ABSTRACT

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SOUTHEASTERN EPT: A GUIDE FOR IDENTIFICATION OF SPECIES OF MAYFLY, STONEFLY, AND CADDISFLY LARVAE OF THE SOUTHEASTERN UNITED STATES (EPA REGION IV) John C. Morse 1 , W. Patrick McCafferty 2 , Bill P. Stark 3 , Ralph W. Holzenthal 4 , Luke M. Jacobus 5 and Nick A. Wiersema 6 1 Clemson University, Clemson, SC, USA, [email protected] , 2 Purdue University, West Lafayette, IN, USA, [email protected] , 3 Mississippi College, Clinton, MS, USA, [email protected] , 4 University of Minnesota, St. Paul, MN, USA, [email protected] , 5 Indiana University Purdue University Columbus, Columbus, IN, USA, [email protected] , 6 NA Wiersema Environmental Consulting LLC, Austin, TX, USA, [email protected] ABSTRACT Species diversity of mayflies, stoneflies, and caddisflies (Ephemeroptera, Plecoptera, and Trichoptera, respectively, or EPT) in the southeastern United States is high, with at least 279 species of mayflies, 267 species of stoneflies, and 651 species of caddisflies known or likely to occur in EPA Region 4 (Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, South Carolina, and Tennessee). Larvae of 265 mayfly species (95%), 166 stonefly species (62%), and 294 caddisfly species (45%) are sufficiently known to permit at least tentative identification with diagnostic keys. An illustrated guide to larvae of these identifiable species will soon be available. In this work, identification is facilitated with dichotomous keys, with supporting illustrations generally on the same page as the diagnostic text or on an adjacent page. Keys for southeastern EPT species of larvae in several genera are provided for the first time. Benthologists studying faunas in nearby states also will likely find these keys useful. The work will be available from Clemson University Public Service Publishing http://www.clemson.edu/psapub lishing / INTRODUCTION The work of Brigham et al. (1982) included species-level identification aids for larvae of mayflies, stoneflies, and caddisflies in the Carolinas, USA. It revealed the limits of our knowledge for species diagnosis at that time and stimulated considerable subsequent research to fill the gaps. We are preparing keys to synthesize diagnostic characters discovered and described in the past 30 years for EPT species occurring or EPHEMEROPTERA W.P. McCafferty, L.M. Jacobus and N.A. Wiersema With respect to mayflies, the southeastern United States is by far the richest and most diverse area of such size in the world. The approximately 279 species in the area (McCafferty et al. 2010) represent over half of those known from the United States and very nearly half of those known from the entire North American continent (McCafferty 2011). CAENIDAE, Caenis spp. 108(103) Dorsal face of forefemora with transverse row of spatulate setae in distal half (e.g., Fig. 2.209); posterior margin of abdominal sternum 9 strongly to barely notched medially (Figs. 2.210, 2.211); abdominal terga 9 and 10 without triad of black dots; abdominal sterna without submedian pairs of black dots . . . . . . . . . . . . .109 108’ Dorsal face of forefemora with scattered simple setae only (e.g., Fig. 2.212); posterior margin of abdominal sternum 9 convexly rounded or straight (Figs. 2.213, 2.214); abdominal terga 9 and 10 usually with triad of black dots; abdominal sterna often with pairs of submedian black dots . . . . . . . . . . . . . . . . . . . . . . . . . . .11 2 109(108) Middle third of caudal filaments (tails) with whorls of short setae on every second or third segment, with setae much shorter than distance between base of whorls (Fig. 2.215) . . . . . . . . . . . . . . . . . . . . Caenis anceps 109’ Middle third of caudal filaments with long lateral setae on every segment, with setae much longer than distance between their bases . . . . . . . . . . . . . . . . . . . . 110 Fig. 2.210 PLECOPTERA B.P. Stark The order Plecoptera includes approximately 3350 extant world species of stoneflies placed in 16 families (DeWalt et al. 2010; Stark et al. 2009). In the Southeast, all nine North American families and 267 species are known. PERLIDAE, Acroneuria spp. (Modified from Hitchcock, 1974. Characters based primarily on pigment patterns are available for 11 of 16 regional species.) 97(37’) Anal gills present on paraprocts (Fig. 3.220) . . . . .98 97’ Anal gills absent from paraprocts . . . . . . . . . . . . . . . . 105 98(97) M-line on head interrupted by dark pigment separating pale band into three pale spots (Fig. 3.221), or head almost completely dark . . . . . . . . . . . . . . . . . . . . . . . 99 98’ M-line on head complete (Fig. 3.222) . . . . . . . . . . . . .100 99(98) Abdominal terga without distinctive pigment bands; predator/engulfers; clingers; AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, OH, OK, PA, WV . . . . . . . . . . . . Acroneuria evoluta 99’ Most abdominal terga with an incomplete, narrow, pale basal band (Fig. 3.223); predator/engulfers; clingers; AL, CT, DC, DE, FL, GA, LA, MA, MD, ME, MS, NC, NJ, NY, PA, PQ, SC, TX, VA, WV . . . . . . . . . . . . . . . . . . . . . . . .Acroneuria arenosa 100(98’) Abdominal terga with broad, pale basal and narrow, dark apical pigment bands (Fig. 3.224); predator/engulfers; clingers; CT, KY, IA, MB, ME, MI, MN, NB, NC, ND, NS, NY, OH, ON, PA, PQ, SK, TN, VA, WI, WV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acroneuria lycorias 100’ Abdominal terga generally with dark basal and pale apical pigment bands (Fig. 3.225) . . . . . . . . . . . . . . . . . . . 101 TRICHOPTERA J.C. Morse and R.W. Holzenthal Of the 1,455 species of caddisflies known from the North American continent north of Mexico (Morse 2011), 651 species (44%) have been reported from in or near the southeastern states, representing 23 of the 27 North American families. BRACHYCENTRIDAE, Micrasema spp. (Modified from Chapin, 1978) 38(25’) Mesonotal sclerites partially or completely divided into 4 plates (Figs. 4.111, 4.112); ventral apotome rectangular, broader than long (Figs. 4.114) or longer than broad (Fig. 4.115); east of Great Plains . . . . . . . . . . . . . . . . . . . . . . . . . . Micrasema rusticum Group, 39 38’ Mesonotal sclerites entire, consisting of 2 plates (Fig. 4.113); ventral apotome trapezoidal (Fig. 4.116); small cold streams, typically in Fontinalis mats grazing on moss leaves and detritus; QC to NB and south to NC and SC . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micrasema sprulesi 39(38) Ventral apotome longer than broad (Fig. 4.115); head dark brown, muscle scars not apparent (Fig. 4.117); case curved, typically of silk but may incorporate some sand (Fig. 4.118); springs and small cold streams of Appalachian Mountains of NY, VA, and GA, NC, SC, TN . . .Micrasema burksi 39’ Ventral apotome broader than long (Fig. 4.114); head, case, and habitat variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40 Fig. 4.111 Fig. 4.112 Fig. 4.113 Fig. 4.114 Fig. 4.115 Fig. 4.116 Fig. 4.117 Fig. 4.118 REFERENCES CITED Brigham, A.R., Brigham, W.U. & Gnilka, A. (Eds.) 1982. Aquatic insects and oligochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. 837 pp. Chapin, J.W. 1978. Systematics of Nearctic Micrasema (Trichoptera: Brachycentridae). Ph.D. dissertation, Clemson University, Clemson, SC. 136 pp. DeWalt, R. E., Neu-Becker, U., and Steuber, G. 2010. Plecoptera species file online .Version 1.1/4.0. Available from http://Plecoptera.SpeciesFile.org/HomePage.aspx (accessed 23 July 2010). Hitchcock, S. W. 1974. Guide to the insects of Connecticut. Part VII. The Plecoptera or stoneflies of Connecticut. State Geological and Natural History Survey of Connecticut, 107:1-262. McCafferty, W. P. 2011. North American mayfly list. Available from http://www.entm.purdue.edu/mayfly/ (accessed 1 August 2011). McCafferty, W. P., D. R. Lenat, L. M. Jacobus, and M. D. Meyer. 2010. The mayflies (Ephemeroptera) of the southeastern United States. Transactions of the Entomological Society of America 136: 221-233.

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SOUTHEASTERN EPT: A GUIDE FOR IDENTIFICATION OF SPECIES OF MAYFLY, STONEFLY, AND CADDISFLY LARVAE OF THE SOUTHEASTERN UNITED STATES (EPA REGION IV)

John C. Morse1, W. Patrick McCafferty2, Bill P. Stark3, Ralph W. Holzenthal4, Luke M. Jacobus5 and Nick A. Wiersema61Clemson University, Clemson, SC, USA, [email protected] ,2Purdue University, West Lafayette, IN, USA, [email protected] , 3Mississippi College, Clinton, MS, USA, [email protected] ,

4University of Minnesota, St. Paul, MN, USA, [email protected] , 5Indiana University Purdue University Columbus, Columbus, IN, USA, [email protected] , 6NA Wiersema Environmental Consulting LLC, Austin, TX, USA, [email protected]

ABSTRACTSpecies diversity of mayflies, stoneflies, and caddisflies (Ephemeroptera, Plecoptera, and Trichoptera, respectively, or EPT) in the southeastern United States is high, with at least 279 species of mayflies, 267 species of stoneflies, and 651 species of caddisflies known or likely to occur in EPA Region 4 (Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, South Carolina, and Tennessee). Larvae of 265 mayfly species (95%), 166 stonefly species (62%), and 294 caddisfly species (45%) are sufficiently known to permit at least tentative identification with diagnostic keys. An illustrated guide to larvae of these identifiable species will soon be available. In this work, identification is facilitated with dichotomous keys, with supporting illustrations generally on the same page as the diagnostic text or on an adjacent page. Keys for southeastern EPT species of larvae in several genera are provided for the first time. Benthologists studying faunas in nearby states also will likely find these keys useful. The work will be available from Clemson University Public Service Publishing http://www.clemson.edu/psapublishing/

INTRODUCTIONThe work of Brigham et al. (1982) included species-level identification aids for larvae of mayflies, stoneflies, and caddisflies in the Carolinas, USA. It revealed the limits of our knowledge for species diagnosis at that time and stimulated considerable subsequent research to fill the gaps. We are preparing keys to synthesize diagnostic characters discovered and described in the past 30 years for EPT species occurring or likely to occur in the Southeast. Keys identify EPT larvae to family, genus, and species. Illustrations are original or borrowed.

EPHEMEROPTERAW.P. McCafferty, L.M. Jacobus and N.A. Wiersema With respect to mayflies, the southeastern United States is by far the richest and most diverse area of such size in the world. The approximately 279 species in the area (McCafferty et al. 2010) represent over half of those known from the United States and very nearly half of those known from the entire North American continent (McCafferty 2011).

CAENIDAE, Caenis spp. 108(103) Dorsal face of forefemora with transverse row of spatulate setae in distal half (e.g., Fig. 2.209); posterior margin of abdominal sternum 9 strongly to barely notched medially (Figs. 2.210, 2.211); abdominal terga 9 and 10 without triad of black dots; abdominal sterna without submedian pairs of black dots . . . . . . . . . . . . .109108’ Dorsal face of forefemora with scattered simple setae only (e.g., Fig. 2.212); posterior margin of abdominal sternum 9 convexly rounded or straight (Figs. 2.213, 2.214); abdominal terga 9 and 10 usually with triad of black dots; abdominal sterna often with pairs of submedian black dots . . . . . . . . . . . . . . . . . . . . . . . . . . .112 109(108) Middle third of caudal filaments (tails) with whorls of short setae on every second or third segment, with setae much shorter than distance between base of whorls (Fig. 2.215) . . . . . . . . . . . . . . . . . . . . Caenis anceps109’ Middle third of caudal filaments with long lateral setae on every segment, with setae much longer than distance between their bases . . . . . . . . . . . . . . . . . . . . 110

Fig. 2.210

Fig. 2.209 Fig. 2.211 Fig. 2.212

Fig. 2.213 Fig. 2.214 Fig. 2.215

PLECOPTERAB.P. Stark

The order Plecoptera includes approximately 3350 extant world species of stoneflies placed in 16 families (DeWalt et al. 2010; Stark et al. 2009). In the Southeast, all nine North American families and 267 species are known.

PERLIDAE, Acroneuria spp. (Modified from Hitchcock, 1974. Characters based primarily on pigment patterns are available for 11 of 16 regional species.) 97(37’) Anal gills present on paraprocts (Fig. 3.220) . . . . .9897’ Anal gills absent from paraprocts . . . . . . . . . . . . . . . . 105 98(97) M-line on head interrupted by dark pigment separating pale band into three pale spots (Fig. 3.221), or head almost completely dark . . . . . . . . . . . . . . . . . . . . . . . 9998’ M-line on head complete (Fig. 3.222) . . . . . . . . . . . . .100 99(98) Abdominal terga without distinctive pigment bands; predator/engulfers; clingers; AL, AR, FL, GA, IL, IN, KS, LA, MO, MS, OH, OK, PA, WV . . . . . . . . . . . . Acroneuria evoluta 99’ Most abdominal terga with an incomplete, narrow, pale basal band (Fig. 3.223); predator/engulfers; clingers; AL, CT, DC, DE, FL, GA, LA, MA, MD, ME, MS, NC, NJ, NY, PA, PQ, SC, TX, VA, WV . . . . . . . . . . . . . . . . . . . . . . . .Acroneuria arenosa 100(98’) Abdominal terga with broad, pale basal and narrow, dark apical pigment bands (Fig. 3.224); predator/engulfers; clingers; CT, KY, IA, MB, ME, MI, MN, NB, NC, ND, NS, NY, OH, ON, PA, PQ, SK, TN, VA, WI, WV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acroneuria lycorias100’ Abdominal terga generally with dark basal and pale apical pigment bands (Fig. 3.225) . . . . . . . . . . . . . . . . . . . 101

Fig. 3.220 Fig. 3.221 Fig. 3.222

Fig. 3.223 Fig. 3.224 Fig. 3.225

ACKNOWLEDGEMENTS

TRICHOPTERAJ.C. Morse and R.W. Holzenthal

Of the 1,455 species of caddisflies known from the North American continent north of Mexico (Morse 2011), 651 species (44%) have been reported from in or near the southeastern states, representing 23 of the 27 North American families.

BRACHYCENTRIDAE, Micrasema spp. (Modified from Chapin, 1978)

38(25’) Mesonotal sclerites partially or completely divided into 4 plates (Figs. 4.111, 4.112); ventral apotome rectangular, broader than long (Figs. 4.114) or longer than broad (Fig. 4.115); east of Great Plains . . . . . . . . . . . . . . . . . . . . . . . . . . Micrasema rusticum Group, 3938’ Mesonotal sclerites entire, consisting of 2 plates (Fig. 4.113); ventral apotome trapezoidal (Fig. 4.116); small cold streams, typically in Fontinalis mats grazing on moss leaves and detritus; QC to NB and south to NC and SC . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micrasema sprulesi

39(38) Ventral apotome longer than broad (Fig. 4.115); head dark brown, muscle scars not apparent (Fig. 4.117); case curved, typically of silk but may incorporate some sand (Fig. 4.118); springs and small cold streams of Appalachian Mountains of NY, VA, and GA, NC, SC, TN . . .Micrasema burksi39’ Ventral apotome broader than long (Fig. 4.114); head, case, and habitat variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . .40

Fig. 4.111 Fig. 4.112 Fig. 4.113

Fig. 4.114 Fig. 4.115 Fig. 4.116 Fig. 4.117 Fig. 4.118

REFERENCES CITEDBrigham, A.R., Brigham, W.U. & Gnilka, A. (Eds.) 1982. Aquatic insects and

oligochaetes of North and South Carolina. Midwest Aquatic Enterprises, Mahomet, Illinois. 837 pp.

Chapin, J.W. 1978. Systematics of Nearctic Micrasema (Trichoptera: Brachycentridae). Ph.D. dissertation, Clemson University, Clemson, SC. 136 pp.

DeWalt, R. E., Neu-Becker, U., and Steuber, G. 2010. Plecoptera species file online .Version 1.1/4.0. Available from http://Plecoptera.SpeciesFile.org/HomePage.aspx (accessed 23 July 2010).

Hitchcock, S. W. 1974. Guide to the insects of Connecticut. Part VII. The Plecoptera or stoneflies of Connecticut. State Geological and Natural History Survey of Connecticut, 107:1-262.

McCafferty, W. P. 2011. North American mayfly list. Available from http://www.entm.purdue.edu/mayfly/ (accessed 1 August 2011).

McCafferty, W. P., D. R. Lenat, L. M. Jacobus, and M. D. Meyer. 2010. The mayflies (Ephemeroptera) of the southeastern United States. Transactions of the Entomological Society of America 136: 221-233.

Morse, J.C. 2011. The Trichoptera World Checklist. Pp. 372-380 in K. Majecka, J. Majecki, & J. Morse (Eds.), Proceedings of the 13th International Symposium on Trichoptera, Zoosymposia 5. 512 pp.

Stark, B. P., Baumann, R. W., and DeWalt, R. E. 2009. Valid stonefly names for North America. Updated as of March 19, 2009. Available from http://plsa.inhs.uiuc.edu/plecoptera/ (accessed 23 October 2009).