Excerpt: Many studies have been undertaken in the past on the food selection, food preferences, and economic damage of various grasshoppers and their allies. Among the more salient of these researches are those of Anderson (1961, 1964); Ball (1 936); Bindra (1958); Boldyrev (1928); Blackith and Blackith (1966); Brues (1946); Chapman (1957); Dibble (1940); Gangwere (1959, 1960, 1961, 1965, 1965a, 1966, 1966a, 1967);Husain etal. (1946); Isely (1938, 1946); Isely and Alexander (1949); Joyce (1952); Mulkern and Anderson (1959); Mulkern, Anderson, and Brusven (1962); Mulkern et al. (1969); Pfadt (1949); Riley (1878); Roonwal (1953); Savin (1927); Weiss (1924); and Williams (1954). Techniques useful in the investigation of food selection are to be found in certain of the above reports. Especially noteworthy in this respect are those by Blackith and Blackith, Chapman, Gangwere (1961), Isely and Alexander, Joyce, Mulkern and Anderson, Pfadt, Roonwal, and Savin.
The relatively new technique of Blackith and Blackith (1966) involves the comparison of colorations in the ileal diverticula of morabine grasshoppers (Orthoptera: Eumastacidae). The digestive caeca of eumastacids were previously discussed by Slifer (1944), and those of other groups by Gangwere (1966) in a comprehensive paper dealing with the mechanical handling of food in the orthopteroid alimentary canal. There is also an extensive literature on the gut physiology of these insects, some of which is appropriate to a consideration of the caeca and diverticula

Tyrkus, Michael

English

Valparaiso University

The Great Lakes Entomologist Volume 4 Number 1 -- Spring 1971 Number 1 -- Spring 1971 Article 4 July 2017 The Feasibility of Use of Caecal and Diverticular Coloration in Field Determination of Grasshopper Diet Michael Tyrkus Wayne State University Follow this and additional works at: https://scholar.valpo.edu/tgle  Part of the Entomology Commons Recommended Citation Tyrkus, Michael 2017. "The Feasibility of Use of Caecal and Diverticular Coloration in Field Determination of Grasshopper Diet," The Great Lakes Entomologist, vol 4 (1) Available at: https://scholar.valpo.edu/tgle/vol4/iss1/4 This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. THE MICHIGAN ENTOMOLOGIST Vol. 4, No. 1 THE FEASIBILITY OF USE OF CAECAL AND DIVERTICULAR COLORATION IN FIELD DETERMINATION OF GRASSHOPPER D IET ' , '  Michael Tyrkus Department of Biology, Wayne State University, Detroit, Michigan 48202 INTRODUCTION Many studies have been undertaken in the past on the food selection, food preferences, and economic damage of various grasshoppers and their allies. Among the more salient of these researches are those of Anderson (1961, 1964); Ball (1 936); Bindra (1958); Boldyrev (1928); Blackith and Blackith (1966); Brues (1946); Chapman (1957); Dibble (1940); Gangwere (1959, 1960, 1961, 1965, 1965a, 1966, 1966a, 1967);Husain etal .  (1946); Isely (1938, 1946); Isely and Alexander (1949); Joyce (1952); Mulkern and Anderson (1959); Mulkern, Anderson, and Brusven (1962); Mulkern e t  al. (1969); Pfadt (1949); Riley (1878); Roonwal (1953); Savin (1927); Weiss (1924); and Williams (1954). Techniques useful in the investigation of food selection are to  be found in certain of the above reports. Especially noteworthy in this respect are those by Blackith and Blackith, Chapman, Gangwere (1961), Isely and Alexander, Joyce, Mulkern and Anderson, Pfadt, Roonwal, and Savin. The relatively new technique of Blackith and Blackith (1966) involves the comparison of colorations in the ileal diverticula of morabine grasshoppers (Orthoptera: Eumastacidae). The digestive caeca of eumastacids were previously discussed by Slifer (1944), and those of other groups by Gangwere (1966) in a comprehensive paper dealing with the mechanical handling of food in the orthopteroid alimentary canal. There is also an extensive literature on the gut physiology of these insects, some of which is appropriate to a consideration of the caeca and diverticula. The possibility that the method of Blackith and Blackith is applicable to other groups of Orthopteroidea prompted the present research, which is, in part, an attempt to test the feasibility of using gastric caecal and diverticular coloration in studying the food-habits of certain members of the Michigan fauna. Moreover, assuming that the method is applicable, and is utilized in an area of known floral composition, it should be indicative of the effect that availability of food plants has on the diets of the various species studied; that possibility is also investigated. MATERIALS AND METHODS During the field season of 1968, various species of Orthopteroidea (largely Acrididae) were collected at Proud Lake and Pontiac Lake Recreation Areas, Oakland County, and at Stoney Creek Metropolitan Park, Macomb County, Michigan. Laboratory populations of the captured insects were maintained for later study using the grasshopper rearing cages described by Gangwere (1960), slightly modified by the use of pie tins both as cage top and as bottom, and were given appropriate maintenance food plants in Erlenmeyer flasks. During actual experimentation, individual grasshoppers were isolated in cages such as were described by Gangwere (1960) for rearing mantids. The insects were starved for 24 to 48 hours (emptying the gut), but during this period were given access to drinking water supplied in cotton-stoppered vials. Following starvation, a known species of food plant was introduced into each cage, and the animals were allowed to feed for 2 to 24 hours. The range of allowed feeding time was determined by the hours necessary to produce particular colorations and to study the mode of color change. The grasshoppers were then killed with ethyl acetate. Under a binocular dissecting microscope, the terminal portion of the abdomen was cut off, the animals were decapitated, and the freed gut was pulled through the foramen magnum to  be placed on a clean, dry microscope slide. As the dissection was immediate, it l ~ o n t r i b u t i o n  No. 247 from the Department of Biology, Wayne State University, Detroit, Michigan 48202. 2 ~ a s e d  on a thesis submitted in partial fulfillment of the degree of Master of Science, Wayne State University. 1Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in FPublished by ValpoScholar, 2017THE MICHIGAN ENTOMOLOGIST Fig. 1.  Alimentary canal of the grasshopper, Syrbula admirabilis (Uhler), showing four of the six caeca and their caudal diverticula. ca, caecum; cr, crop; di, diverticulum; mt, anterior intestine; mt, malpighian tubule; a ,  oesophagus; r, rectum; rp, rectal "pad"; vt, ventriculus ("stomach"). Fig. 2. Alimentary canal of Syrbula adrnirabilis showing the first sequence of color change in the caeca and diverticula. Sequence denoted respectively by a, b, c, and d. Fig. 3. Alimentary canal of Syrbula admirabilis showing the second sequence of color change in the caeca and diverticula. Sequence denoted respectively by a, b, and c. Fig. 4. Alimentary canal of Syrbula admirabilis showing the third sequence of color change in the caeca and diverticula. Sequence denoted respectively by a and b. 2The Great Lakes Entomologist, Vol. 4, No. 1 [2017], Art. 4https://scholar.valpo.edu/tgle/vol4/iss1/416 THE MICHIGAN ENTOMOLOGIST Vol. 4, No. I was not necessary to use an isotonic solution to  maintain the gut tissues. This avoidance of the use of saline solution was actually desirable so as not to  mix the gut contents. Coloration of the caeca and diverticula was recorded immediately, for the observed colors proved highly ephemeral. The following species are investigated: Acrididae: Gomphocerine (Slant-Facted Grasshoppers) Chorthippus c. curtipennis (Hams) Acridadae: Oedipodinae (Band-Winged Grasshoppers) Arphia p. pseudonietana (Thomas) Dissosteira carolina (Linnaeus) Encoptolophus s. sordidus (Burmeister) Pardalophora apiculata (Harris) Spharagemon b. bolli (Scudder) Acrididae: Catantopinae (Spine-Breasted Grasshoppers) Melanoplus confusus (Scudder) Melanoplus f -r femur-rubrum (De Geer) Melarzoplus s. sanguinipes (Fabricius) Tettigoniidae: Phaneropterinae (Bush and Round-Headed Katydids) Scudderia c. curvica~tda (De Geer) RESULTS The results of the study are presented in Table 1 (Caecal and Diverticular Coloration of Selected Orthoptera). DISCUSSION The orthopteroid insects possess a number of blind pouches at the proximal end of the ventriculus (Fig. 1). They include the caeca (those pouches that project cephalad) and the diverticula (those that project caudad). Distinct colors and color patterns were observable in the caeca and diverticula of the species fed and studied in the present investigation (Table 1). These patterns proved to be superficially visible when the gut was first removed from the body, and were even brighter and more intense on its interior walls (as was indicated by observations on the slit gut). Moreover, within the exposed lumen there was, at times, a small volume of fluid with a slight hue generally indicative of the wall coloration. These several colors and color patterns were highly ephemeral, which necessitated immediate observation before they faded and were replaced by the dark hues of the death condition. The guts of starved animals were always dark brown to black in color. This "starved" coloration changed promptly with feeding. Sequentially timed dissections after feeding demonstrated disappearance of the "starved" coloration in a rather prescribed manner: the "starved" coloration first disappeared from the proximal end of the caeca. This new coloration (in part attributable to  the food source) then spread from the proximal to  the distal end of the caeca, while simultaneously it replaced the "starved" condition in the proximal portion of the diverticula. The distal portion of the diverticula was generally the last area in which the coloration underwent food-induced change (Fig. 2). Two relatively common variations to the above general pattern of color change were observed. In the first, coloration began simultaneously both in the proximal caeca and in the proximal diverticula, and then spread through the remaining area of the caeca and then of the diverticula (Fig. 3). In the second variation, there was a simultaneous, relatively synchronized spreading of color throughout the length both of the caeca and of the diverticula (Fig. 4), but at times lines or stripes of the "starved" condition coloration persisted, usually slightly lighter than their initial shade. Large variations in coloration were seen, both within given species arid between them. However, in all cases, the intensity and distinctiveness of the observed coloration appeared related to  the amount of food present in the midgut. Thus, when the gut contents were concentrated in the hindgut, the caecal and diverticular coloration was less well defined. 3Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in FPublished by ValpoScholar, 20171971 THE MICHIGAN ENTOMOLOGIST 17 The hoped for (but not expected) specificity of color based on food source was not encountered. There proved to be a wide range of color variation, but it was not sufficiently consistent to  submit to classification. Individuals of a given grasshopper species were sometimes observed to develop different coloration when eating different parts-and sometimes even identical parts-of the same plant species. It is apparent that gut color variation is due, in part, to food source, but is also greatly modified by the animal's internal physiological state, which rules out use of the technique for field determination of Michigan grasshopper food-habits. The possibility that caecal coloration is a function of genetic factors inherent within a single grasshopper population is also dismissed, based on the complete absence of any noticeable difference in the animals collected from different stations. It seems more likely that the observed patterns of coloration are the result of the extraction of plant pigments from the food, as modified and influenced by digestive enzymes and other substances present in the feeder's alimentary tract. CONCLUSIONS In light of the results of the present study, it appears that the gut coloration in non-morabine grasshoppers-at least in the species here investigated-is not sufficiently specific to submit to  classification, and cannot be used as an index of food-habit. The observed colors and color patterns prove ephemeral, for they change with the onset of feeding, and are continuously altered as the food progresses along the length of the alimentary tract. These colors doubtlessly derive from substances within the food source, as modified by the internal state of the feeder. ACKNOWLEDGEMENTS The author is greatly indebted to  Professor S. K. Gangwere, Department of Biology, Wayne State University, who served as major advisor of the Master's Committee to  which the foregoing report was originally submitted; Miss Rosemarie Rozsa, who drew figures 1-4; Miss Marlene Nigosian, Department of Gynecology and Obstetrics, Wayne State University School of Medicine, and Miss Mary Talos, Department of Biology, Wayne State University, who typed the various revisions of the manuscript; and finally to  his wife, Marian, who served as secretary and laboratory assistant throughout the entire project. LITERATURE CITED Anderson, N. L. 196 1. Seasonal losses in rangeland vegetation due to grasshoppers. J. Econ. Entomol. 54:369-378. Anderson, N. L. 1964. Some relationships between grasshoppers and vegetation. Ann. Entomol. Soc. Amer. 57:736-742. Ball, E. D. 1936. Food plants of some Arizona grasshoppers. J. Econ. Entomol. 29:679-684. Bindra, 0 .  S. 1958. Food preferences of Poecilocerus pictus Fabr., a pest of some horticultural plants at Gwalior. Ind. J. Hort. Sobour. 15:80-86. Blackith, R. E., and R. M. Blackith. 1966. The food of morabine grasshoppers. Aust. J. Zool. 14: 877-894. Boldyrev, B. T. 1928. Biological studies on Bradyporus rnultituberculatus F. W. Eos. 4: 13-56. Brues, C. F. 1946. Insect dietary. Harvard Univ. Press, Cambridge Mass. Chapman, R. F. 1957. Observaticns on the feeding of adults of the red locust (Nomadacris septemfasciata [Serville]). Brit. J .  Animal Behav. 5:60-75. Dibble, C. B. 1940. Grasshoppers, a factor in soil erosion in Michigan. J. Econ. Entomol. 33:498-499. 4The Great Lakes Entomologist, Vol. 4, No. 1 [2017], Art. 4https://scholar.valpo.edu/tgle/vol4/iss1/418 THE MICHIGAN ENTOMOLOGIST Vol. 4, No. 1 Gangwere, S. K. 1959. Experiments upon the food consumption of the grasshopper Melanoplus s. scudderi (Uhler), Papers Mich. Acad. Sci. Arts Lett. 44:93-96. Gangwere, S. K. 1960. The feeding and culturing of Orthoptera in ine laboratory. Entomol. News 71 :7-13, 37-44. Gangwere, S. K. 1961. A monograph on food selection in Orthoptera. Trans. Entomol. Soc. Amer. 87:67-230. Gangwere, S. K. 1965. Food selection in the oedipodine grasshopper Arphia sulphurea (Fabricius). Amer. Midl. Nat. 74:67-75. Gangwere, S. K. 1965a. The strucrural adaptations of mouthparts in Orthoptera and allies. EOS 40167-85. Gangwere, S. K. 1966. The mechanical handling of food by the alimentary canal of Orthoptera and allies. Eos 41 :247-265. Gangwere, S. K. 1966a. Relationships between the mandibles, feeding behavior. and damage inflicted on plants by the feeding of certain acridids (Orthoptera). Mich. Entomol. 1:13-16. Gangwere, S. K. 1967. The feeding behavior of Atlanticus testaceus (Orthoptera: Tettigoniidae). Ann. Entomol. Soc. Amer. 60:74-81. Husain, M. A,, C. B. Mathur, and M. L. Roonwal. 1946. Studies on Schistocerca gregaria (Forskal) XIII. Food and feeding habits of the desert locust. Ind. J. Entomol. 8: 141-163. Isely, F. B. 1938. The relations of Texas Acrididae to plants and soils. Ecol. Mon. 8:551-604. Isely, F. B. 1946. Differential feeding in relation to local distribution of grasshoppers. Ecol. 27: 128-138. Isely, F. B., and G. Alexander. 1949. Analysis of insect food habits by crop examination. Sci. 109:115-116. Joyce, R. J. V. 1952. The ecology of grasshoppers in East Central Sudan. Anti-Locust Bull., Anti-Locust Res. Centre 1 1 : 1-99. Mulkern, G. B., and J. F. Anderson. 1959. A technique for studying the food habits and preferences of grasshoppers. J. Econ. Entornol. 52:342. Mulkern, G. B., J. F. Anderson, and M. A. Brusven. 1962. Biology and ecology of North Dakota grasshoppers. 1. Food habits and preferences of grasshoppers associated with alfalfa fields. Res. Rep. N. Dak. Agric. Exp. Stat. 7. Mulkern, G. B., K. P. Preuss, H. Knutson, A. F. Hagen, J. B. Campbell, and J. D. Larnbley. 1969. Food habits and preferences of grassland grasshoppers of the north central great plains. Bull. N. Dak. Agric. Exp. Stat. 481: 1-32. Pfadt, R. E. 1949. Food plants as factors in the ecology of the lesser migratory grasshopper, Melanoplus mexicanus (Saussure). Bull. Wyom. Agric. Exp. Stat. 290. Riley, C. V. 1878. The rocky mountain locust. Its habits and natural history. First Rept. U. S. Entomol. Comm. (1877):212-257. Roonwal. M. L. 1953. Food- reference ex~eriments on the desert locust. Schistocerca gregaria (Forskal), in its permanent breeding grounds in Mekran. J. Zool. Soc. Ind. 5:44-58. Savin, M. B. 1927. Food preferences of the black cricket (Gryllus assirnilis) with speciaI reference to the damage done to fabrics. Entomol. News 38:4-10. Slifer, E. H. 1944. Ileal caecae in the Eumastacidae (Orthoptera). Ann. Entomol. Soc. Amer. 37:441-446. Weiss, H. B. 1924. Insect food habits and vegetation. Ohio J. Sci. 24: 100-106. Williams, L. H. 1954. The feeding habits and food preferences of Acrididae and the factors which determine them. Trans. Roy. Entomol. Soc. London 105:423-454. 5Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in FPublished by ValpoScholar, 2017Table 1. Caecal and diverticular coloration of selected Orthoptera - w 2 Foods Plant parts Feeders Caecal Diverticular ingested3 coloration coloration Gramineae Agrostis sp. ,> >, I ,  7 ,  Eleusine indica ,, , 7, 3 ,  7 , )  ,, 7 Grass sp. ,, ,, Pardalophora apiculata 7, ,, Melanoplus s. sanguinipes 9 ,  39 7, Dissosteira carolina Melanoplus f:-r. femur- rubrum Encoptolophus s. sordidus Golden; brown striped Golden; brown striped Golden; brown striped Yellow-green Brown Olive Yellow-green Yellow-brown Golden; brown striped Yellow-white Yellow-white with reddish brown tips Yellow-brown striped Yellow Yellow-brown striped Amber Dark blue (distally); creamy brown (prox- imally) White Dull pink Pale golden Golden Golden Golden Y ellow-green Brown Olive Yellow-green Black Golden-brown striped Dark green Black Yellow-brown striped Yellow Yellow-brown striped Amber Creamy brown White Dull pink Golden brown & brown striped 6The Great Lakes Entomologist, Vol. 4, No. 1 [2017], Art. 4https://scholar.valpo.edu/tgle/vol4/iss1/4THE MICHIGAN ENTOMOLOGIST Vol. 4. No. 1 7Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in FPublished by ValpoScholar, 2017Table 1 (Continued) Cruciferae Lepidium virginicum Golden; brown striped Golden; brown striped Golden Melanoplus confusus Melanoplus s. sanguinlpes Golden; brown striped Fabaceae L'espedeza sp. , , Melanoplus s. sanguinipes 3 7 ,? Milky white White Milky white White Lythraceae Lythrum salicaria Umbelliferae Daucus carota 6 9 Scudderia c. curvicauda Purple Purple 2 m Yellow Z Pale green z Transparent z 0 * Dark brown z m (starved animal) t j  Black (starved animal) 2 Black (starved animal) g E r n  Spharagemon b. bolli Melanoplus confusus Yellow Pale green Golden Asclepiadaceae Asclepias tuberosa Melanoplus s. sanguinipes Dark brown (starved animal) Black (starved animal) Black (starved animal) Melanoplus confusus 9 ,  Labiatae Monarda fistulosa 3 3,  4 Y ellow-white Yellow t o  gold White Melanoplus s. sanguinipes 9, 9 ,  7, Golden green White Composit ae Galinosoga parviflora Lactuca sp. Melanoplus s. sanguinipes Pardalophora apiculata Yellow-brown striped Pale t o  bright yellow (distally t o  prox- imally) Whitish yellow Green Golden Yellow Cream Whitish yellow Pale black - 1 3  Golden Melanoplus s. sanguinipes Melanoplus confusus 9, 99 8The Great Lakes Entomologist, Vol. 4, No. 1 [2017], Art. 4https://scholar.valpo.edu/tgle/vol4/iss1/4THE MICHIGAN ENTOMOLOGIST 9Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in FPublished by ValpoScholar, 2017

The Feasibility of Use of Caecal and Diverticular Coloration in Field Determination of Grasshopper Diet

ValpoScholar

The Great Lakes Entomologist 
Volume 4 
Number 1 -- Spring 1971 Number 1 -- Spring 
1971 
Article 4 
July 2017 
The Feasibility of Use of Caecal and Diverticular Coloration in 
Field Determination of Grasshopper Diet 
Michael Tyrkus 
Wayne State University 
Follow this and additional works at: https://scholar.valpo.edu/tgle 
 Part of the Entomology Commons 
Recommended Citation 
Tyrkus, Michael 2017. "The Feasibility of Use of Caecal and Diverticular Coloration in Field Determination 
of Grasshopper Diet," The Great Lakes Entomologist, vol 4 (1) 
Available at: https://scholar.valpo.edu/tgle/vol4/iss1/4 
This Peer-Review Article is brought to you for free and open access by the Department of Biology at ValpoScholar. 
It has been accepted for inclusion in The Great Lakes Entomologist by an authorized administrator of ValpoScholar. 
For more information, please contact a ValpoScholar staff member at scholar@valpo.edu. 
THE MICHIGAN ENTOMOLOGIST Vol. 4, No. 1 
THE FEASIBILITY OF USE OF CAECAL AND DIVERTICULAR 
COLORATION IN FIELD DETERMINATION OF GRASSHOPPER D IET ' , '  
Michael Tyrkus 
Department of Biology, Wayne State University, Detroit, Michigan 48202 
INTRODUCTION 
Many studies have been undertaken in the past on the food selection, food preferences, 
and economic damage of various grasshoppers and their allies. Among the more salient of 
these researches are those of Anderson (1961, 1964); Ball (1 936); Bindra (1958); Boldyrev 
(1928); Blackith and Blackith (1966); Brues (1946); Chapman (1957); Dibble (1940); 
Gangwere (1959, 1960, 1961, 1965, 1965a, 1966, 1966a, 1967);Husain etal .  (1946); Isely 
(1938, 1946); Isely and Alexander (1949); Joyce (1952); Mulkern and Anderson (1959); 
Mulkern, Anderson, and Brusven (1962); Mulkern e t  al. (1969); Pfadt (1949); Riley (1878); 
Roonwal (1953); Savin (1927); Weiss (1924); and Williams (1954). Techniques useful in the 
investigation of food selection are to  be found in certain of the above reports. Especially 
noteworthy in this respect are those by Blackith and Blackith, Chapman, Gangwere (1961), 
Isely and Alexander, Joyce, Mulkern and Anderson, Pfadt, Roonwal, and Savin. 
The relatively new technique of Blackith and Blackith (1966) involves the comparison of 
colorations in the ileal diverticula of morabine grasshoppers (Orthoptera: Eumastacidae). 
The digestive caeca of eumastacids were previously discussed by Slifer (1944), and those of 
other groups by Gangwere (1966) in a comprehensive paper dealing with the mechanical 
handling of food in the orthopteroid alimentary canal. There is also an extensive literature 
on the gut physiology of these insects, some of which is appropriate to a consideration of 
the caeca and diverticula. 
The possibility that the method of Blackith and Blackith is applicable to other groups of 
Orthopteroidea prompted the present research, which is, in part, an attempt to test the 
feasibility of using gastric caecal and diverticular coloration in studying the food-habits of 
certain members of the Michigan fauna. Moreover, assuming that the method is applicable, 
and is utilized in an area of known floral composition, it should be indicative of the effect 
that availability of food plants has on the diets of the various species studied; that 
possibility is also investigated. 
MATERIALS AND METHODS 
During the field season of 1968, various species of Orthopteroidea (largely Acrididae) 
were collected at Proud Lake and Pontiac Lake Recreation Areas, Oakland County, and at 
Stoney Creek Metropolitan Park, Macomb County, Michigan. Laboratory populations of the 
captured insects were maintained for later study using the grasshopper rearing cages 
described by Gangwere (1960), slightly modified by the use of pie tins both as cage top and 
as bottom, and were given appropriate maintenance food plants in Erlenmeyer flasks. 
During actual experimentation, individual grasshoppers were isolated in cages such as 
were described by Gangwere (1960) for rearing mantids. The insects were starved for 24 to 
48 hours (emptying the gut), but during this period were given access to drinking water 
supplied in cotton-stoppered vials. Following starvation, a known species of food plant was 
introduced into each cage, and the animals were allowed to feed for 2 to 24 hours. The 
range of allowed feeding time was determined by the hours necessary to produce particular 
colorations and to study the mode of color change. The grasshoppers were then killed with 
ethyl acetate. Under a binocular dissecting microscope, the terminal portion of the abdomen 
was cut off, the animals were decapitated, and the freed gut was pulled through the foramen 
magnum to  be placed on a clean, dry microscope slide. As the dissection was immediate, it 
l ~ o n t r i b u t i o n  No. 247 from the Department of Biology, Wayne State University, Detroit, Michigan 
48202. 
2 ~ a s e d  on a thesis submitted in partial fulfillment of the degree of Master of Science, Wayne State 
University. 
1
Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in F
Published by ValpoScholar, 2017
THE MICHIGAN ENTOMOLOGIST 
Fig. 1.  Alimentary canal of the grasshopper, Syrbula admirabilis (Uhler), showing four of the six caeca and 
their caudal diverticula. ca, caecum; cr, crop; di, diverticulum; mt, anterior intestine; mt, malpighian 
tubule; a ,  oesophagus; r, rectum; rp, rectal "pad"; vt, ventriculus ("stomach"). 
Fig. 2. Alimentary canal of Syrbula adrnirabilis showing the first sequence of color change in the caeca and 
diverticula. Sequence denoted respectively by a, b, c, and d. 
Fig. 3. Alimentary canal of Syrbula admirabilis showing the second sequence of color change in the caeca 
and diverticula. Sequence denoted respectively by a, b, and c. 
Fig. 4. Alimentary canal of Syrbula admirabilis showing the third sequence of color change in the caeca and 
diverticula. Sequence denoted respectively by a and b. 
2
The Great Lakes Entomologist, Vol. 4, No. 1 [2017], Art. 4
https://scholar.valpo.edu/tgle/vol4/iss1/4
16 THE MICHIGAN ENTOMOLOGIST Vol. 4, No. I 
was not necessary to use an isotonic solution to  maintain the gut tissues. This avoidance of 
the use of saline solution was actually desirable so as not to  mix the gut contents. 
Coloration of the caeca and diverticula was recorded immediately, for the observed colors 
proved highly ephemeral. 
The following species are investigated: 
Acrididae: Gomphocerine (Slant-Facted Grasshoppers) 
Chorthippus c. curtipennis (Hams) 
Acridadae: Oedipodinae (Band-Winged Grasshoppers) 
Arphia p. pseudonietana (Thomas) 
Dissosteira carolina (Linnaeus) 
Encoptolophus s. sordidus (Burmeister) 
Pardalophora apiculata (Harris) 
Spharagemon b. bolli (Scudder) 
Acrididae: Catantopinae (Spine-Breasted Grasshoppers) 
Melanoplus confusus (Scudder) 
Melanoplus f -r femur-rubrum (De Geer) 
Melarzoplus s. sanguinipes (Fabricius) 
Tettigoniidae: Phaneropterinae (Bush and Round-Headed Katydids) 
Scudderia c. curvica~tda (De Geer) 
RESULTS 
The results of the study are presented in Table 1 (Caecal and Diverticular Coloration of 
Selected Orthoptera). 
DISCUSSION 
The orthopteroid insects possess a number of blind pouches at the proximal end of the 
ventriculus (Fig. 1). They include the caeca (those pouches that project cephalad) and the 
diverticula (those that project caudad). Distinct colors and color patterns were observable in 
the caeca and diverticula of the species fed and studied in the present investigation (Table 
1). These patterns proved to be superficially visible when the gut was first removed from the 
body, and were even brighter and more intense on its interior walls (as was indicated by 
observations on the slit gut). Moreover, within the exposed lumen there was, at times, a 
small volume of fluid with a slight hue generally indicative of the wall coloration. These 
several colors and color patterns were highly ephemeral, which necessitated immediate 
observation before they faded and were replaced by the dark hues of the death condition. 
The guts of starved animals were always dark brown to black in color. This "starved" 
coloration changed promptly with feeding. Sequentially timed dissections after feeding 
demonstrated disappearance of the "starved" coloration in a rather prescribed manner: the 
"starved" coloration first disappeared from the proximal end of the caeca. This new 
coloration (in part attributable to  the food source) then spread from the proximal to  the 
distal end of the caeca, while simultaneously it replaced the "starved" condition in the 
proximal portion of the diverticula. The distal portion of the diverticula was generally the 
last area in which the coloration underwent food-induced change (Fig. 2). 
Two relatively common variations to the above general pattern of color change were 
observed. In the first, coloration began simultaneously both in the proximal caeca and in the 
proximal diverticula, and then spread through the remaining area of the caeca and then of 
the diverticula (Fig. 3). In the second variation, there was a simultaneous, relatively 
synchronized spreading of color throughout the length both of the caeca and of the 
diverticula (Fig. 4), but at times lines or stripes of the "starved" condition coloration 
persisted, usually slightly lighter than their initial shade. 
Large variations in coloration were seen, both within given species arid between them. 
However, in all cases, the intensity and distinctiveness of the observed coloration appeared 
related to  the amount of food present in the midgut. Thus, when the gut contents were 
concentrated in the hindgut, the caecal and diverticular coloration was less well defined. 
3
Tyrkus: The Feasibility of Use of Caecal and Diverticular Coloration in F
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1971 THE MICHIGAN ENTOMOLOGIST 17 
The hoped for (but not expected) specificity of color based on food source was not 
encountered. There proved to be a wide range of color variation, but it was not sufficiently 
consistent to  submit to classification. Individuals of a given grasshopper species were 
sometimes observed to develop different coloration when eating different parts-and 
sometimes even identical parts-of the same plant species. It is apparent that gut color 
variation is due, in part, to food source, but is also greatly modified by the animal's internal 
physiological state, which rules out use of the technique for field determination of Michigan 
grasshopper food-habits. 
The possibility that caecal coloration is a function of genetic factors inherent within a 
single grasshopper population is also dismissed, based on the complete absence of any 
noticeable difference in the animals collected from different stations. It seems more likely 
that the observed patterns of coloration are the result of the extraction of plant pigments 
from the food, as modified and influenced by digestive enzymes and other substances 
present in the feeder's alimentary tract. 
CONCLUSIONS 
In light of the results of the present study, it appears that the gut coloration in 
non-morabine grasshoppers-at least in the species here investigated-is not sufficiently 
specific to submit to  classification, and cannot be used as an index of food-habit. The 
observed colors and color patterns prove ephemeral, for they change with the onset of 
feeding, and are continuously altered as the food progresses along the length of the 
alimentary tract. These colors doubtlessly derive from substances within the food source, as 
modified by the internal state of the feeder. 
ACKNOWLEDGEMENTS 
The author is greatly indebted to  Professor S. K. Gangwere, Department of Biology, 
Wayne State University, who served as major advisor of the Master's Committee to  which 
the foregoing report was originally submitted; Miss Rosemarie Rozsa, who drew figures 1-4; 
Miss Marlene Nigosian, Department of Gynecology and Obstetrics, Wayne State University 
School of Medicine, and Miss Mary Talos, Department of Biology, Wayne State University, 
who typed the various revisions of the manuscript; and finally to  his wife, Marian, who 
served as secretary and laboratory assistant throughout the entire project. 
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18 THE MICHIGAN ENTOMOLOGIST Vol. 4, No. 1 
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Table 1. Caecal and diverticular coloration of selected Orthoptera 
- w 
2 
Foods Plant parts Feeders Caecal Diverticular 
ingested3 coloration coloration 
Gramineae 
Agrostis sp. 
,> >, 
I ,  7 ,  
Eleusine indica 
,, 
, 7, 
3 ,  7 , )  
,, 
7 
Grass sp. 
,, ,, 
Pardalophora apiculata 
7, 
,, 
Melanoplus s. sanguinipes 
9 ,  39 7, 
Dissosteira carolina 
Melanoplus f:-r. femur- 
rubrum 
Encoptolophus s. sordidus 
Golden; brown striped 
Golden; brown striped 
Golden; brown striped 
Yellow-green 
Brown 
Olive 
Yellow-green 
Yellow-brown 
Golden; brown striped 
Yellow-white 
Yellow-white with 
reddish brown tips 
Yellow-brown striped 
Yellow 
Yellow-brown striped 
Amber 
Dark blue (distally); 
creamy brown (prox- 
imally) 
White 
Dull pink 
Pale golden 
Golden 
Golden 
Golden 
Y ellow-green 
Brown 
Olive 
Yellow-green 
Black 
Golden-brown 
striped 
Dark green 
Black 
Yellow-brown 
striped 
Yellow 
Yellow-brown 
striped 
Amber 
Creamy brown 
White 
Dull pink 
Golden brown & 
brown striped 
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Table 1 (Continued) 
Cruciferae 
Lepidium virginicum Golden; brown striped Golden; brown 
striped 
Golden 
Melanoplus confusus 
Melanoplus s. sanguinlpes Golden; brown striped 
Fabaceae 
L'espedeza sp. 
, , 
Melanoplus s. sanguinipes 
3 7 ,? 
Milky white 
White 
Milky white 
White 
Lythraceae 
Lythrum salicaria 
Umbelliferae 
Daucus carota 6 
9 
Scudderia c. curvicauda Purple Purple 2 
m 
Yellow Z 
Pale green z 
Transparent z 0 
* 
Dark brown z m (starved animal) t j  
Black (starved 
animal) 2 
Black (starved 
animal) g E 
r n  
Spharagemon b. bolli 
Melanoplus confusus 
Yellow 
Pale green 
Golden 
Asclepiadaceae 
Asclepias tuberosa Melanoplus s. sanguinipes Dark brown (starved 
animal) 
Black (starved 
animal) 
Black (starved 
animal) 
Melanoplus confusus 
9 ,  
Labiatae 
Monarda fistulosa 
3 3,  
4 Y ellow-white 
Yellow t o  gold 
White 
Melanoplus s. sanguinipes 
9, 9 ,  7, Golden green 
White 
Composit ae 
Galinosoga parviflora 
Lactuca sp. 
Melanoplus s. sanguinipes 
Pardalophora apiculata 
Yellow-brown striped 
Pale t o  bright yellow 
(distally t o  prox- 
imally) 
Whitish yellow 
Green 
Golden 
Yellow 
Cream 
Whitish yellow 
Pale black - 1 3  
Golden 
Melanoplus s. sanguinipes 
Melanoplus confusus 
9, 99 
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The Feasibility of Use of Caecal and Diverticular Coloration in Field Determination of Grasshopper Diet

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