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Orthoptera: Acrididae) 204 Florida Entomologist 88(2) June 2005 MANDIBULAR MORPHOLOGY OF SOME FLORIDIAN GRASSHOPPERS (ORTHOPTERA: ACRIDIDAE) TREVOR RANDALL SMITH AND JOHN L. CAPINERA University of Florida, Department of Entomology and Nematology, Gainesville, FL 32611 The relationship between mouthpart structure zen until examination. Mandibles were removed and diet has been known for years. This connec- from thawed specimens by lifting the labrum and tion between mouthpart morphology and specific pulling out each mandible separately with for- food types is incredibly pronounced in the class In- ceps. Only young adults were used in an effort to secta (Snodgrass 1935). As insects have evolved avoid confusion of mandible type due to mandible and adapted to new food sources, their mouthparts erosion (Chapman 1964; Uvarov 1977). An exam- have changed accordingly. This is an extremely im- ple of moderate erosion can be seen in Figure 1 (I). portant trait for evolutionary biologists (Brues This process was replicated with 10 individuals 1939) as well as systematists (Mulkern 1967). from each species. After air-drying, each mandi- Isley (1944) was one of the first to study grass- ble was glued to the head of a #3 or #2 insect pin, hopper mouthparts in detail. He described three depending on its size, for easier manipulation, groups of mandibles according to general struc- and examined microscopically. ture and characteristic diet. These three groups, We used Isley’s (1944) description of mandible still used today, were graminivorous (grass-feed- types and their adaptive functions to divide the ing type) with grinding molars and incisors typi- mandibles into 3 major categories: forbivorous cally fused into a scythe-like cutting edge, for- (forb-feeding), graminivorous (grass-feeding), bivorous (forb or broadleaf plant-feeding type) and herbivorous (mixed-feeding). which have a molar region consisting of a depres- Mandibles were lightly brushed with 80 per- sion surrounded by raised teeth and sharp inter- cent ETOH and distilled water in an effort to re- locking incisor teeth, and herbivorous (mixed- move most of the sand and debris adhering to the feeding type) that have characteristics of both of mouthparts. Photographs were taken with the the aforementioned groups. The original findings Syncroscopy Auto-Montage system (University of by Isley (1944) have since been proven to be wide- Florida, Entomology and Nematology Dept.). spread in grasshoppers. Additional studies have The mandible structure of 36 species of grass- been conducted by Snodgrass (1928), Gangwere hopper, from five subfamilies (Acridinae, Cyrta- (1965, 1966), Gangwere et al. (1976), and Patter- canthacridinae, Gomphocerinae, Oedipodinae, son (1984) in North America; Lieberman (1968) and Romaleinae), found in Florida was micro- and Gangwere & Ronderos (1975) in South Amer- scopically examined. These grasshoppers were ica; Williams (1954), Kaufmann (1965), and collected from a variety of habitats including dis- Gangwere & Morales (1973) in Europe; Gangwere turbed freshwater marsh, high pine, swamp, and & Spiller (1995) and Gangwere et al. (1998) in the oak hammocks. All grasshoppers had distinctive Mediterranean islands; Feroz & Chaudhry mouthparts that could be described as forbivo- (1975), Gapud (1968), and Kang et al. (1999) in rous (forb-feeding type), herbivorous (mixed-feed- Asia; and Chapman (1964) in Africa. ing type), or graminivorous (grass-feeding type) The relationship between grasshopper mouth- (Fig. 1, A-L). A list of each species studied and the parts and food is far from precise. Mulkern (1967) mandible type is given in Table 1. was convinced that only the grossest determina- Of the subfamilies examined, the Cyrtacan- tions could be made between mandibular struc- thacridinae demonstrated the most diversity in ture and diet (i.e., graminivorous, forbivorous, and mandible type; however, most of them displayed herbivorous). Occasionally, grasshoppers with either herbivorous or forbivorous mandibles, indi- forb-feeding mandibles regularly feed on grasses cating a tendency toward forb-feeding. These or vice versa (Chapman 1964). Nevertheless, there grasshoppers can be found in a wide range of hab- is some value in assessing mouthpart structure itats, usually in dense vegetation or woodland ar- relative to predicting diet and habitat of grasshop- eas, and are quite active in both walking and fly- pers, especially for the many rare or non-economic ing. It is interesting to note that both the grass- species that are unlikely to be studied in detail. hoppers in this subfamily that did display Thus, the morphological characteristics and struc- graminivorous type mandibles (L. marginicollis tural adaptations of the mouthparts of 36 of the 71 and S. vitreipennis) also have extremely slender, grasshoppers occurring in Florida were examined. elongated bodies and can be found on the edges of Grasshoppers were collected from various habi- ponds or in freshwater marshes (Isley 1944; Squi- tats throughout north-central Florida in 2001 and tier & Capinera 2002b; Smith & Capinera 2005). 2002. Thirty-six of the most common Floridian These grasshoppers typically grasp the stems of grasshopper species were identified with the tax- emergent grass or grass-like vegetation such as onomic key found in Smith et al. (2004) and fro- sedges or cattails, blending in almost perfectly. Scientific Notes 205 Fig. 1. Mandibles of Amblytropidia mysteca, a representative graminivorous species: right incisor region (A), left incisor region (B), right molar region (C), and left molar region (D); Schistocerca ceratiola, a representative forbiv- orous species: right incisor region (E), left incisor region (F), right molar region (G), and left molar region (H); and Spharagemon cristatum, a representative herbivorous species: right incisor region (I), left incisor region (J), right molar region (K), and left molar region (L). 206 Florida Entomologist 88(2) June 2005 TABLE 1. ACRIDID SPECIES EXAMINED AND THEIR MOUTHPART MORPHOLOGY. Graminivorous type mandibles Herbivorous type mandibles Forbivorous type mandibles Acridinae Cyrtacanthacridinae Cyrtacanthacridinae Metaleptea brevicornis (Johannson) Gymnoscirtetes pusillus Scudder Aptenopedes aptera Scudder Melanoplus bispinosus Scudder Aptenopedes sphenarioides Scudder Cyrtacanthacridinae Melanoplus sanguinipes (Fabricius) Melanoplus keeleri (Thomas) Leptysma marginicollis (Serville) Schistocerca alutacea (Harris) Melanoplus ordwayae Deyrup Stenacris vitreipennis (Marschall) Schistocerca americana (Drury) Melanoplus propinquus Scudder Schistocerca obscura (Fabricius) Melanoplus punctulatus Scudder Gomphocerinae Schistocerca rubiginosa (Scudder) Melanoplus querneus Rehn & Hebard Achurum carinatum (F. Walker) Melanoplus rotundipennis Scudder Amblytropidia mysteca (Saussure) Oedipodinae Paroxya atlantica Scudder Dichromorpha viridis (Scudder) Chortophaga australior (Rehn & Hebard) Paroxya clavuliger (Serville) Eritettix obscurus (Scudder) Spharagemon crepitans (Saussure) Schistocerca ceratiola Hubbell & Walker Mermiria intertexta Scudder Spharagemon cristatum (Scudder) Schistocerca damnifica (Saussure) Orphulella pelidna (Burmeister) Syrbula admirabilis (Uhler) Romaleinae Romalea microptera (Beauvois) Oedipodinae Arphia granulata (Saussure) Hippiscus ocelote (Saussure) Pardalophora phoenicoptera (Burmeister) However, the overwhelming majority of these these subfamilies was limited. The Acridinae are grasshoppers display either herbivorous or for- typically considered to be grass-feeders, display- bivorous mandibles (Isley 1944; Gangwere 1965, ing the classic graminivorous type mandibles 1966; Squitier & Capinera 2002a). (Chapman 1964; Isley 1944). Very rarely a species The Oedipodinae were split between two man- in this subfamily will display herbivorous type dible types: graminivorous and herbivorous. This mandibles (Chapman 1964). The Romaleinae are signifies a more grass-dominated diet. However, always forb feeders and always display forbivo- these grasshoppers are much more divergent and rous type mandibles (Isley 1944; Squitier & Cap- some may be completely graminivorous or forbivo- inera 2002a; Smith & Capinera 2005). rous. Most of the species in this subfamily were Many thanks to David Almquist for help in found on the ground in open areas on bare soil, taking photographs with the auto-montage sys- rarely on plants or grasses. As a general rule, the tem. This research was supported by the Florida Oedipodinae show the most mandible diversity of Agricultural Experiment Station, and approved all the grasshopper subfamilies. Isley (1944), for publication as Journal Series No. R-10456. Gangwere (1966), and Kang et al. (1999) found a fairly even distribution of the three mouthpart SUMMARY types in this group. The gomphocerinae all had graminivorous Mouthpart consistency within subfamilies in- mandibles, indicating a consistent diet of grasses. dicates that evolution is just as important as eco- These findings are reinforced by the preferred logical factors in determining food plants; for habitats of this subfamily, usually open grassy most subfamilies there is a strong association fields and pastures. Virtually all Gomphocerinae with a particular form of vegetation. It is evident are graminivorous (Lockwood et al. 1994), or at that the ability, or tendency, of grasshoppers to least have graminivorous type mandibles. Occa- change hosts is partly limited by the structure of sionally a gomphocerine will display graminivo- their mandibles. However, because there are ex- rous type
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