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Feeding Habits of the Predaceous Nymphs of Dolania Americana in Northwestern Florida (Ephemeroptera : Behningiidae)

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Feeding Habits of the Predaceous Nymphs of Dolania Americana in Northwestern Florida (Ephemeroptera : Behningiidae) Hydrobiologia vol . 67, 2, pag. 119-123, 1979 FEEDING HABITS OF THE PREDACEOUS NYMPHS OF DOLANIA AMERICANA IN NORTHWESTERN FLORIDA (EPHEMEROPTERA : BEHNINGIIDAE) Philip T. P. TSUI* & Michael D . HUBBARD Laboratory of Aquatic Entomology, Florida A & M University, Tallahassee, Florida 32307 USA * Present Address : Mobil Oil Canada, Ltd., Environmental Planning Management Dept ., Box 800, Calgary, Alberta T2P2J7, Canada Received November 9, 1978 Keywords : Behningiidae, Chironomidae, competition, Dolania, Ephemeroptera, feeding, mayflies, predation . Abstract Dolania is a monotypic genus of the family Behningii- dae which also contains the Palaearctic genera Behningia Feeding habits of nymphs of the predatory mayfly Dolania ame- Lestage, with three species, and Protobehningia Tsherno- ricana Edmunds & Traver in northwestern Florida were va, with one species . Dolania americana is known from examined during 1971 and 1972 . Larvae of Chironomidae (Dip- northwestern Florida, southwestern Louisiana, and the tera) formed the bulk of prey items . The size of the predaceous D. americana nymphs was correlated with the size of chironomid Savannah River drainage in Georgia and South Carolina . prey taken. Differential microhabit utilization allows partition Further general biological and zoogeographic data on D . of prey resources with the other major predator species present. americana can be found in Peters & Peters (1977) . The relationship between predator and prey biomass throughout Nymphs examined in this study were collected from two the year is discussed . sampling stations on the Blackwater River . Both sites were located in the vicinity of the Florida A & M Universi- ty Biological Field Station near Holt, Okaloosa County, Introduction Florida . The physical and chemical characteristics of the Blackwater River were described by Beck (1973), and the During a long-term study of the sand burrowing mayfly, flora and fauna were discussed by Peters & Jones (1973) . Dolania americana Edmunds & Traver, in the Black- water River, northwestern Florida, we discovered that its nymphs are carnivorous . Predatory feeding is relatively Methods uncommon among the predominately detritivorous- herbivorous order Ephemeroptera and we felt that the Nine collections of D. americana nymphs were made be- feeding habits of D. americana merited further investiga- tween June, 1971, and May, 1972, in which a total of 208 tion . The objectives of the present study were : to deter- nymphs were dissected for gut analyses. Each nymph mine the nature of the food (prey) consumed by the examined for gut contents was split meso-ventrally and nymphs of D . americana; to determine whether food the entire gut was removed and spread in a drop of CMC- consumption showed seasonal variations ; to determine 10 (Turtox) mounting medium on a standard micro- the size relationship between the nymphs of D. americana scope slide . The entire gut was examined and the contents and their prey, if any; and to examine resource partition- enumerated . Size of the prey organisms was determined ing among macroinvertebrate predators in the Black- (in the case of dipteran larvae, the length of the head cap- water River . sule was measured), and the size of the D. americana nymphs was measured by the interorbital distance (shortest distance between the eyes) . Dr. W. Junk b. v. Publishers - The Hague, The Netherlands Estimates of chironomid (Diptera) population densi- "9 ties at the study areas were made in the months of February, March, and April, 1974 . Five one-litre sand samples were taken at hazard each month from the river bottom, preserved in ethanol, and stained with Rose Ben- gal (10o mg 1 ' preservative) . The entire sample was examined and the total number of dipteran larvae in each JUN litre of sand was determined . Sand particle size distribution was examined by running a transect perpendicular to the shore and taking 1 periodic bottom samples . The samples were then AUG ,-,+ + + + + processed through standard soil sieves, separated into 9 size-classes, air-dried, and weighed, and percentage com- SEP + + + + + + + + *TTrt + + + + 7 position was calculated. Horizontal and vertical distribution of macroinverte- I brate predators was examined by taking periodic samples OCT moo + +TTL + + + + along a transect perpendicular to the shore . The samples JAN + + + + + . + + were examined and the density and kind of predators 1 JIONIL+ + + + present were determined . 9 FEB + + + 4--- + + + • IIII • + + + + 7 Results and discussion 2 MAR +++4--46++ . In the Blackwater River nymphs of D. americana are found burrowing in about the top five centimetres of APR + +fir+ + + clean, shifting sand . The shifting sand biotope is one of I I I I I I I I+ I I I I I 1 I 1# I 0 5 10 15 the major habitats found in the river and occupies a great SIZE CLASS IN MM majority of the river bottom. The sand is loose and con- Fig. I . Seasonal size (body length) distribution of tinuously shifting with grain sizes mostly in the range of Dolania americana nymphs in the Blackwater River. 0.25-1 .0 mm. Morphologically, the nymph is well adapted for burrowing in this substrate and has evolved various protective structures to counteract the molar action of the acheived through differential microhabitat utilization . In shifting sand grains (Edmunds & Traver, 1959). Prelimi- contrast to the burrowing habit of D. americana and P. nary studies indicated that D. americana has primarily a obscurus, the nymphs of P. meridionalis occupy the univoltine life cycle in the Blackwater River, and nymphs surface of the sand, not burrowing beneath the substrate . are present in the river throughout the year (Fig . 1). Emer- In addition to this vertical partitioning, horizontal parti- gence occurs in late April to May (Peters & Peters, 1977). tioning also occurs . Nymphs of P. obscurus burrow A smaller cohort of D. americana nymphs in the Black- mostly at the peripheral areas of the stream, whereas water River apparently does not emerge the first year but D. americana nymphs burrow in more submarginal to remains in the river until the following spring . The reasons midstream areas. Samples along transects from the for this dichotomy remain unknown . shore reveal that the density of P. obscurus lessens with Other predators coexist with D. americana in lesser increasing water depth (i.e., distance from shore) and that densities in the shifting sand biotope of the Blackwater the density of D. americana increases concurrently until River. The two major competitors of D. americana are almost no P. obscurus occur (Fig . 2). Shore transects also the dragonfly Progomphus obscurus (Rambur) (Odo- indicate that the sand particle size of the substrate like- nata : Gomphidae) and the mayfly Pseudiron meridio- wise increases with increasing water depth, with the per- nalis Traver (Ephemeroptera : Heptageniidae) . Prelimi- centage of sand particles 0 .5 mm or greater in diameter nary study indicated that although a dietary overlap oc- increasing from about 30% at 50 mm depth to 65% at 350- curs among these three carnivores, partitioning of re- 500 mm depth . sources and a lessening of potential competition is Based on gut analyses of D. americana nymphs during 1 2 0 4 0 Progomphue obsourue months after hatching from the egg stage . Obvious varia- Dolania americana tion in size range of the nymphs occurred throughout the 3 study period. Such size variability within the predator s 0 2 population might be an important adaptive feature if the s feeding habits of the predator are size-dependent since this would allow better partition of the available food re- -o o sources . Fig. 3 is a graphic presentation of the size relationship 10 20 30 40 50 WATER DEPTH (CM) between the nymphs of D. americana and their chirono- mid prey . Small larvae were consumed by nymphs of all Fig . 2 . Density of nymphs of Dolania americana and Progomphus obscurus along a transect perpendicular to sizes, as illustrated by the essentially unchanging lower the shore in the Blackwater River (Kennedy Bridge, 16-11- size limits of the prey, while the upper, mean, and median 1 974) . size limits increased with increasing predator size, in- dicating a size dependent relationship on the upper end of the prey size distribution . The product moment correla- the study period we discovered that 95% or more of the tion coefficients for predator size and the largest, mean, prey organisms were Chironomidae larvae (Diptera). The and median prey sizes were .732, .707, and .817 respective- overwhelming majority of these chironomid larvae ly. All of these values are statistically significant (p < .01). belong to an undescribed species in the subfamily Chiro- Thus, as nymphs of D. americana grow larger, they con- nominae . Other prey organisms found in much lesser sume an increasingly wider range of chironomid prey numbers in gut contents of D. americana included micro- size . This fact, coupled with the extensive variation in size crustacea, ceratopogonid larvae, nematodes, and range of the Dolania nymphs at any one time, results in occasionally tardigrades . These food organisms are all a greatly lessened prey size overlap . This feeding pattern is elements of the interstitial fauna of the sandy river similar to that of another aquatic insect predator, Doro- bottom . Keffermiiller (1959), in a brief investigation of neuria californica (Banks) (Plecoptera : Perlidae), re- gut contents of the related species Behningia lestagei ported (as Acroneuria) by Sheldon (1969). Motas & Bacesco in the Warta River, Poland, was unable to determine the type of food utilized except that the 0 .7 standard digestive canal contained traces of `small grains of sand, deviation diatoms, small pieces of sponge spicules, and an in- 0 .8 mean determinable organic matter'. • 1 .0-I .1 Because chironomid larvae formed the main bulk of 1 .2-1 .3 the prey items consumed by Dolania nymphs we there- range • 1 .4-1 .5 fore focused our attention on this prey category. Mean • 1 .6-1 .7 numbers of chironomid larvae per litre of sand were 65, • 76, and 9o for the months of February, March, and April, 1 .8-1 .9 1974, respectively .
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