Hierarchical Relationships of North American States and Provinces: an Area Cladistic Analysis Based on the Distribution of Stoneflies (Insecta: Plecoptera)

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Zeitschrift/Journal: Illiesia

Jahr/Year: 2008

Band/Volume: 04

Autor(en)/Author(s): Nelson Charles H.

Artikel/Article: Hierarchical relationships of North American States and Provinces: an area cladistic analysis based on the distribution of stoneflies (Insecta: Plecoptera). 176-204

Nelson, C.H. 2008. Hierarchical relationships of North American states and provinces: An area cladistic analysis based on the distribution of stoneflies (Insecta: Plecoptera). Illiesia, 4(18):176‐204. Available online: http://www2.pms‐lj.si/illiesia/Illiesia04‐18.pdf

HIERARCHICAL RELATIONSHIPS OF NORTH AMERICAN STATES AND PROVINCES: AN AREA CLADISTIC ANALYSIS BASED ON THE DISTRIBUTION OF STONEFLIES (INSECTA: PLECOPTERA)

Charles H. Nelson Department of Biological and Environmental Sciences, The University of Tennessee at Chattanooga, Chattanooga, Tennessee, U. S. A. 37403 E‐mail: charles‐[email protected]

ABSTRACT The distribution pattern of 67 genera, 507 species and 2 subspecies of North American Plecoptera is examined using a variant of Parsimony Analysis of Endemicity (PAE), Cladistic Analysis of Distributions and Endemisms (CADE), to assess hierarchical relationships of 85 large political units belonging to Canada, the USA and Mexico. Consensus area cladograms generated from the 146 minimum‐length area cladograms revealed three main assemblages of area clades: The states of northwestern Mexico (Chihuahua, Sonora, Baja California) together with the states and provinces of the western USA and Canada (Nunavut, Manitoba, Saskatchewan, Northwest Territories, Arizona, South Dakota, Nevada, New Mexico, Colorado, Utah, Wyoming, Alaska, Yukon, Montana, Idaho, Alberta, British Colombia, California, Oregon, Washington); the states and provinces of the eastern USA and Canada (Rhode Island, New Hampshire, Vermont, Newfoundland, Labrador, Prince Edward Island, Nebraska, North Dakota, Texas, Louisiana, Florida, Mississippi, Alabama, Georgia, South Carolina, North Carolina, Tennessee, District of Columbia, New Jersey, Delaware, Iowa, Minnesota, Wisconsin, Michigan, Ontario, Quebec, Nova Scotia, New Brunswick, Massachusetts, Connecticut, Maine, New York, Pennsylvania, Maryland, Virginia, West Virginia, Kansas, Oklahoma, Arkansas, Missouri, Ohio, Kentucky, Indiana, Illinois); and the remaining states of Mexico (Sinaloa, Tabasco, San Luis Potosi, Puebla, Tamaulipas, Nuevo Leon, Nayarit, Michoacan, Morelos, Distrito Federal, Mexico, Jalisco, Durango, Coahuila, Chiapas, Vera Cruz, Oaxaca, Guerrero). The major clades of the consensus area cladograms may be reflecting historical phenomena as these can be correlated with geologic events that either fragmented or joined principal North American land areas in the late Jurassic (clade 67), early and mid‐Cretaceous, and Miocene (clades 2 and 24). Low values for the consistency (CI), retention (RI) and rescaled consistency (RC) indices of the 146 minimum‐length area cladograms very likely reflect one or some combination of (1) the extinction of taxa; (2) the influence of random distribution of taxa; (3) the high number as well as the artificial nature of the geographic area units; or (4) incomplete information concerning the distribution of North American Plecoptera genera and species. The hierarchical patterns of this PAE/CADE analysis will need to be corroborated by species phylogenies of North American Plecoptera when they become available.

Keywords: Parsimony Analysis of Endemicity, PAE, Cladistic Analysis of Distributions and Endemisms, CADE, distribution, Plecoptera, states, provinces, territories, North America

INTRODUCTION North American Plecoptera have been confined to Comprehensive analyses on the biogeography of studies by Hynes (1988) and Stewart & Stark (1988,

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2002). These workers concluded that there were at species representatives of a large number of least three basic groupings of the North American Plecoptera genera, have generated a cladistic stonefly fauna: Eastern Boreal, Western Boreal, and hypothesis on family and generic relationships Austral. Moreover, they considered that these within the order, the state of knowledge regarding groupings are compatible with fragmentation and the phylogeny of species within each North merging of geographic areas associated with plate American genus remains largely incomplete. tectonics. The goal of this study is to add to these Therefore, a useful first‐step to assess geographic findings by determining North American area area relationships in North America would be to relationships using taxonomic and distributional data apply a method that relies less heavily on of plecopteran species. phylogenetic information. Based on the assumption A number of methodological approaches have that shared distribution patterns of a biota reflect an been employed to analyze the distributions of underlying common historical explanation, this organisms. Morrone (2005) organized these in a study employs a panbiogeographic method that taxonomy of methods that are initially divided into applies a variant of the Parsimony Analysis of two classes. The first consist of dispersalist Endemicity (PAE) approach termed Cladistic approaches that use dispersal from previously Analysis of Distributions and Endemisms (CADE) identified centers of origin or ancestral areas to (Cracraft 1991; Porzecanski & Cracraft 2005). This explain distribution of organisms. The second method uses parsimony to construct a cladogram include vicarance approaches. These consist of the depicting the hierarchical pattern of relationships methods of panbiogeography, which first construct a between geographic areas based on distributional track of distribution between related organisms and data of taxa. Geographic areas that have been used in then look for congruence between tracks of unrelated PAE analyses include artificially determined units organisms, and cladistic biogeography, which such as quadrants of varying sizes and natural units construct relationships between the geographic areas such as islands, oceans, continents, ecoregions or a group of organisms inhabit by employing the biogeographic regions. In this study states, provinces, phylogeny of that group. territories and federal districts of Canada, the USA, There appears to be agreement among those and Mexico were used as the area units. The use of workers who use vicariance approaches that it would PAE or CADE in biogeographic studies has been be preferable to apply the comparative phylogenetic criticized by Brooks and van Veller (2003), but has approach of cladistic biogeographic methods to been defended by Porcecanski & Cracraft (2005) and resolve the problem of the historical biogeography of Vazquez‐Miranda et al. (2007). a group of organisms (Cracraft 1991; Luna‐Vega et al. 1999; Morrone & Escalante 2002; Porzecanski & MATERIAL AND METHODS Cracraft 2005). When North American Plecoptera The basic requirements of CADE (Cracraft 1991; species are considered, only three such studies have Porcecanski & Cracraft 2005) are: been undertaken: Nelson’s (1988) analysis of the 1. predetermined areas of endemism; species of Pteronarcyidae using a reduced area 2. a sufficiently large sample of taxa to cladogram incorporating broad geographic regions, adequately reflect the region’s biota; Stark & Nelson’s (1994) analysis of the species of 3. incorporation of hierarchical information Yoraperla in the Peltoperlidae using Brooks into the data matrix by coding the Parsimony Analysis on Asian and western North distribution of subspecies, species, and American mountain chains, and Nelson’s (2004) genera; study of the Capnia californica species group using a 4. parsimony analysis of the data matrix to comparative phylogenetic analysis to assess produce a cladogram showing the zoogeographic affinities between six western states relationship between the endemic areas. and northwestern Mexico. A total of 85 states, provinces, territories and Although Terry & Whiting (In Preparation), using federal districts from which stoneflies have been morphological and molecular evidence derived from reported from Mexico, Canada, and the USA were

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Figure 1. Provinces, territories, states and one federal district representing endemic areas in Canada and the USA that report the presence of stoneflies. Canada: AB (Alberta), BC (British Columbia), LB (Labrador), MB (Manitoba), NB (New Brunswick), NE (Newfoundland), NS (Nova Scotia), NT (Northwest Territories), NU (Nunavut), ON (Ontario), PE (Prince Edward Island), PQ (Quebec), SK (Saskatchewan), YK (Yukon). USA: AL (Alabama), AK (Alaska), AR (Arkansas), AZ (Arizona), CA (California), CO (Colorado), CT (Connecticut), DC (District of Columbia), DE (Delaware), FL (Florida), GA (Georgia), IA (Iowa), ID (Idaho), IL (Illinois), IN (Indiana), KS (Kansas), KY (Kentucky), LA (Louisiana), MA (Massachusetts), MD (Maryland), ME (Maine), MI (Michigan), MN (Minnesota), MO (Missouri), MS (Mississippi), MT (Montana), NC (North Carolina), ND (North Dakota), NE (Nebraska), NH (New Hampshire), NJ (New Jersey), NM (New Mexico), NV (Nevada), NY (New York), OH (Ohio), OK (Oklahoma), OR (Oregon), PA (Pennsylvania), RI (Rhode Island), SC (South Carolina), SD (South Dakota), TN (Tennessee), TX (Texas), UT (Utah), VA (Virginia), VT (Vermont), WN (Washington), WI (Wisconsin), WV (West Virginia), WY (Wyoming).

considered to represent possible areas of endemism required by CADE. Nihei (2006), for example, has and were used in this analysis (Figs. 1, 2). To assist in criticized the use of non‐natural area units in PAE as the discernment of relationships between these North they may distort area relationships. Nonetheless, American area units, they were treated as belonging endemism is difficult to define. Crisci et al. (2003) list to a monophyletic entity despite published evidence five definitions for this concept and observe that of a number of taxon relationships between the defining endemism in biogeography is analogous to Eastern Palearctic and Western Nearctic, the Eastern the attempt of defining the species concept in Palearctic and Eastern Nearctic and the Western systematic biology. A second justification for using Palearctic and Eastern Nearctic. Since states, states, provinces, territories and federal districts is provinces, territories and federal districts are political that, although some Plecoptera species have a or artificial area units rather than natural area units, widespread distribution (Stewart & Stark 2002), most it can be argued that these are not the endemic areas members of this order have limited

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Fig. 2. States and federal district representing endemic areas in Mexico that report the presence of stoneflies. BJ (Baja California), CH (Chihuahua), CI (Coahuila), CP (Chiapas), DF (Distrito Federal), DG (Durango), GU (Guerero), JA (Jalisco), MC (Michoacan), MR (Morelos), MX (Mexico), NA (Nayarit), NL (Nuevo Leon), OX (Oaxaca), PU (Puebla), SI (Sinaloa), SL (San Luis Potosi), SO (Sonora), TA (Tamaulipas), TB (Tabasco), VZ (Vera Cruz).

dispersal abilities (Ross & Ricker 1971, Stark & apomorphic condition. A hypothetical outgroup is Gaufin 1976; Hynes 1988; Sargent et al. 1991; Stewart used since, as Rosen (1988) noted, there is no & Stark 2002) and are likely to be contained within a convincing means of identifying a ‘primitive’ area. few political area units. In this study 176 or 26% of Moreover, coding the outgroup with 1’s would the 688 Plecoptera taxa whose distribution were suggest that area relationships should be recognized examined are confined to one state, province, or by the shared loss of taxa or extinction. territory. An additional 86 taxa or 12% are restricted The taxa employed in this study included 67 to two states, provinces, or territories. Lastly, states, genera, 507 species and 2 subspecies occurring in the provinces, territories and federal districts are the USA, Canada, and Mexico (Appendix 1). most frequently cited geographic area units for North Incorporating both genera and species was done to America Plecoptera distribution (Stark et al. 1986; amplify the historical ‘signal’ present in the data set Stark et al. 2008; Stewart & Stark 2002; Stewart & (Porzecanski & Cracraft 2005; Vazquez‐Miranda et al. Oswood 2006). Nonetheless, ongoing efforts to 2007). Species or genera autapomorphic for a single compile a database of North American stonefly state or province were excluded as they were not distribution records at the county level (Kondratieff informative with respect to grouping the taxa of this & Baumann 2000) should be able to provide more study (Bryant 1995). CADE also provides for the detailed information for future analyses. incorporation of the distribution of subspecies. While A terminal area consisting of all 0’s (absences) this latter category is not widely employed in was also included and is used to root the tree Plecoptera, the one instance in North American (Cracraft 1991; Porcecanski & Cracraft 2005). The Plecoptera, Cultus decisus decisus and C. decisus terminal area is, thus, a hypothetical outgroup and isolatus, is included. The main sources of information coding it in this manner assumes that the absence of for the distribution of Plecoptera taxa in Canada, the taxa is the plesiomorphic condition and that the USA, and Mexico were Stark & Kondratieff (2004), sharing of taxa between two or more areas is the Stewart & Oswood (2006) and Stark et al. (2008).

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Presence or absence of a taxon in a political area unit bootstrap support at the 50 % confidence level. was coded as 1 or 0 respectively. Basal clade 2 is characterized by the distribution Parsimony analysis of the data matrix was of the genus Capnia (Appendix 2). Within this clade undertaken using PAUP* (version 4.0b10, Swofford subordinate clade 3 comprised of the 20 states, 2002). The heuristic search option was used in lieu of provinces, and territories of the western USA and exact approaches such as the branch‐and‐bound and Canada appears as the sister group of subordinate exhaustive searches. The latter are computationally clade 22 comprised of the 3 states of northwestern intractable with large‐sized data sets. However, use Mexico. Although subordinate clade 3 does not of the former does not guarantee the identification of receive bootstrap support, 14 of its subordinate the minimum‐length area cladogram(s). The heuristic clades are supported (Fig. 3). Subordinate clade 22, search option was performed using 100 replicates, however, receives support as a monophyletic group 100 random addition sequence replicates, random (Fig. 3). Plecoptera species and genera whose step‐wise addition, tree‐bisection and reconnection distribution represents unambiguous changes for branch swapping, collapse of zero length branches, subordinate clades 3 and 22 and subordinate clades Multrees option in effect, deepest descent option not included within them (4 through 21 and 23 in effect. Clade support was assessed using bootstrap respectively) are listed in Appendix 2. In subordinate frequencies (Felsenstein 1985). Bootstrap branch clade 3 five subordinate clades exhibit a pattern values were determined in PAUP* from using the consisting of a pair of geographically adjacent heuristic search option above. Strict Consensus and political area units: 16 (Oregon, Washington), 17 Majority Rule Consensus area cladograms were (Idaho, Montana), 18 (Yukon, Alaska), 19 (Colorado, constructed when more than one equally New Mexico), and 21 (Saskatchewan, Manitoba). In parsimonious cladogram resulted. MacClade 4.0 subordinate clade 22 the two states comprising (Maddison & Maddison 2000) was used to trace subordinate clade 23 (Sonora, Chihuahua) are character transformations for specific clades on the geographically adjacent political area units. resulting Majority Rule Consensus area cladogram. Basal clade 24 is characterized by the distribution of the genus Allocapnia (Appendix 2) and includes 43 RESULTS states, provinces, and territories of the eastern USA The heuristic approach of PAUP* identified 146 and Canada as well as the federal district of the USA. area cladograms with a length of 2267 steps. These Plecoptera species and genera whose distributions area cladograms have a consistency index (CI) of represent unambiguous changes for the subordinate 0.2532, a retention index (RI) of 0.6704, a rescaled clades within basal clade 24 (25 through 66) are listed consistency index (RC) of 0.1698, and a homoplasy in Appendix 2. This clade is largely grouped into index (HI) of 0.7468. The Strict Consensus area three major subordinate clades 28, 36 and 63 (Fig. 4). cladogram (Fig. 3) exhibits three basal clades of Major subordinate clade 28 includes the states of the states, provinces and territories but political entities southeastern USA and most of its subordinate clades from Mexico are largely unresolved. The Majority (30 through 35) receive bootstrap support (Fig. 3). In Rule Consensus area cladogram (Fig. 4) shows clades clade 28 the six states comprising subordinate clade found in 50% of the 146 most parsimonious trees. It 31 (Mississippi, Alabama, Georgia, South Carolina, identifies the same three basal clades found in the North Carolina, Tennessee) exhibit a nesting of Strict Consensus area cladogram, but exhibits greater geographically adjacent political area units that resolution with respect to the areas from Mexico. extend in an arc from west to east. Major subordinate The more highly resolved Majority Rule clade 36 includes the central and northeastern Consensus area cladogram is the basis of the political area units of the USA and Canada. following discussion of relationships between the However, it does not receive bootstrap support. political area units comprising Canada, the USA, and Subordinate clade 36 largely comprised two Mexico. The three basal clusters of the Majority Rule subordinate clades 40 and 47 (Fig. 4). The former Consensus area cladogram are identified as clades 2, includes states from the central USA and it and its 24, and 67 (Fig. 4). These clades, however, lack subordinate clades (41 through 46) receive bootstrap

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Fig. 3. Strict Consensus area cladogram of the 146 most parsimonious area cladograms of states, provinces, territories, and federal districts of Canada, USA, and Mexico based on Plecoptera distributional data. Numbers above the branch indicate bootstrap values above 50%, otherwise less than 50%.

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Fig. 4. Majority Rule Consensus area cladogram of the 146 most parsimonious area cladograms of states, provinces, territories, and federal districts of Canada, USA, and Mexico based on Plecoptera distributional data. Numbers identify clades discussed in text. Percentage of a particular clade appearing among the 146 most parsimonious area cladograms indicated within parentheses, otherwise 100%.

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support (Fig. 3). States that comprise two of the adjacent political area units. subordinate clades of clade 40, 42 (Illinois, Indiana, Kentucky, Ohio) and 45 (Missouri, Arkansas, DISCUSSION Oklahoma), each exhibit a nesting of geographically The three major area clades of the consensus area adjacent political area units. Subordinate clade 47 cladograms agree with the previous observations of includes political area units in central and Atlantic Hynes (1988) and Stewart & Stark (1988, 2002) Canada and the northeast, upper south Atlantic and concerning the grouping of North American stonefly upper Midwest USA. This clade did not receive fauna into western boreal (clade 2), eastern boreal bootstrap support and only two of its subordinate (clade 24) and austral (clade 67) areas. Within clade 2, clades, 57 and 59, are supported. The latter clades are the relationship between subordinate clades 22 (the included within subordinate clade 56. The five states three states of northwestern Mexico) and 3 (the comprising this clade (New York, Pennsylvania, political units of the western USA and Canada) Maryland, Virginia, West Virginia) exhibit a nesting agrees with the study of Sargent et al. (1991) of geographically adjacent political area units that regarding the zoogeographic affinities of extends linearly from north to south. Two additional northwestern Mexico. The pattern of grouping in clades within subordinate clade 47, 60 (Connecticut, clade 2 agrees with the Rocky Mountains and Sierra‐ Massachusetts) and 61 (New Brunswick, Nova Cascade‐Coast Ranges centers of endemism noted by Scotia), each consist of a pair of geographically Stewart & Stark (2002). Clade 7 and its subordinate adjacent political area units. Major subordinate clade clades are comprised of political area units that 63 of basal clade 24 includes two provinces from include sections of one or both of these mountain Atlantic Canada (Prince Edward Island, systems or are hypothesized to have faunal affinities Newfoundland and Labrador) and two with these systems (Stewart & Oswood 2006). geographically adjacent New England states (New A single Appalachian‐Ozark center of endemism Hampshire, Vermont). This clade and its subordinate reported by Stewart & Stark (2002) may correspond clades did not receive support as monophyletic to clade 24. However, political area units containing groupings. sections of one or both of these mountain systems are Basal clade 67 is characterized by the distribution distributed among subordinate clades in clade 24 that of the genus Anacroneuria (Appendix 2) and is also include political area units lacking sections of comprised of 17 states and the federal district of these systems. Major subordinate clades in clade 24 Mexico (Fig. 4). Plecoptera species and genera whose suggest a relationship between states located in the distributions represent unambiguous changes for the southeastern USA (clade 28, Fig. 4), states located in subordinate clades within 67 (68 through 81) are the central USA (clade 40, Fig. 4), and states and listed in Appendix 2. The two subordinate sister provinces located in central and Atlantic Canada as clades, 68 and 75, that originate within clade 67 are well as northeast, upper south Atlantic and upper not present on the Strict Consensus area cladogram Midwest USA (clades 47 and 63, Fig. 4). but are present on 81% and 70% of the minimum‐ The political area units comprising subordinate length trees respectively. Subordinate clade 68 clades 68 and 75 of clade 67 indicate a historical includes the states of eastern and southern Mexico. relationship between the states from the eastern and States comprising one of its subordinate clades, 71 southern areas of Mexico and a historical relationship (Guerrero, Oaxaca, Vera Cruz, and Chiapas), show a between the states of the west‐central area of Mexico nesting of geographically adjacent political area units respectively. that extends north to south. Another subordinate The three basal clusters of clades found in the clade, 74, includes two states (Nuevo Leon, consensus area cladograms are compatible with Tamaulipas) that are geographically adjacent political vicariance events associated with plate tectonics. The area units. Subordinate clade 75 includes the political split of the boreal area into the western and eastern units from west‐central Mexico. In this clade the state regions of the USA and Canada (Fig. 4, clades 2 and and federal district (Mexico, Distrito Federal) 24) is in agreement with either the late Cretaceous comprising subordinate clade 81 are geographically mid‐continental seaway that divided the Nearctic

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(Allen 1983, Nelson 1988) or the later Miocene examination of fossil stoneflies, Sinitshenkova (1997) orogenies of the Rockies and Sierra Nevada in the reported that separate northern and southern west along with the isostatic uplift of the hemisphere groupings of the order actually date Appalachians in the east or both. from the late Permian. Although this earlier date It is not evident what intercontinental vicariant would make the late Jurassic vicariant event events are responsible for subsequent branching compatible with the separation of an ancestral within clades 2 and 24. In general, the relatively Anacroneuria lineage from a sister Nearctic lineage, recent Pleistocene geological events (Ross & Ricker the incomplete fossil data and the lack of information 1971, Hynes 1988, Sargent et al. 1991, Stewart & Stark on phylogenetic relationships does not allow for any 1988, 2002) are thought to have been important. definitive conclusion. Pleistocene ice at its maximum covered Canada and The low CI, RI, and RC values exhibited by the parts of the northern USA and is considered to have 146 minimum‐length area cladograms as well as the forced Plecoptera into refugia located in the lack of bootstrap support of many of the clades (Fig. southeastern USA, western USA, and Alaska. As the 3) are indicative of numerous reversals and ice subsided taxa dispersed from these refugia. parallelisms (Schuh 2000). Biogeographically, these However, the great age of the primary divisions of have has been interpreted to represent the influence the North American Plecoptera implies that the later of extinction and random long‐distance dispersal diversification within clades 2 and 24 had an earlier respectively (Craw et al. 1999) and suggests that history than the Pleistocene. The Miocene uplift of these two phenomena may have impacted the the Rockies and Sierra Nevada along with the distribution of North American Plecoptera genera secondary uplift of the Appalachians by fragmenting and species. Extinction of taxa, as reflected by formerly continuous habitats and creating new ones shared‐absences of genera and species, was exhibited could, through parapatric and allopatric speciation, by 27 of the 69 clades (Appendix 2) of the Majority be responsible for the hierarchical patterns seen Rule Consensus area cladogram (Fig. 4). These within these subordinate clades. ranged from 3.8‐100% of the unambiguous taxon Clade 67, the austral clade, is characterized by the changes for each of these clades with a mean of Perlidae genus Anacroneuria that invaded this area 34.9%. Eight clades, 11, 15, 18, 53, 58, 60, 61 and 81, from the Neotropic during the Pliocene (Stark & are characterized by having shared‐absences Gaufin 1976, Hynes 1988, Stewart & Stark 1988, 2002) constitute a majority of the observed unambiguous when North America and South America were re‐ changes (Appendix 2). The extent of how greatly united by the newly re‐established Isthmus of extinction and random long‐distance dispersal might Panama. Anacroneuria, however, originated much have distorted any historical signal embedded in the earlier than this date. Stark & Gaufin (1976) distribution data of Plecoptera is difficult to assess at hypothesize that starting in the early Cretaceous, this time. As distribution data in the same political prior to the breakup of Gondwanaland, ancestral area units also exist for Ephemeroptera (Randolph, progenitors of this genus dispersed from Europe or 2002) and Odonata (Mauffray & Beckenbach 2005a, from Asia via Africa to South America. Subsequently 2005b), two aquatic groups with origins this group became extinct in Africa. phylogenetically older than Plecoptera, one approach Another possibility is faunal interchange between would be to undertake a PAE/CADE analysis of the North and South America prior to the break‐up of distribution of these groups similar to that done for Pangaea in the late Jurassic. This vicarance event has stoneflies. Coincident hierarchical patterns exhibited instead been taken by Zwick (2000) to correspond to by these groups with that resulting from this analysis the fragmentation of the order itself into two would be indicative of common underlying historical suborders ‐ one representing a northern faunal causes while their absence would imply the assemblage and the other a southern faunal importance of extinction and random long‐distance assemblage. This would imply a later vicariant dispersal. However, unlike Plecoptera, which is separation was responsible for the origin of the generally restricted to lotic habitats, Ephemeroptera Neotropic Anacroneuria. However, based on an and Odonata occupy both lotic and lentic habitats

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and this may increase the influence of extinction and units. random long‐distance dispersal on the distribution patterns of their taxa. ACKNOWLEDGMENTS The high level of homoplasy on the minimum‐ I thank C. Riley Nelson and R. Edward DeWalt length area cladograms may also reflect both the for insightful observations and useful suggestions large number of area units and their artificial nature. concerning the manuscript. I also thank Mark Schorr Morrone & Escalante (2002) employing PAE analysis and Joey Shaw for making relevant comments on an involving the distribution of Mexican terrestrial earlier version of the manuscript. animals found that the number of steps decreased and the number of synapomorphies increased on REFERENCES their resulting area cladograms when the total Allen, R.T. 1983. Distribution patterns among number of the geographic area units decreased while arthropods of the north temperate deciduous simultaneously the area of the individual units forest biota. Annals Missouri Botanical Garden, increased as well as became more natural. 70:616‐628. PAE/CADE studies on North American Plecoptera Brooks, D.R. & M.P.G. van Veller. 2003. Critique of distributional data associated with ecoregions or parsimony analysis of endemicity as a method of biogeographic provinces would be desirable, but historical biogeography. Journal of Biogeography, accurate distributional data based on these area units 30:819‐825. are not available in a published database. Such Bryant, H.N. 1995. Why autapomorphies should be information is crucial, however, and will have to be removed: A reply to Yeates. Cladistics, 11(4):381‐ gathered by digitization of specimen data presently 384. housed in a number of museum, university and Cracraft, J. 1991. Patterns of diversification within personal collections before more finely grained continental biotas: Hierarchical congruence PAE/CADE analyses can be expected. among areas of endemism of Australian It is also possible, despite the efforts of many vertebrates. Australian Systematic Botany, 4:211‐ workers, that the low level of shared taxa for many 227. clades of the minimum‐length area cladograms may Craw, R.C., J.R. Grehan, & M.J. Heads. 1999. represent an artifact of incomplete information on the Panbiogeography: tracking the history of life. distribution of North American stonefly genera and Oxford Biogeography Series No. 11, Oxford species. Accumulating essential additional stonefly University Press, New York, NY. 229 pp. distributional data from stream ecosystems in Crisci, J.V., L. Katinas, & P. Posadas. 2003. Historical Canada, the USA and Mexico would address this biogeography: an introduction. Harvard concern as well as possibly clarify unresolved area University Press, Cambridge, London, i–vii, 1‐250. relationships depicted on the Majority Rule Felsenstein, J. 1985. Confidence limits on Consensus area cladogram of this study (Fig. 4). phylogenies: an approach using the bootstrap. Ultimately corroboration of this PAE/CADE Evolution, 39:783‐791. analysis will need to be determined by using as yet Hynes, H.B.N. 1988. Biogeography and origins of the unpublished resolved phylogenies of species North American stoneflies (Plecoptera). Memoirs comprising North American stonefly genera. Several of the Entomological Society of Canada, 144:31‐37. genera each with over thirty described species could Kondratieff, Boris C. & Richard W. Baumann provide a basis for generating useful biogeographical (coordinators). 2000. Stoneflies of the USA. hypotheses ‐ Allocapnia, Capnia, Alloperla, Sweltsa, and Jamestown, ND: Northern Prairie Wildlife Isoperla. Congruence between results obtained from Research Center Online. these analyses with that provided by this study http://www.npwrc.usgs.gov/resource/distr/insects would be a further indication that the latter correctly /sfly/index.htm (Version 12DEC2003). captured the historical signal in existing stonefly Luna‐Vega, I., O. Alcantra Ayala, D. Espinosa distributional data concerning hierarchical Organista, & J.J. Morrone. 1999. Historical relationships of North American large political area relationships of the Mexican cloud forests: a

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preliminary vicariance model applying Biogeographic Review of North American Mayflies parsimony analysis of endemicity to vascular (Ephemeroptera). PhD Dissertation, Department of plant taxa. Journal of Biogeography, 26:1299‐1305. Entomology, Purdue University. URL address Maddison, D.R. & W.P. Maddison. 2000. MacClade http://www1.usu.edu/buglab/projects/Mayflylist.pdf 4.0. Analysis of phylogeny and character Rosen, B.R. 1988. From fossils to earth history: evolution, version 4.0. Sinauer Associates, applied historical biogeography. In Analytical Sunderland, MA, USA. Biogeography: An Integrated Approach to the Study of Mauffray, W. & L. Beckenbach. 2005a. Distribution Animal and Plant Distributions (A. A. Myers and P. Summary of North American Anisoptera. S. Giller, eds.). pp.437‐481. New York: Chapman Compiled by George H. Bick and Bill Mauffray. & Hall. Last update 17 June 2005. URL address Ross, H.H. & W.E. Ricker. 1971. The classification, http://www.iodonata.net/ evolution, and dispersal of the winter stonefly Mauffray, W. & L. Beckenbach. 2005b. Distribution genus Allocapnia. Illinois Biological Monographs, Summary of North American Anisoptera. University of Illinois Press, Urbana, Chicago, and Compiled originally from the “The Damselflies of London 45:1‐166. North America” by Minter J. Westfall, Jr. and Sargent, B.J., R.W. Baumann, & B.C. Kondratieff. Michael L. May (1996) with permission of 1991. Zoogeographic affinities of the Nearctic Scientific Publishers Inc., Gainesville, FL. Last stonefly (Plecoptera) fauna of Mexico. The update 25 October 2005. URL address Southwest Naturalist, 36 (3):323‐331. http://www.iodonata.net/ Schuh, R.T. 2000. Biological systematics: principles Morrone, J.J. 2005. Cladistic biogeography: identity and applications. Cornell University Press, Ithaca and place. Journal of Biogeography, 32:1281‐1286. and London, i‐ix, 1‐236. Morrone, J.J. & T. Escalante. 2002. Parsimony Sinitshenkova, N.D. 1997. Paleontology of stoneflies. analysis of endemicity (PAE) of Mexican In Landolt, P. and M. Sartori (eds.) terrestrial mammals at different area units: when Ephemeroptera & Plecoptera Biology‐Ecology‐ size matters. Journal of Biogeography, 29:1095‐ Systematics, Fribourg: Mauron + Tinguely & 1104. Lachat SA, 561‐565. Nelson, C.H. 1988. Note on the phylogenetic Stark, B.P. & A.R. Gaufin. 1976. The Nearctic genera systematics of the family Pteronarcyidae of Perlidae (Plecoptera). Miscellaneous (Plecoptera), with a description of the eggs and Publications of the Entomological Society of nymphs of the Asian species. Annals of the America, 10:1‐80. Entomological Society of America, 81:560‐576. Stark, B.P. & B.C. Kondratieff. 2004. Anacroneuria Nelson, C.R. 2004. Systematics of the Capnia from Mexico and upper Mesoamerica (Plecoptera: californica species group, including a Perlidae). Monographs of the Western North morphological phylogeny, zoogeography, and American Naturalist, 2:1‐64. description of Capnia kersti, new species Stark, B.P. & C.R. Nelson. 1994. Systematics, (Plecoptera: Capniidae). Annals of the phylogeny and zoogeography of genus Yoroperla Entomological Society of America, 97 (1):97‐104. (Plecoptera: Peltoperlidae). Entomolgica Nihei, S.S. 2006. Misconceptions about parsimony of Scandinavica, 25:241‐243. endemicity. Journal of Biogeography, 33:2099‐ Stark, B.P., S.W. Szczytko,& R.W. Baumann. 1986. 2106. North American stoneflies (Plecoptera): Porzecanski, A.L. & J. Cracraft. 2005. Cladistic systematics, distribution and taxonomic analysis of distributions and endemism (CADE): references. The Great Basin Naturalist, 46:383‐397. using raw distributions of birds to unravel the Stark, B.P., R.W. Baumann & R.E. DeWalt. 2008. biogeography of South American arid lands. Valid stonefly names for North America: Updated Journal of Biogeography, 32:261‐265. as of 28 January 2008. Plecoptera Society of North Randolph, R.P. 2002. Distribution of mayfly species America. URL address in North America. List compiled from Atlas and http://plsa.inhs.uiuc.edu/plecoptera/validnames.aspx

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Stewart, K.W. & B.P. Stark. 1988. Nymphs of the North American stonefly genera (Plecoptera). Entomological Society of America, Thomas Say Foundation, Volume 12:i‐xiii, 1‐460. Stewart, K.W. & B.P. Stark. 2002. Nymphs of the North American stonefly genera (Plecoptera), second edition. The Caddis Press, i‐xi, 1‐510. Stewart, K.W. & M.W. Oswood. 2006. The stoneflies (Plecoptera) of Alaska and Western Canada. The Caddis Press, i‐vi, 1‐325. Swofford, D.L. 2002. PAUP* Phylogenetic analysis using parsimony (and other methods). Version 4.0b10 for Apple Macintosh. Sinauer Associates, Sunderland, MA, USA. Terry, M.D. & M.F. Whiting. In Preparation. Phylogenetic systematics of Plecoptera: evidence from morphology and six genes. Vazquez‐Miranda, H., A.G. Navarro‐Siguenza, & J.J. Morrone. 2007. Biogeographical patterns of the avifaunas of the Caribbean Basin: a parsimony perspective. Cladistics, 23:180‐200. Zwick, P. 2000. Phylogenetic system and zoogeography of the Plecoptera. Annual Review of Entomology, 45:709‐746.

Received 3 April, Accepted 29 April, Published 18 December 2008

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Appendix 1 Data matrix for the 574 Plecoptera taxa and genera employed as characters for 85 Canadian, United States and Mexican provinces. Presence of taxa indicated by one, absence by zero. An outgroup (OG) consisting of all zeros is included in the matrix. Area acronyms as noted for Figures 1 and 2.

AL AB 10000111000000100000000000000100000001000000000000000000000100000000000000000000000000 00001010000000000000000000000000000001000000000000000000000000000100000000000000000000 00001000000000000000000000000000000001000000000000000000000000000000000000000000000000 10000100000000000000000000000100000000000000000000000000000000000000000000000000000000 00000100000000000000000000000000000000000000000000000000000100000000000000000000000000 10000101000000100000000000000100000001000000000000000000000000000000000000000000000000 01000000111000011010010100000011000010100011000101100010011011000000000000000000000000 00000000000000001010000000000000000000000011000000000000000001000000000000000000000000 00000000000000000000000000000010000000000010000101000000000000000000000000000000000000 00010000000000000000000000000000000000000000100000000000000000000000000000000000000000 01000000000000000000000000001000000000000000000000000000000000000000000000000000000000 00000000110000000010010111110010100100100011010101100010011011000000000000000000000000 01000000111010011011010110000001000100100011010101010010010011000000000000000000000000 01010000011011010110000100111001000010000011110100000010010011000000000000000000000000 00010000000000000000000000010000000000000000100000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000100000000010000000000000000000000000000 01000000000000011010000000000000000000000001000000000000000000000000000000000000000000 00000000000000000000000000000001000000000000000000000010010000000000000000000000000000 00000000000000000001000000001000000000000000000000000000000000000000000000000000000000 01000000000000000000000000000000000000000000000000000010000000000000000000000000000000 01000000001010000001000000001001000000000000000000010000010000000000000000000000000000 00010000010001011110000101110010010000000011110101000010010011000000000000000000000000 00010000000000000000000000000000000000000000000000000000000000000000000000000000000000 00000000011010000000000100000001000000000000000100010010010000000000000000000000000000 00000000100000000010000110000010000000000011000100000010010001000000000000000000000000 11111111100110111111111111111111110101110111001111011110110111110000100000000000000000 01000000010010000000000100001001000000000000000100010010010000000000000000000000000000 00000000000000000010000000000000000000000000000000000010000000000000000000000000000000 00000000000000000010000100000000000000000010000100000000010001000000000000000000000000 00000000000000011010000000000000000000000001000000000010000001000000000000000000000000 00000000000000000010000100000000000000000011000100000010010011000000000000000000000000 00000000000000000000000100000000000000000000000000000000000001000000000000000000000000 00000000000000000000000000000001000000000000000000000010010000000000000000000000000000 01010000010001011111001101111000000010000011110101000011010011000000000000000000000000 00000000000010000000000000000001000000000000000000010010010000000000000000000000000000 10101101000000100000000000000100000001010100001000000100100000110000000000000000000000 10100010000000000000000000000000000000000100001000000000000100000000000000000000000000 00000000000000000010001110000001000100000010000101010000010001000000000000000000000000 00000000000000000000000100000000000000000000000100000000010001000000000000000000000000 00000000000000011000000110100000000000000011010001000000010011000000000000000000000000 00000000000000001010000000000000000000000011000000000000000000000000000000000000000000 00010000000000000000000000010000000000000000010000000000000000000000000000000000000000 00000000000000000000000000000000000000000000000000000010010001000000000000000000000000 00010000000000000001000000000000000000000000100000000001000000000000000000000000000000 00000000100000000000010110000010000100100010000101000000011001000000000000000000000000 00000000100000000000010011100010001100100010010001000000001010000000000000000000000000 00010000000000000000000000010000000000000000100000000000000000000000000000000000000000 01010000000010011010000000011000000000000001000000000010010001000000000000000000000000 10101111000000100000000000000100000001010100001000000100100100110000000000000000000000 01000000100010000010010110000011000100100010000111010000010001000000000000000000000000 00101111000000100000000000000100000001000000001000000000100100110000100000000000000000 00000000000000000010000010000010000000000010000100000000010001000000000000000000000000 11100101000100001111101111101111000000010011000101010110010011110000000000000000000000 00010000100000011110001001110000110000000001000100001010000010000000000000000000000000 00000110000000100000000000000000000000000100001000000000100100000000000000000000000000

badia regularis spenceri sasquatchi milami

scotti proteus biloba californica comstocki dorsata pictetii princeps

aurora cunninghami curiosa forbesi frisoni frumi fumosa granulata harperi illinoensis indianae jeanae loshada malverna maria minima mohri mystica nivicola ohioensis pechumani peltoides polemistis pygmaea recta rickeri sandersoni simmonsi smithi stannardi starki tennessa virginiana vivipara wrayi zola

californica caryi Bolshecapnia Pteronarcys Allocapnia Pteronarcella

Capnia Capnia Genus Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Bolshecapnia Genus Bolshecapnia Pteronarcys Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Taxa/Area Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Allocapnia Genus Bolshecapnia Pteronarcella Pteronarcella Genus Pteronarcys Pteronarcys Pteronarcys Pteronarcys Pteronarcys Pteronarcys Pteronarcys 1 2 3 4 5 6 7 8 9 54 55 53 13 14 15 16 17 18 19 20 52 12 51 11 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 10

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Appendix 1 Continued

AL AB 10000100000000000000000000000100000000000000000000000000000000010000000000000000000000 00100101000000100000000000000100000001000000000000000100100000110000000000000000000000 10100111000000100000001000000100000001010100000000001100100100110000000000000000000000 10000001000000000000000000000000000001000000000000000000000000001100000000000001000000 00100110000000000000000000000000000000000000001000000000000100010000000000000000000000 00100110000000000000000000000000000000000000000000000000000100000000000000000000000000 10100111000000100000000000000100000001010000001000001100100100110000000000000000000000 10101111000000100000000000000100000001000000001001000001100100011100000000000001000000 10101111000000100000000000000100000001000100001000001000100100110000000000000000000000 00100110000000000000000000000000000000000000001000000000000100000000000000000000000000 10100100000000000000000000000000000000000000001000000000000100000000000000000000000000 10100001000000100000000000000100000001000000000000000100100000110000000000000000000000 10100110000000100000000000000100000000010000001000000000100100100000000000000000000000 10000001000000100000000000000100000000000000001000000000100000000000000000000000000000 10100000000000100000000000000100000000000000000000001000000000110000000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 00000000000000100000000000000000000000000000001000000000000100000000000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 00000010000000000000000000000000000000000100000000000000100000000000000000000000000000 10000101000000100000101001100110000001011001010001000100100010100000000000000000000000 10101111000000100000101000100110000001011101011001001100100111111100000000000001000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00001001000000000000000000000000000001000000000000000000000000000100000000000000000000 00000000000000100000000000000000000000000100001000000000100000000000000000000000000000 00000000100000000000011011000010000000000000010001000000001010000000000000000000000000 00000000000000100000000000000100000000000000001000000000000100000000000000000000000000 00000011000000000000000000000000010001000100000000000000100000100000000000000000000000 00001011100000100000011011000110010001000100011001000000101110100100000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 10100111000000100000001000000100000001000100001000001100100100111000000000000000000000 00000010000000000000000000000000000000000100000000000000000000000000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00000010000000100000000000000000000000000000000000000000000000000000000000000000000000 00100110000000000000000000000000000000000000001000000000000100000000000000000000000000 10100101000000100000000000000100000000000000001000000000100100010000000000000000000000 00100100000000000000001000000000000000011000010000000000000000010000000000000000000000 10100100000000100000000000000100000000000000001000000000100000110000000000000000000000 00000110000000000000000000000000000000000000000000000000000000010000000000000000000000 00000110000000000000000000000000000000000000001000000000000100000000000000000000000000 10000100000000100000000000000100000000000000001000000000000100000000000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 00000001000000100000000000000000000001000100000000000000100001000000000000000000000000 00001000000000000000000000000000000001000000000000000000000000000000000000000001000000 00100100000000000000000000000000000000000000001000000000000100000000000000000000000000 00000011000000000000000000000000000001000000000000000001100000001100000000000001000000 00000010000000100000000000000000000000000000001000000000000000001000000000000000000000 10100100000000000000000000000100000000000000001000000000000100010000000000000000000000 00000010000000000000000000000000000000000000001000000000000100000000000000000000000000 00100110000000000000000000000000000000000000001000000000000100000000000000000000000000 00100100000000000000000000000000000000000000000000000000000000010000000000000000000000 00001010000000000000000000000000000001000000000000000000000000000100000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00100110000000000000000000000000000000000000001000000000000100000000000000000000000000 10100111000000100000000000000100000001000000000000000000100100010000000000000000000000

brevicauda

arizonensis autumna frisoni lapwae oenone porrecta projecta variabilis werneri yoloensis

abbreviata agassizi crinita grandis hyalita integra mogila palousa rickeri spenceri vedderensis elevata fibula intermontana manitoba venosa wanica

umpqua utahensis vernalis cheama coloradensis confusa decepta elongata excavata glabra gracilaria lacustra licina lineata melia nana nearctica petila pileata promota sextuberculata tumida uintahi Capnia Capnura Mesocapnia Isocapnia

Isocapnia Isocapnia Isocapnia Isocapnia Isocapnia Isocapnia Isocapnia Isocapnia Taxa/Area Capnia Capnia Capnia Genus Capnura Capnura Capnura Capnura Capnura Capnura Genus Eucapnopsis Mesocapnia Mesocapnia Mesocapnia Mesocapnia Mesocapnia Mesocapnia Mesocapnia Mesocapnia Mesocapnia Genus Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Capnia Isocapnia Capnia Capnia Capnia Capnia Capnia Capnia Mesocapnia Isocapnia Isocapnia Genus 88 89 90 91 92 93 94 95 77 78 79 80 81 82 83 84 85 86 87 56 63 64 65 66 67 68 69 70 71 72 73 74 75 76 96 57 58 59 60 61 62 97 98 99 101 102 103 104 105 106 107 108 109 110 100

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Appendix 1 Continued

AL AB 00000010000000000000000000000000000000000100000000000000000000000000000000000000000000 10100111000000100000100000000100000001010100001001001000100100110000000000000000000000 00100110000000100000000000000100000000000000001000000000000100000000000000000000000000 00000000000000000010000100000001000000000000000100010010010001000000000000000000000000 01000000000000011010000000000000000000000000000000000000000000000000000000000000000000 01000000000010000000000000000001000000000000010000000010010000000000000000000000000000 00000001000000000000000000000100000001000000000000000000100000100000000000000000000000 00000000000000100000000000000000000000000000001000000000000000000000000000000000000000 00000001000000000000000000000000000001000000000000000000100000100000000000000000000000 00000001000000100000000000000000000000000100000000001000100000100000000000000000000000 00000000000000000000100000000000000000000000000001000000000000000000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 10000000000000100000000000000100000000000000000000000000000000100000000000000000000000 10100110000000000000000000000100000000010000001000000000000000010000000000000000000000 00000000100000000000100010100010001010000010010011000000000010000000000000000000000000 00010011101000010010111111110011001110000011111111001110010111100000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00010001101000010010111111010001001100000011100101001110010011100000000000000000000000 10100111000000100000000000000100000001000000001000000000100100100000000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 10000100000000100000000000000100000000000000001000000000000000110000000000000000000000 10000110000000100000000000000100000000000000001000000000000000110000000000000000000000 00010000000000000000000000000000000000000000100000000000000000000000000000000000000000 00000000000000000000000100000001000000000000000100000010010000000000000000000000000000 01000000000100000000000000001000000000000000000000000000000000000000000000000000000000 00000000100000000000000110000010000000100010010111000000010001000000000000000000000000 01000000101100010010010111101011001000100011000101010100010011000000000000000000000000 00000000000000000000010110000011000000000000000100000000000001000000000000000000000000 00000000100000000000000010000000000100000000010101000000001001000000000000000000000000 00000000000000000000000000000001000000000000000000000000010000000000000000000000000000 01000000000010000000000000000000000000000000000000010000000000000000000000000000000000 00000000000000000000000000000000000000000000000000000010010000000000000000000000000000 00000000000000000000000000000001000000000000000000000010000000000000000000000000000000 01000000000001011010000100001000000000000001000000000000010001000000000000000000000000 00000000101000011010010110000011000000100011010111010010010011000000000000000000000000 00000000100000000000110110101010000010100010010101000000010011000000000000000000000000 01010000101001011010010110111010000010100011110101010010010011000000000000000000000000 00000000000100000000000000000001000000000010000001010000010000000000000000000000000000 00000000100000000000000010000010001000100010000101000000010001000000000000000000000000 00000000000000000000010110000000000100000000000100000000011000000000000000000000000000 01010000101111011011110111111011001110100011110111010110011011000000000000000000000000 00100110000000100000000000000000000000000000001000000000000100000000000000000000000000 10000111000000000000000000000100000001000000001000000000100100100000000000000000000000 01010000000100011000000000001001000000000000000001010000010000000000000000000000000000 00010000000000001101000000000000000000000000100000000001000000100000000000000000000000 00000000000000011010000000000000000000000001000000000011000000000000000000000000000000 00010000000000010000000000010000000000000000100000000011000000000000000000000000000000 00010000000000000000000000010000000000000000100000000000000000000000000000000000000000 10000110000000100000001100000101000000000000001100010010010101000000000000000000000000 01010000000000011110000000010000000000000001100000000011000001000000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000000000000000000100000000000000000000000100000000010001000000000000000000000000 00000000000000100000000000000000000000000000001000000000000100000000000000000000000000 10000100000000000000001000000100000000000000000000000000000100000000000000000000000000 00000000000000000000000000000001000000000000000000010010010000000000000000000000000000

complicata flinti kincaidi stigmata williamsae andersoni purcellana carolina

opis ensicala angulata fraxina narfi warreni cherokee claasseni

tahoensis poda nedia logana lemoniana labradora imbera distincta columbiana collaris utahensis

augusta

alexanderi alta biloba carolinensis cottaquilla duplicata ferruginea grandis maria mitchellensis moha monticola nephophila rickeri sibleyi tenella tenuis triloba truncata variabilis Utacapnia Paracapnia Pomoleuctra Zealeuctra Leuctra Perlomyia Megaleuctra

Genus Despaxia Leuctra Leuctra Leuctra Leuctra Utacapnia Utacapnia Utacapnia Utacapnia Utacapnia Utacapnia Utacapnia Utacapnia Genus Utacapnia Paracapnia Paracapnia Pomoleuctra Pomoleuctra Genus Zealeuctra Zealeuctra Zealeuctra Genus Zealeuctra Zealeuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Leuctra Genus Perlomyia Perlomyia Genus Paracapnia Nemocapnia Megaleuctra Megaleuctra Megaleuctra Megaleuctra Megaleuctra Genus Taxa/Area 125 126 127 128 129 130 124 123 122 121 120 119 118 117 115 116 114 113 151 152 153 156 157 158 159 154 155 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 112 111 160 161 162 163 164 165

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Appendix 1 Continued

AL AB 10000001000000100000000000000100000001000000001000000000100100100000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000000000100000000000000100000000000100001000000000100100000000000000000000000000 10001111000000100000001000000100000001000100001000001100100100101000000000000000000000 00000000000000000000000000000100000000000000000000000000000100000000000000000000000000 10100100000000000000001000000000000000010000000001001000000000110000000000000000000000 00100000000000000000000000000000000000010000000000000000000000000000000000000000000000 00100000000000000000001000000000000000000000000000000100000000010000000000000000000000 00000000000001010000101011000010000000100011010111001000000010100000000000000000000000 00000000100000000000010110000011000100100010010111000010011001000000000000000000000000 10100101000000100000000000000100000000000000001000000000100100110000000000000000000000 10100111000000100000000000000100000001000100001000000100100100110000000000000000000000 00000000000000000000100010000010001000100000000011000000000000000000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00100100000000000000000000000000000000000000000000000000000000010000000000000000000000 10100111000000100000100010000110001001100100001011000100100100110000000000000000000000 10100111000000100000000000000100000001000100001000001000100100110000000000000000000000 01010000001001011010010111111011000000100000110111010000010011000000000000000000000000 00000000000000000000000000000001000000000000000000000000010000000000000000000000000000 00000000101000001010010011110000000000000001000101010000010011000000000000000000000000 11110111101001111010010111111111000001100101111111011000110111110000000000000000000000 11110000000001010000001111101011000000010000010001010100010010000000000000000000000000 00000000001000000000000100000001000000000000000100000010010001000000000000000000000000 00000100000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000000000000000000100000001000000000000000000000000000000000000000000000000000000 00000000100000000000010111000000000100100010010101000000010001000000000000000000000000 00000000100000000000010100000010000000100010010101000000010001000000000000000000000000 00000100000000000000000000000000000000000000001000000000000100000000000000000000000000 00000100000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000001000000000000010000010000100000000000101000000010001000000000000000000000000 00000000100000001010010110000000000000000011010101000000010001000000000000000000000000 10100110000001010000101011000010000000110011010111001100000010110000000000000000000000 00000100101000001010010111000010000100100011011101000000010101000000000000000000000000 00000000000000000000000010000010000100100000000011000000000000000000000000000000000000 00000000100000000000010110000011000100100010010111000010011001000000000000000000000000 00000010000000000000000000000000000000000000001000000000000000000000000000000000000000 00000110000000000000000000000000000000000000001000000000000100000000000000000000000000 00001001000000000000000000000000000001000000000000001000100000100000000000000000000000 10000111000000100000001000000100000001000100001000000100100100100000000000000000000000 00000010000000000000000000000000000000000000000000000000000000001000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000101010000010010110000011000010100010000111010010010001000000000000000000000000 11111101101111111111111111011111000011110011110111011111110011110100100110101000001100 00001000000000000000000000000000000000000000000000000000000000000000000100000000001100 00000000000000011010000000000000000000000000000000000000000010000000000000000000000000 00000000000000000001000000000000000000000000000000000001000000000000000000000000000000 01010000101110011010100110011011000000100011010111010010010001000000000000000000000000 00001000000000000000000000000000000001000000000000000000100000000000000000000000000000 00000000000000000000000000000000000000000000000000000000000000000100000110000000001100 10100101000000000000101001000000000000010000010101001100000010010000000000000000000000 01010000000011011110000101011000000000000001110101000010010011000000000000000000000000 00001001000000100000000000000100000001000000000000001000100000100000000000000000000000 00000000000010000000000000000001000000000000000100010010010000000000000000000000000000 00001000000000000000000000000000000000000000000000000000000000000100100000000000000000 01000000000000000010000000000000000000000000000000000000000000000000000000000000000000

wui venusta varshava texana nigritta mogollonica mexicana linda delosa banksi appalachia apache alabama

claasseni perfecta

complexa dimicki foersteri prolongata truncata albidipennis

decepta delicatula macdunnoughi obscura weberi arctica normani rickeri trispinosa

carolinensis interrupta kondratieffi

depressa cornuta coloradensis californica biloba bifurcata flexura perplexa tina

besametsa completa hallasi similis

Amphinemura Paranemoura Malenka Nemoura Podmosta Prostoia Ostrocerca tumana rotunda

Malenka Malenka Malenka Malenka Malenka Genus Malenka Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Amphinemura Malenka Malenka Malenka Genus Lednia Nemoura Nemoura Nemoura Nemoura Genus Taxa/Area Podmosta Podmosta Podmosta Podmosta Podmosta Genus Prostoia Prostoia Prostoia Prostoia Genus Shipsa Soyedina Soyedina Soyedina Ostrocerca Ostrocerca Ostrocerca Ostrocerca Ostrocerca Genus Ostrocerca Paranemoura Paranemoura Genus 185 184 183 182 181 179 180 178 177 176 175 174 173 172 171 170 169 168 167 166 186 187 188 189 190 191 192 193 194 195 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 197 198 199 200 201 202 196 203 204 205

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Appendix 1 Continued

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appalachia arkansae cucullata fasciata limata

lita lonicera maura metequi nivalis parvula ugola burksi

glacialis contorta fosketti atlanticum jacobii jewetti kincaidii oregonense pacificum pallidum uinta umatilla

ovibovis

rossi

aracoma atlantica banksi biserrata caudata chloris concolor nevadensis potteri producta vallicularia washingtoni

occidentalis

chila cinctipes columbiana cordillera frigida haysi katahdin oregonensis Oemopteryx Soyedina Taeniopteryx Zapada Strophopteryx Taenionema

cataractae

Oemopteryx Genus Soyedina Soyedina Soyedina Soyedina Soyedina Genus Visoka Taeniopteryx Taeniopteryx Taeniopteryx Taeniopteryx Taeniopteryx Taeniopteryx Genus Taxa/Area Strophopteryx Zapada Zapada Zapada Zapada Zapada Zapada Zapada Zapada Genus Alaskaperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Strophopteryx Doddsia Oemopteryx Oemopteryx Bolotoperla Strophopteryx Strophopteryx Strophopteryx Genus Taeniopteryx Taenionema Taenionema Taenionema Taenionema Taenionema Taenionema Taenionema Taenionema Taenionema Genus Taeniopteryx 241 242 221 222 223 224 225 226 227 261 262 263 264 265 266 267 243 228 229 230 231 232 233 234 235 236 268 269 270 271 272 273 274 275 244 238 239 240 237 245 246 247 248 260 249 250 251 252 253 254 255 256 257 258 259

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Appendix 1 Continued

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diversa orpha brevis chilnualna

pastina delicata fraterna hamata idei imbecilla leonarda medveda nanina neglecta petasata pilosa serrata severa usa voinae vostoki

autumna dubia forcipata lineosa marginata pallidula starki wardi

terna

hondo lamba lateralis mediana naica occidens onkos oregonensis pacifica palearata pocahontas revelstoka townesi urticae voshelli adamantea albertensis borealis coloradensis exquista fidelis gaufini

Alloperla Haploperla Suwallia

Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Alloperla Haploperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Alloperla Genus Bisancora Haploperla Taxa/Area Genus Haploperla Plumaperla Rasvena Suwallia Suwallia Suwallia Suwallia Suwallia Suwallia Suwallia Suwallia Genus Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa Sweltsa 316 317 318 319 320 321 322 323 324 325 326 327 328 329 330 276 296 277 278 279 280 281 282 283 284 285 286 287 288 289 290 291 292 293 294 297 295 298 299 300 301 302 303 304 305 306 307 308 309 310 311 312 313 314 315

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Appendix 1 Continued

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perdita takhoma baumanni theodora evoluta filicis frisoni internata kirchneri lycorias ozarkensis perplexa petersi abnormis arenosa arida carolinensis covelli

ruralis cora

georgiana stewarti

californica ada

anna cornelia elisa laurie maria arcuata tarteri

brevis mariana nigrisoma siletz frontalis wilsoni

pintada signata thyra quadrispinula salish gaspesiana sopladora

Acroneuria Sweltsa Triznaka Soliperla Kathroperla Tallaperla Yoraperla Beloneuria Paraperla Utaperla Peltoperla

Acroneuria Acroneuria Acroneuria Acroneuria Acroneuria Acroneuria Acroneuria Acroneuria Genus Genus Soliperla Triznaka Triznaka Genus Genus Taxa/Area Attaneuria Kathroperla Kathroperla Genus Tallaperla Tallaperla Tallaperla Tallaperla Tallaperla Genus Veihoperla Yoraperla Yoraperla Yoraperla Yoraperla Genus Acroneuria Acroneuria Acroneuria Acroneuria Acroneuria Acroneuria Beloneuria Beloneuria Genus Paraperla Paraperla Genus Utaperla Utaperla Genus Calineuria Doroneuria Doroneuria Peltoperla Peltoperla Genus Sierraperla Soliperla Soliperla 370 371 372 373 374 375 376 377 378 331 350 332 333 334 351 379 335 336 337 352 353 354 355 356 357 358 359 360 361 362 363 364 365 366 367 368 369 380 381 382 338 339 340 341 342 343 383 384 385 344 345 346 347 348 349

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Appendix 1 Continued

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appalachia hokolesqua lineata litura negrolineata perplexa planicollis quadriloba ratcliffei aethiops annulicauda costana crenulata flavominuta hoogstrali

pacifica

xanthenes drymo ephyre zwicki

carlsoni catharae choctaw clymene coosa falayah gaufini harpi

nelsoni nitida placida puttmanni robeli shawnee shubuta taeysia xube adena baumanni bolukta browni cinctipes dakota decipiens frisoni fusca golconda lagoi napacola

Perlesta Anacroneuria Doroneuria Hansonoperla Perlinella Hesperoperla

Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Genus Anacroneuria Anacroneuria Anacroneuria Anacroneuria Anacroneuria Anacroneuria Genus Genus Eccoptura Hansonoperla Hansonoperla Genus Taxa/Area Perlinella Perlinella Perlinella Genus Anacroneuria Anacroneuria Anacroneuria Anacroneuria Anacroneuria Anacroneuria Anacroneuria Neoperla Neoperla Neoperla Neoperla Neoperla Neoperla Neoperla Neoperla Hesperoperla Genus Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta Perlesta 406 407 408 409 410 411 412 413 414 426 427 428 429 430 431 432 386 387 388 389 390 415 416 417 418 419 420 421 422 423 424 425 433 434 435 436 437 438 439 440 391 392 393 394 395 396 397 398 399 400 401 402 403 404 405

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Appendix 1 Continued

AL AB 00000000000000010000000010000000000000000001010000000000000000000000000000000000000000 01000000000000011010000010001000000000100011010110010010010010000000000000000000000000 00010000000001000000000000010000000000000000100000000000000000000000000000000000000000 00010000000000010000000100011000000000000000100100000010000010000000000000000000000000 01000000000000011010000110101000000000000001000100000010010011000000000000000000000000 01010000101111011010010101011011000000000001110100010010010011000000000000000000000000 00000000000010000000000000000001000000000000000000000000000000000000000000000000000000 00000010000000100000000000000000000000000000001000000000000100000000000000000000000000 01000001100001011110011011111011111000000011010101001100010011000000000000000000000000 00010000000000011010000100010001000000000001100000010000010001000000000000000000000000 01010000000111011011000110111001000000100011110100010011010011000000000000000000000000 01000000010100001001000100001000000000000000000100010000010001000000000000000000000000 00010000100001000010010111110010000100100011010100000000000011000000000000000000000000 01010000000010011010000100110001000000000011100100010010010011000000000000000000000000 01010000110111011011010111111011000100100011110100010010010011000000000000000000000000 10001111000000100000001000000100000001011000011001001100100100110000000000000000000000 01000000010110000001000000001001000000000000000000010001010000000000000000000000000000 00000000000000000000000000000001000000000000000000010010010000000000000000000000000000 00000000101010000010010110000011000100100010000101010010010001000000000000000000000000 01010000000110011101000000011001000000000000000000010000000000000000000000000000000000 00010000101001011010001111110010000100000011010101000100010011000000000000000000000000 01010000111111011111011111111011000100100011010101010111010011000000000000000000000000 00000010000000000000000000000000000000000000001000000000000100000000000000000000000000 00000110000000100000000000000100000000011000001000000000000100000000000000000000000000 00000000000000000000010011100011000000100000010101000010010011000000000000000000000000 10100101000001110100001000110100100001010000000001001100100000100000000000000000000000 00000001000001011110001011100010010110100000110101010100010011000000000000000000000000 00000010000000000000000000000000000000000100001000000000000100000000000000000000000000 00010000000000010101000000010000000000000000100000000000000000000000000000000000000000 00000000101000000000000110100000000100100010010101000000000000000000000000000000000000 00001111000000100000000000000100000001000000001000000000100100101000000000000000000000 00010000000000001010000110010001000000000001100100000000010001000000000000000000000000 00000000000000011010000001000000000000000001010101000000000010000000000000000000000000 00000000100100000010000010100011000100100011000101010010011001000000000000000000000000 00010000000000000000000000010000000000000000100000000000000000000000000000000000000000 10100101000000100000000000000100000000010000000000000100100100110000000000000000000000 00000001000000100000000000000000000001000000001000001000100000100000000000000000000000 10000111000000100000000000000100000000000100001000000000100100101000000000000000000000 10000111000000100100000000000100010001000100001000001100100000101000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000000000000000010011000010000000100000010001010000000011000000000000000000000000 00010000001000000001000000001000000000000000100000000001010000000000000000000000000000 01000000000000000000000000000000000000000000000000010000000000000000000000000000000000 00000000000000011010000000010000000000000001010000000000000000000000000000000000000000 00100100000000000000001000000000000000010000000010000100000000010000000000000000000000 01010000100101001010011111111011000000000000010101010010010011000000000000000000000000 00000000000000000000000000000001000000000000000000010010000000000000000000000000000000 00000000100000000000010010000000000000000010000101000000000000000000000000000000000000 00000000101000001000001011100011000000100000010111000000000010000000000000000000000000 10001111000000100000000000000100000001000100001000000000100100100000000000000000000000 10000100000000100000000000000100000000000000001000000000000100110000000000000000000000 00000000100000000000000100000000000000000010000001000000000001000000000000000000000000 00000000000000000000000000000000000000000000001000000000000100000000000000000000000000 00000000101010000010010110000011000000100010000101010010010001000000000000000000000000 00000001000000000000000000000000000001000000000000000001000000000000000000000000000000

trictura

fumosa ichusa immarginata kansensis media

sabulosa

luctuosa annulipes capitata flavescens

clio mainensis occipitalis osage robisoni stewarti

longiseta marlynia marmorata mohri montana mormona namata nana orata ouachita petersoni phalerata pinta quinquepunctata rainiera bellona bifurcata bilineata burksi cotta coushatta davisi decepta decolorata dicala distincta francesca frisoni fulva fusca gibbsae gravitans holochlora jewetti lata

Neoperla Agnetina Paragnetina

Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Neoperla Neoperla Neoperla Neoperla Genus Neoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Taxa/Area Agnetina Agnetina Agnetina Genus Claassenia Paragnetina Paragnetina Paragnetina Paragnetina Paragnetina Genus Calliperla Cascadoperla Clioperla 481 482 483 484 485 486 487 488 489 490 491 492 493 494 495 442 443 444 445 446 441 461 462 463 464 465 466 467 468 469 470 471 472 473 474 475 476 477 478 479 480 447 448 449 450 451 452 453 454 455 456 457 458 459 460

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Appendix 1 Continued

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decisus isolatus

compacta

misnomus

duplicata kanawholensis morgani robusta aurea expansa

bogaloosa nalatus subvarians irregularis signata subtruncata watertoni yosemite

yakimae

bilobatus kirchneri

nonus picticeps modestus tibialis bradleyi richardsoni roguensis signata similis slossonae sobria sordida tilasqua transmarina

sorpta

americana curvata Setvena Isoperla Kogotus Pictetiella Skwala Cultus Megarcys Remenus Diura Diploperla aestivalis decisus decisus pilatus tostonus verticalis

bicaudata knowltoni nanseni

Setvena Genus Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Isoperla Genus Isoperla Taxa/Area Genus Osobenus Pictetiella Genus Skwala Skwala Genus Cultus Cultus Cultus Cultus Cultus Cultus Genus Arcynopteryx Frisonia Megarcys Megarcys Megarcys Megarcys Megarcys Genus Remenus Remenus Genus Rickera Chernokrilus Diura Diura Diura Genus Diploperla Diploperla Diploperla Diploperla Genus Kogotus Kogotus Perlinodes Helopicus Helopicus Helopicus Setvena 516 517 497 498 499 500 501 502 503 504 505 496 535 536 537 538 518 519 520 521 522 523 524 525 526 527 506 507 508 509 510 511 512 513 539 540 541 542 543 544 545 546 547 528 529 530 531 532 533 534 514 548 549 550 515

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Appendix 1 Continued

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innubila crosbyi fugitans phormidia colubrinus doratus elongatus frontalis hansoni krumholzi olivaceus varians zionensis

hastatus iroquois

Helopicus Yugus Hydroperla Isogenoides Malerikus bulbosus kirchneri kondratieffi arinus

Genus Taxa/Area Yugus Yugus Yugus Genus Hydroperla Hydroperla Hydroperla Genus Isogenoides Isogenoides Isogenoides Isogenoides Isogenoides Isogenoides Isogenoides Isogenoides Isogenoides Genus Malirekus Malerikus Genus Oconoperla Yugus 551 571 572 573 574 552 553 554 555 556 557 558 559 560 561 562 563 564 565 566 567 568 569 570

Illiesia – http://www2.pms‐lj.si/illiesia/ Volume 4 – Number 18 – Page 198

Appendix 2 List of unambiguous species and genera changes for the clades noted in Figure 4. * indicates reversals. Clade Unambiguous species and genera changes

2 Genus Capnia

3 Claassenia sabulosa, Genus Isogenoides, I. colubrinus

4 Genus Pteronarcys, genus Pteronarcella, P. badia, genus Amphinemura, genus Suwallia, genus Isoperla

5 Eucapnopsis brevicauda, Amphinemura banksi, Malenka coloradensis, genus Sweltsa, S. coloradensis, genus Triznaka, T. pintada, genus Hesperoperla, H. pacifica, Isogenoides elongatus

6 Capnia confusa, C. gracilaria, genus Utacapnia, U. lemoniana, genus Prostoia, P. besametsa, genus Zapada, Z. cinctipes, Sweltsa borealis, Isoperla quinquepunctata

7 Capnia uintahi, Capnura wanica, Malenka californica, genus Podmosta, P. delicatula, Zapada frigida, Z. haysi, Z. oregonensis, Doddsia occidentalis, Taenionema pallidum, T. uninta, genus Alloperla, A. severa, Plumaperla diversa, Swelta fidelis, Isoperla pinta, genus Megarcys, M. signata, genus Diura, D. knowltoni

8 Pteronarcys californica, Capnia coloradensis, C. vernalis, Isocapnia crinata, I. vedderensis, genus Mesocapnia, Utacapnia logana, U. poda, genus Perlomyia, P. utahensis, Malenka flexura, Taenionema pacificum, Suwallia starki, Sweltsa lamba, Isoperla mormona, I. phalerata, genus Cultus, C. aestivalis, genus Kogotus, K. modestus

9 Capnia petila, Isocapnia grandis

10 Pteronarcys dorsata, Capnia uintahi*, Isocapnia integra, genus Pomoleuctra, P. purcellana, genus Nemoura, N. arctica, Alloperla medveda, A. serrata, Sweltsa revelstoka, Isoperla fusca, Arcynopteryx compacta, Kogotus nonus

11 Capnura wanica*, Mesocapnia oenone, Utacapnia columbiana, U. lemoniana*, U. logana*, U. poda*, Amphinemura banksi*, Malenka coloradensis*, Taenionema uinta*, Suwallia forcipata*, Sweltsa lamba*, Triznaka pintada*, genus Kathroperla, K. perdita, Isoperla phalerata*

12 Genus Bolshecapnia, B. milami, Capnia sextuberculata, genus Megaleuctra, Sweltsa albertensis, Calineuria californica, Megarcys watertoni, genus Setvena, S. bradleyi

13 Despaxia augusta, genus Soyedina, Sweltsa occidens, Cascadoperla trictura, Megarcys subtruncata, Skwala curvata, Cultus pilatus, C. tostonus

Illiesia – http://www2.pms‐lj.si/illiesia/ Volume 4 – Number 18 – Page 199

Appendix 2 Continued Clade Unambiguous species and genera changes

14 Capnia elongata, C. excavata, C. melia, C. promota, Isocapnia abbreviata, Isocapnia crinita*, I. integra*, I. spenceri, Mesocapnia projecta, Malenka cornuta, Soyedina producta, Taenionema kincaidii, Alloperla fraterna, genus Haploperla, H. chilnualna, Sweltsa exquista, S. pacifica, Yoraperla mariana, Y. siletz, Doroneuria baumanni, Frisonia picticeps, Setvena tibialis, Osobenus yakimae

15 Pteronarcella regularis, Pteronarcys dorsata*, Bolshecapnia milami*, Bolshecapnia spenceri*, Capnia coloradensis*, C. vernalis*, Mesocapnia porrecta, genus Paracapnia, genus Amphinemura*, Alloperla medveda*, Triznaka pintada, genus Utaperla*, U. sopladora*, genus Soliperla, Yoraperla nigrisoma, Doroneuria theodora*, Calliperla luctuosa, Isoperla bifurcata, I. longiseta*, I. marmorata, Arcynopteryx compacta*, Megarcys signata*, M. watertoni*, Setvena bradleyi*,Cultus aestivalis*, Kogotus modestus*, Rickera sorpta

16 Capnia licina, Capnura elevata, C. venosa, Isocapnia palousa, I. rickeri, Paracapnia ensicala, Utacapnia imbera, Megaleuctra complicata, M. kincaidi, Malenka bifurcata, M. perplexa, M. tina, genus Nemoura*, Podmosta obscura, Taenionema jewetti, T. oregonense, Sweltsa adamantea, Isoperla gravitans, I. rainiera, I. tilasqua, Isogenoides colubrinus*

17 Capnura venosa, Amphinemura banksi, Malenka tina, genus Nemoura*, Soyedina potteri, Sweltsa lamba, Soliperla salish, genus Pictetiella, P. expansa

18 Capnia elongata, C. nearctica, C. pileata, Capnura vernalis*, Mesocapnia variabilis, genus Perlomyia*, P. utahensis*, Amphinemura linda, genus Malenka*, M. californica*, M. flexura*, Nemoura rickeri, Podmosta weberi, Taenionema kincaidii, genus Taeniopteryx*, Alaskaperla ovibovus, Isoperla decolorata, I. fulva*, I. pinta*, I. quinquepunctata*, Diura bicaudata, Isogenoides elongatus*

19 Genus Bolshecapnia, B. milami, Capnia decepta, Capnura fibula, Taeniopteryx parvula, Alloperla pilosa, Sweltsa hondo, genus Perlesta, P. decipiens, Isoperla jewetti, Isogenoides zionensis

20 Pteronarcys dorsata, Amphinemura linda, genus Nemoura, Shipsa rotunda, Isoperla decolorata

21 Capnia confusa, C. gracilaria, genus Paracapnia, P. angulata, Malenka californica, Nemoura rickeri, genus Oemopteryx, O. fosketti, genus Taeniopteryx, T. nivalis, genus Acroneuria, A. abnormis, A. lycorias, genus Paragnetina, P. media, Isoperla bilineata, I. marlynia, I. transmarina, Isogenoides frontalis

22 Capnia decepta, genus Mesocapnia, M. frisoni

23 Genus Anacroneuria

24 Genus Allocapnia

25 Genus Isoperla

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26 Genus Perlesta

27 Allocapnia granulata, Acroneuria arenosa, genus Perlinella, P. drymo, genus Paragnetina

28 Taeniopteryx burksi, T. lita, T. lonicera, Acroneuria evoluta, genus Neoperla, N. carlsoni, N. clymene

29 Genus Pteronarcys, P. dorsata, genus Leuctra, Acroneuria abnormis, Perlinella ephyre, genus Agnetina, A. annulipes, Paragnetina kansensis, genus Helopicus, H. bogaloosa

30 Nemocapnia carolina, Leuctra cottaquilla, L. ferruginea, Amphinemura nigritta, genus Haploperla, Attaneuria ruralis, Eccoptura xanthenes, Perlinella zwicki, Clioperla clio, Isoperla dicala

31 Allocapnia aurora, A. mystica, A. rickeri, Leuctra tenuis, Amphinemura delosa, genus Prostoia, P. completa, Shipsa rotunda, genus Strophopteryx, S. fasciata, Taeniopteryx maura, genus Alloperla, Acroneuria carolinensis, Neoperla occipitalis, genus Diploperla, D. duplicata

32 Pteronarcys biloba, Allocapnia recta, Leuctra biloba, L. moha, Alloperla atlantica, A. usa, genus Sweltsa, Tallaperla laurie, T. maria, Acroneuria filicis, genus Beloneuria, Agnetina flavescens

33 Pteronarcys scotti, Allocapnia granulata*, A. wrayi, Leuctra cottaquilla*, Amphinemura appalachia, A. wui, Alloperla chloris, A. nanina, A. petasata, Sweltsa lateralis, Tallaperla anna, T. cornelia, Veihoperla ada, Beloneuria stewarti, Paragnetina immarginata, Isoperla holochlora, genus Remenus, genus Malerikus, M. hastatus, genus Yugus, Y. arinus

34 Leuctra alexanderi, Leuctra sibleyi, genus Megaleuctra, M. williamsae, Strophopteryx appalachia, Perlesta frisoni, P. nelsoni, Paragnetina ichusa, Isoperla distincta, I. orata, I. similis, Remenus bilobatus, genus Hydroperla, genus Isogenoides, I. varians, Oconoperla innubila

35 Allocapnia fumosa, A. nivicola, A. stannardi, genus Paracapnia, P. angulata, Leuctra carolinensis, L. moha*, L. nephophila, genus Paranemoura, P. perfecta, genus Soydina, S. carolinensis, genus Zapada, Z. chila, genus Oemopteryx, O. contorta, Strophopteryx limata, genus Taenionema, T. atlanticum, Alloperla neglecta, Rasvena terna, Tallaperla elisa, Acroneuria frisoni, A. lycorias, Isoperla lata, Cultus verticalis

36 Allocapnia nivicola, A. rickeri, genus Haploperla, H. brevis

37 Genus Paracapnia, genus Leuctra, L. tenuis, Isoperla transmarina

38 Genus Prostoia, P. completa, genus Strophopteryx, S. fasciata, Taeniopteryx burksi, Acroneuria abnormis, Paragnitina media, Clioperla clio

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39 Genus Pteronarcys, P. pictetii, Allocapnia nivicola*, A. vivipara, Amphinemura delosa, Acroneuria arenosa*, Attaneuria ruralis, Isoperla bilineata, I. longiseta, genus Isogenoides, I. varians

40 Genus Zealeuctra, Z. claasseni, Taeniopteryx metequi, Acroneuria evoluta, A. frisoni, Paragnetina kansensis, Isoperla mohri, I. transmarina*, genus Helopicus, genus Hydroperla, H. crosbyi, H. fugitans

41 Allocapnia mystica, Paracapnia angulata, Amphinemura nigritta, Taeniopteryx parvula, genus Alloperla, A. caudata, Acroneuria filicis, A. internata, A. perplexa, Perlesta decipiens, Neoperla catharae, Agnetina flavescens, Isoperla burksi

42 Allocapnia forbesi, A. illinoensis, A. nivicola, A. recta, A. smithi, Leuctra alta, L. rickeri, L. sibleyi, Zealeuctra fraxina, Amphinemura varshava, Soyedina vallicularia, Perlesta napacola, Neoperla clymene, N. occipitalis, N. stewarti, I. nana

43 Pteronarcys dorsata, Allocapnia indianae, A. ohioensis, genus Ostrocerca, O. truncata, Prostoia similis, Taenipteryx maura, genus Sweltsa, S. onkos, Perlesta adena, Neoperla gaufini, Isoperla longiseta*, I. mohri*, genus Diploperla , D. robusta

44 Allocapnia frisoni, A. pygmaea, A. zola, genus Taenionema, T. atlanticum, Alloperla caudata*, A. chloris, A. idei, A. imbecilla, A. usa, genus Peltoperla, P. arcuata, Acroneuria carolinensis, A. lycorias, Eccoptura xanthenes, Perlesta nitida, Agnetina capitata, Paragnetina kansensis*, Isoperla orata, genus Hydroperla*, H. crosbyi*, H. fugitans*, genus Isogenoides*, I. varians*, genus Malirekus, M. hastatus

45 Allocapnia mohri, A. sandersoni, Strophopteryx arkansae, S. cucullata,Acroneuria abnormis*, Perlesta browni, P. fusca, Neoperla choctaw, N. falayah, N. osage, Isoperla ouachita, I. signata, genus Isogenoides*, I. varians*

46 Allocapnia malverna, A. peltoides, Zealeuctra cherokee, Taeniopteryx maura, Perlesta baumanni, I. bilineata*, I. coushatta, I. longiseta

47 Allocapnia pygmaea, Shipsa rotunda, Acroneuria lycorias, Agnetina capitata, Isoperla richardsoni, I. signata, I. slossonae

48 Pteronarcys dorsata, Allocapnia illinoensis, A. minima, genus Capnia, C. vernalis, Paracapnia opis, Leuctra ferruginea, L. tenella, Prostoia similis, genus Oemopteryx, O. glacialis, Taeniopteryx parvula, Haploperla orpha, Acroneuria internata, Isoperla frisoni, I. lata, Isogenoides frontalis, I. olivaceus

49 Genus Capnura, Capnura manitoba, Paracapnia angulata, Isoperla cotta, I. longiseta*, I. nana

50 Genus Ostrocerca, O. albidipennis, Alloperla banksi, Attaneuria ruralis*, Isoperla richardsoni*, genus Cultus, genus Helopicus

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51 Pteronarcys pictetii*, Leuctra duplicata, Amphinemura nigritta, Ostrocerca complexa, O. truncata, genus Paranemoura, P. perfecta, Alloperla concolor, A. petasata, genus Sweltsa, S. onkos, Acroneuria carolinensis, A. internata*, genus Perlesta*, Perlinella ephyre*, Isoperla montana, Helopicus subvarians

52 Pteronarcys biloba, Allocapnia maria, A. nivicola, Leuctra truncata Amphinemura wui, genus Taenionema, T. atlanticum, Alloperla chloris, A. voinae, Sweltsa lateralis, S. naica, Paragnetina immarginata, Isoperla holochlora, I. orata, I. similis, Isogenoides hansoni

53 Allocapnia granulata*, Amphinemura delosa*, A. linda*, Oemopteryx glacialis*, Isoperla nana*, Isogenoides olivaceous*

54 Oemopteryx contorta, Taeniopteryx maura, genus Tallaperla, T. maria, genus Perlesta, Perlinella ephyre

55 Pteronarcys comstocki, P. proteus, Bolotoperla rossi, Rasvena terna, genus Utaperla, U. gaspesiana, Neoperla occipitalis

56 Allocapnia curiosa, A. frisoni, A. granulata, A. rickeri, genus Capnura*, C manitoba*, Alloperla imbecilla, genus Peltoperla, P. arcuata, Attaneuria ruralis, Eccoptura xanthenes, Agnetina flavescens, Isoperla cotta*, genus Remenus, R. bilobatus, genus Malerikus, M. iroquois

57 Allocapnia aurora, A. harperi, A. minima*, A. wrayi, Paracapnia opis*, Leuctra alexanderi, L. carolinensis, genus Megaleuctra, M. flinti, Amphinemura delosa, Soyedina carolinensis, Strophopteryx appalachia, Taeniopteryx metequi, T. ugola, Alloperla aracoma, A. banksi*, A. biserrata, A. usa, A. voinae*, Acroneuria filicis, A. frisoni, genus Hansonoperla, H. appalachia, Perlesta nelsoni, P. taeysia, Agnetina annulipes, genus Diploperla, D. duplicata, D. robusta, genus Yugus, Y. kirchneri

58 Leuctra rickeri, genus Nemoura*, N. trispinosa*, Alloperla vostoki*, Sweltsa palearata, Isoperla burksi, I. francesca*

59 Allocapnia loshada, A. mystica, Taeniopteryx lita, T. nivalis*, Alloperla idei, Peltoperla tarteri, Acroneuria internata, A. kirchneri, Perlesta frisoni, Isoperla montana*, Diploperla kanawholensis, Malerikus iroquois*

60 Pteronarcys dorsata*, genus Nemoura*, N. trispinosa*, Alloperla banksi*, Sweltsa naica*, Perlesta nitida, Isoperla marlynia*

61 Paracapnia angulata*, Ostrocerca truncata*, Prostoia similis*, genus Oemopteryx*, genus Taeniopteryx*, T. burksi*, T. nivalis*, T. parvula*, Acroneuria carolinensis*, genus Helopicus*, H. subvarians*

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62 Genus Pteronarcys, P. pictetii, Acroneuria abnormis, Perlesta decipiens, P. xube, Isoperla bilineata

63 Genus Leuctra, genus Taenionema, T. atlanticum, genus Alloperla, genus Sweltsa, S. naica, S. onkos

64 Paracapnia opis, genus Podmosta, P. macdunnoughi, Isoperla transmarina

65 Genus Pteronarcys, genus Allocapnia*, genus Amphinemura, A. nigritta, genus Nemoura, N. trispinosa

66 Allocapnia maria, A. pygmaea, Leuctra maria, L. variabilis, genus Paranemoura, P. perfecta, genus Haploperla, H. brevis, Rasvena terna, genus Suwallia, S. marginata, Sweltsa lateralis, Isoperla orata, genus Malerikus, M. Iroquois

67 Genus Anacroneuria

72 Anacroneuria aethiops, A. costana, A. quadriloba

73 Anacroneuria ratcliffei

77 Genus Amphinemura

79 Amphinemura mexicana

80 Amphinemura venusta

81 Genus Anacroneuria

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