show it to be "nested" within the traditional family Acrididae, and not separable as a distinct family group [unless Acrididae is much further divided]. Romaleinae is a composite of groups of genera that share certain anotomical features, but because of the huge diversity in morphology and behavior, generalizations about the Romaleinae are difficult to make. For example the cryptic rock-like species with no prosternal spine (genera Phrynotettix & Tytthotyle) seem to share only a little in common with the large smooth almost polished "typical" Lubbers that have a prominent prosternal spine (genera Romalea & Taeniopoda); the two groups are probably related, but perhaps only distantly so. It is likely that ongoing molecular studies will result in some rearrangement of the included genera, and some genera may be removed from the group. At least some tropical genera seem to be related to the South American subfamily [or family] Ommexechinae.

First version of this page contributed by Troy Bartlett as family Romeidae in 2004
Print References
The Century Dictionary--entry for Romalea (1)
Arnett, p. 169 (3)
Capinera, Field Guide to Grasshoppers,..., p. 148 (2)
Flook, P. K. & Rowell, C. H. F. 1997. The phylogeny of the Caelifera (Insecta, Orthoptera) as deduced from mitochondrial rRNA gene sequences. Molecular Phylogenetics & Evolution 8: 89-103.
Flook, P. K. & Rowell, C. H. F. 1998. Inferences about orthopteroid phylogeny and molecular evolution from small subunit nuclear ribosomal RNA sequences. Insect Mol. Biol. 7: 163-178.
Flook, P. K. Klee, S., & Rowell, C. H. F. 1999. A combined molecular phylogenetic analysis of the Orthoptera and its implications for their higher systematics. Syst. Biol. 48: 233-253.
Rowell, C.H.F., and P.K. Flook, Dec.1998. 'Phylogeny of Caelifera and the Orthoptera as Derived from Ribosomal Gene Sequences', Journal of Orthoptera Research 7, 147-156.