Taxonomy and Natural History of Cattail Aphids, Rhopalosiphum enigmae (Hemiptera: Aphidomorpha: Aphididae), Including a New Synonymy and Notes on Ant and Parasitoid Associates of Rhopalosiphum Skvarla, Michael J. Miller, Gary L. Bauchan, Gary Lewis, Matthew Foottit, Robert Maw, Eric Insect Systematics and Diversity 2018 2018-03-01 2 2018 2 1 14 KF KF [145,1765,558,577] Insecta Aphididae Melanaphis Animalia Aphidomorpha 5 6 Arthropoda species donacis    R. enigmaeAB Bonnyville Truman, Hwy55 54°28.074′N, 111°11.160′W  Typha latifolia 14-July-2009E. Maw, R.G. CNC#HEM063643 GU668766at Sand River Foottit   R. enigmaeON Ottawa Mer Bleu Cons. 45°24.252′N, 75°33.882′W  Typha latifolia 1-August-2008E. Maw et al. CNC#HEM061388 KR038471.1 Area   R. enigmaeON Ottawa Ottawa 45°23.652′N, 75°42.198′W  Typha latifolia 25-September-2009G. Miller, CNC#HEM064177 GU668786E. Maw   R. enigmaeDE KentFelton 39°00.730′N, 75°35.746′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-006 MF123452& G. Miller   R. enigmaeDE Sussex Georgetown 38°44.570′N, 75°25.602′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-001 MF123417& G. Miller   R. enigmaeDE Sussex Selbyville 38°28.778′N, 75°14.093′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-004 & G. Miller   R. enigmaeDE Sussex Selbyville 38°28.778′N, 75°14.093′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-005 MF123422& G. Miller   R. enigmaeFL Collier Immokalee Suctiontrap 10–17-February-2016S. Halbert E2016-536-2 MF123420   R. enigmaeFL Collier Immokalee Suctiontrap 24-February-2016– S. Halbert E2016-731-1 MF123449 2-March-2016   R. enigmaeFL Polk Lake Alfred 28°08.869′N, 81°44.301′W  Typha 25-September-2016M. J. Skvarla MS 16-0925-001 MF123434   R. enigmaeFL Polk Winter Haven 28°03.350′N, 81°44.050′Wsuction trap 11–18-February-2016S. Halbert E2016-535-2 MF123437   R. enigmaeFL St. Johns St. Augustine 29°54.959′N, 81°24.846′W  Typha 22-September-2016M. J. Skvarla MS 16-0922-001 MF123448   R. enigmaeFL Walton Point Washington 30°22.185′N, 86°06.391′W  Typha 25-April-2016K. E. Schnepp KS 16-0425-001 MF123433   R. enigmaeGA McIntosh Darien 31°22.536′N, 81°25.862′W  Typha 22-September-2016M. J. Skvarla MS 16-0922-003 MF123435   R. enigmaeMD Anne Arundel Russett 39° 06.267′N, 76°48.004′W  Typha 10-May-2016M. J. Skvarla MS 16-0510-001 MF123421   R. enigmaeMD Anne Arundel Russett 39° 06.267′N, 76°48.004′W  Typha 21-April-2016M. J. Skvarla MS 16-0421-001 MF123425   R. enigmaeMD Anne Arundel Russett 39° 06.267′N, 76°48.004′W  Typha 19-April-2017M. J. Skvarla MS 16-0419-001   R. enigmaeMD Dorchester Linkwood 38°33.105′N, 75°57.521′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-009 MF123453, MF123430& G. Miller   R. enigmaeMD Frederick Frederick 39°22.860′N, 77°24.410′W  Typha 15-June-2016M. J. Skvarla MS 16-0615-001 MF123450, MF123447   R. enigmaeMDQueen Anne’s Chester 38°57.294′N, 76°17.981′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-002 MF123432& G. Miller   R. enigmaeMDQueen Anne’s Grasonville 38°57.850′N, 76°13.253′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-008 MF123424, MF123439& G. Miller   R. enigmaeMDQueen Anne’s Grasonville 38°57.850′N, 76°13.253′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-010 & G. Miller   R. enigmaeMD Talbot Easton 38°49.014′N, 76°03.686′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-003 MF123418, MF123445& G. Miller   R. enigmaeMD WashingtonHancock 39°42.142′N, 78°11.429′W  Typha 15-July-2016M. J. Skvarla MS 16-0715-002   R. enigmaeMD WashingtonHancock 39°42.142′N, 78°11.429′W  Typha 15-July-2016M. J. Skvarla MS 16-0715-003   R. enigmaeMD Wicomico Salisbury 38°22.178′N, 75°32.160′W  Typha 28-July-2016M. J. Skvarla MS 16-0728-007 MF123431& G. Miller   Table 3.Continued    Plant associa-  tion/ collection GenBank Accession  Species State/ province County Locality Coordinates method Date Collected by Collection number number    R. enigmae NC Brunswick Bolivia 34°02.502′N, 78°14.617′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-011    R. enigmae NC Carteret Bouge 34°41.834′N, 77°03.287′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-010    R. enigmae NC Craven Ernul 35°14.611′N, 77°03.616′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-001    R. enigmae NC Craven New Bern 35°02.983′N, 77°00.086′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-002    R. enigmae NC Halifax Tillery 36°13.909′N, 77°27.336′W   Typha 20-September-2016 M. J. Skvarla MS 16-0920-004    R. enigmae NC Onslow Jacksonville 34°44.522′N, 77°29.848′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-005    R. enigmae NC Onslow Swansboro 34°41.544′N, 77°07.147′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-003    R. enigmae NC Onslow Swansboro 34°41.544′N, 77°07.147′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-004    R. enigmae NC Pender Montague 34°27.275′N, 78°03.059′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-009    R. enigmae NC Pender Watha 34°38.618′N, 77°54.196′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-006    R. enigmae NC Pitt Greenville 35°30.425′N, 77°19.405′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-007    R. enigmae NC Pitt Greenville 35°30.425′N, 77°19.405′W   Typha 21-September-2016 M. J. Skvarla MS 16-0921-008    R. enigmae PA Adams Gettysburg NMP 39°48.106′N, 77°14.078′W   Typha 09-June-2016 M. J. Skvarla MS 16-0609-001    R. enigmae PA Bedford Bedford 40°02.120′N, 78°31.200′W   Typha 15-July-2016 M. J. Skvarla MS 16-0715-004  MF123440    R. enigmae PA Lancaster Lancaster 40°02.602′N, 76°14.706′W   Typha 4-June-2016 M. J. Skvarla MS 16-0603-001    R. enigmae PA Lancaster New Danville 39°59.175′N, 76°19.523′W   Typha 4-June-2016 M. J. Skvarla MS 16-0603-002    R. enigmae PA Westmoreland Ligonier 40°15.874′N, 72°16.021′W   Typha 15-July-2016 M. J. Skvarla MS 16-0715-005  MF123416, MF123443    R. enigmae PA Westmoreland Paintertown 40°22.115′N, 79°42.011′W   Typha 6-May-2016 M. J. Skvarla MS 16-0506-001    R. enigmae PA Westmoreland Paintertown 40°22.115′N, 79°42.011′W   Typha 6-May-2016 M. J. Skvarla MS 16-0506-002    R. enigmae SC Colleton Waterboro 32°52.612′N, 80°42.765′W   Typha 22-September-2016 M. J. Skvarla MS 16-0922-002  MF123427, MF123426    R. enigmae SC Jasper Hardeeville 32°16.336′N, 81°04.678′W   Typha 22-September-2016 M. J. Skvarla MS 16-0922-004  MF123451, MF123444    R. enigmae SC Sumter Pinewood 33°44.301′N, 80°27.855′W   Typha 22-September-2016 M. J. Skvarla MS 16-0922-005  MF123413    R. enigmae VA Chesterfield Chester 37°21.139′N, 77°24.199′W   Typha 20-September-2016 M. J. Skvarla MS 16-0920-002    R. enigmae VA Chesterfield Chester 37°21.139′N, 77°24.199′W   Typha 20-September-2016 M. J. Skvarla MS 16-0920-003    R. enigmae WV Raleigh Bradley 37°52.831′N, 81°13.521′W   Iris? 7-May-2016 M. J. Skvarla MS 16-0507-001  MF123414, MF123436    R. enigmae WV Raleigh Bradley 37°52.831′N, 81°13.521′W   Iris? 7-May-2016 M. J. Skvarla MS 16-0507-002  MF123415    R. musae OR Lake 8 mi NW of   Prunus 3-May-2016 A. Jensen MS 16-1206-001  MF123446, MF123419  Lakeview  subcordata    R. musae EU179242.1    R. nymphaeae BC Vancouver Vancouver, Queen 54°41.160′N, 124°56.220′W   Callitriche 12-June-2005 C.-K. Chan CNC#HEM054279 KR045003.1  Elizabeth Park  stagnali    R. nymphaeae MD Frederick Adamstown 39°17.696′N, 77°25.887′W   Nymphaea 14-September-2016 M. J. Skvarla MS 16-0914-001  MF123438, MF123441,   MF123442    R. nymphaeae ON Ottawa Ottawa 45°23.580′N, 75°42.240′W   Butomus 2-October-2009 E. Maw CNC#HEM064179   umbellatus    R. nymphaeae OR Lake 3 mi N of   Alisma 1-September-2016 A. Jensen MS 16-1206-002  MF123411  Lakeview member of the subtribe Rhopalosiphina, is thought to be closely related to  Rhopalosiphumand was used to root the phylogenetic tree. The COI sequence for  M. donaciswas obtained from GenBank ( Table 3).  R. musaeand  R. nymphaeaewere included in order to help root the phylogenetic tree, and to determine the percent difference in COI between species and thus provide a baseline comparison for the percent difference in COI between  R. enigmaeand  R. laconae. In addition to COI sequences produced de novo and available through GenBank, seven  R. enigmae, one  R. nymphaeae, and one  R. musaewere available through the Barcode of Life Database and included in analyses ( Table 3). Bayesian analyses were performed with MrBayes (3.2.6) using the Extreme Science and Engineering Discovery Environment (XSEDE) infrastructure on the Cipres Portal ( Miller et al. 2010). Each analysis consisted of four simultaneous runs, each with four chains sampling every 1,000 generations for 1.11 million generations, under a GTR+I+Γ model of molecular evolution. The analysis was automatically ended when the split frequencies fell below 0.01; 25% of the trees were discarded as burn-in. The resulting majority-rule consensus trees were viewed with Dendroscope 3 (v. 3.5.7) ( Huson and Scornavacca 2012); tree image files were then exported in PDF format and edited for final figures in Adobe Illustrator CS6 (Adobe Systems, San Jose, CA). Percent difference in COI was determined by comparing sequence data for pairs of individuals in BioEdit ( Hall 1999) using the ‘calculate identity/similarity for two sequences’ function. At least one such pair comparison was made within and between each clade and additional within-clade comparisons were made when multiple morphologies existed within a single clade, such that 19 pair comparisons were made within the larger  R. enigmae+  R. laconaeclade, 1 pair comparison was made within the  R. musaeclade, 2 pair comparisons were made within the  R. nymphaeaeclade, and 1 pair comparison was made between  R. enigmae+  R. laconaeand each of the outgroup clades.