Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: 
Pteronarcyidae).  Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
http://zoobank.org/ urn:lsid:zoobank.org:pub:68D355B3-CD8A-4699-AC4E-94527D18FE24 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 192 
 
 
LARVAE OF NORTH AMERICAN SPECIES OF PTERONARCYS 
(PLECOPTERA: PTERONARCYIDAE) 
 
L.W. Myers1 and B.C. Kondratieff2 
 
1 Lake Champlain Research Institute, SUNY Plattsburgh, Plattsburgh, NY 12901, U.S.A. 
E-mail: myerslw@plattsburgh.edu 
 
2 Department of Bioagricultural Sciences and Pest Management, Colorado State University, 
Fort Collins, Colorado, 80523, U.S.A. 
E-mail: boris.kondratieff@colostate.edu 
 
 
 
 
 
ABSTRACT 
Larvae of the eight North American Pteronarcys (Plecoptera: Pteronarcyidae) species have been difficult or 
impossible to identify over the past century. This stems from the lack of rigorous comparisons of reared 
material. The absence of a reliable key diminishes the importance of Pteronarcys larvae in aquatic ecological 
and biomonitoring studies. We provide comparative larval descriptions and a key illustrated with high 
resolution photographs of important diagnostic characters for the eight North American species of 
Pteronarcys. Earlier descriptions are reviewed and supplemented with new photographs, illustrations and 
morphometric data to aid in the separation of morphologically similar species. 
 
Keywords: Stonefly larvae, Plecoptera, Pteronarcyidae, Pteronarcys, North America 
 
 
 
 
 
INTRODUCTION 
The stonefly genus Pteronarcys Newman, 1838 is 
represented by eight valid species in North America 
(Stark & Szczytko 1982, Nelson 2000, DeWalt et al. 
2017). Two more species, P. reticulata (Burmeister, 
1839) and P. sachalina Klapálek, 1908, are known 
from Mongolia (Judson & Nelson 2012) and western 
Asia (Teslenko 2009), neither of which are the focus 
of this treatment. Stark & Szczytko (1982) 
determined that Pteronarcys sensu Zwick (1973) was 
a polyphyletic taxon, finding that the genus 
Allonarcys Needham & Claassen, 1925 was a 
synonym of Pteronarcys, supporting the 
suppositions of Nelson & Hanson (1971). Nelson & 
Hanson (1971) originally recognized six species 
groups, whereas Stark & Szczytko (1982) only 
recognized three species groups within Pteronarcys. 
Larvae of species that possessed abdominal spines 
were formerly placed in Allonarcys as the P. scotti 
group, and those without abdominal spines in two 
groups as the P. californica group (western) and the 
P. dorsata group (largely eastern). 
Given their large size and widespread 
distribution, larvae and adults of the stonefly genus 
Pteronarcys have often captured the attention of 
aquatic scientists. Larval descriptions and 
illustrations have been provided for P. biloba 
Newman, 1838 (Smith 1917, Claassen 1931, Ricker 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 193 
 
Table 1. Abbreviations and terminology used in morphometric analysis of Pteronarcys larvae. 
 
 
 
1952, Tarter et al. 1975), P. californica Newport, 1848 
(Smith 1917, Claassen 1931, McCafferty 1981, Ward 
et al. 2002), P. comstocki Smith, 1917 (Frison 1942, 
Ricker 1952, Nelson et al. 1977), P. dorsata (Say, 1823) 
(Smith 1917, Shepard & Stewart 1983, Stewart & 
Stark 1988, Stark et al. 1998), P. pictetii Hagen, 1873 
(Frison 1935, Poulton & Stewart 1991), P. proteus 
Newman, 1838 (Smith 1917, Claassen 1931, Ricker 
1952), P. princeps Banks, 1907 (Claassen 1931), and P. 
scotti Ricker, 1952 (Ricker 1952, Shepard & Stewart 
1983). 
Smith (1917) provided one of the first treatments 
of this family in North America, providing useful 
descriptions and illustrations for both larvae and 
adults of P. biloba, P. californica, P. dorsata and P. 
proteus and the adult female of P. comstocki. Claassen 
(1931) provided another much needed review of this 
family with more complete larval descriptions for 
existing members of the family and the first larval 
description and illustration of P. princeps. The first 
larval description of P. pictetii was provided by 
Frison (1935), as P. nobilis Hagen, 1861. Later the 
larvae of P. comstocki were described and illustrated 
by Frison (1942) as Pteronarcys species. Frison (1942) 
also discussed his inability to provide a reliable 
means to separate the larvae of P. pictetii with P. 
dorsata despite the availability of reared, associated 
material. Ricker (1952) described the larvae and 
adults of P. scotti and was also the first to associate 
P. comstocki with Frison’s (1942) description of 
Pteronarcys species. Ricker (1952) also provided a 
means of separating P. proteus and P. scotti based on 
characteristics of the abdominal spines. Despite his 
efforts to distinguish these two species in areas of 
the southern Appalachians where the ranges 
overlap, separation is difficult using his characters. 
A more recent key to mature larvae from the 
southeastern United States was provided by Stark 
APP (Anterior Pronotal Projection) Fig. 1 Projection located at the anterior corners of the pronotum. 
PPP (Posterior Pronotal Projection) Fig. 1 Projection located at the posterior corners of the pronotum. 
APPBW (Anterior Pronotal Projection Basal 
Width) Fig. 1 
A measure of the basal width of the anterior pronotal projection. This 
feature is measured in dorsal view. 
APPMH (Anterior Pronotal Projection 
Median Height) Fig. 1 
The measurement of a line drawn from the midpoint of the base to the 
apex of the projection. Measured in dorsal view.  
APPA (Anterior Pronotal Projection Angle) 
Fig. 1 
Angle formed at the intersection of two straight lines extended from 
the edges of the anterior pronotal projection. 
PPPA (Posterior Pronotal Projection Angle) 
Fig. 1 
Angle formed at the intersection of two straight lines extended from 
the edges of the posterior pronotal projection.  
AAMP (Mean Angle of Male Plate) Fig. 3 Angle measured ventrally at the apex of two lines drawn from the 
lateral edges of the plate on the 8th segment of males of P. dorsata and 
P. pictetii. 
SL (Abdominal Segment Length) Fig. 2 A straight line measured medially and extending from the anterior to 
posterior edge of respective abdominal segment. 
SW (Abdominal Segment Width) Fig. 2 A straight line measured medially from the two lateral edges of the 
respective segment. 
PIM (Projection Inner Margin) Fig. 2 A straight line measured on the inner (posterior) margin of abdominal 
projections that extends from the base to the apex of the projection. 
Measured in dorsal view. 
POM (Projection Outer Margin) Fig. 2 A straight line measured on the outer (anterior) margin of abdominal 
projections that extends from the base to the apex of the projection. 
Measured in dorsal view. 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 194 
 
         
 
Fig. 1. Pronotal projection morphometrics; APP Angle (Anterior Pronotal Projection Angle), APPMH 
(Anterior Pronotal Projection Median Height), APPBW (Anterior Pronotal Projection Basal Width), PPP 
Angle (Posterior Pronotal Projection Angle), PPMH (Posterior Pronotal Projection Median Height), PPPBW 
(Posterior Pronotal Projection Basal Width). 
Fig. 2. Abdominal projection morphometrics; SL (Segment Length), SW (Segment Width), POM (Projection 
Outer Margin), PIM (Projection Inner Margin). 
Fig. 3. Process of AB9 in male of P. pictetii showing measured AAMP angle. 
 
 
(2017), but it lacks a comprehensive analysis of the 
usefulness of the characters for separating the 
species. 
Despite the fact that written descriptions exist for 
all eight species, some descriptions are incomplete 
and none of them present a rigorous analysis 
capable of putting this century-old problem to rest. 
Because of this, no synoptic key to the larvae exists 
that includes all eight North American species. The 
purpose of this paper is to provide a comprehensive 
review of larval characters and assess each for their 
ability to separate larvae of all eight species. Using 
these characters we present a key that separates 
mature larvae of North America species of 
Pteronarcys. 
 
MATERIALS AND METHODS 
Reared adult specimens, associated exuvia, and 
directly associated mature larvae (adult characters 
under larval cuticle) were examined using a Zeiss 
Stemi 200-C dissecting stereomicroscope. 
Photographs of important characters were taken 
using a Cannon Rebel T3i attached to the dissecting 
microscope. Multiple digital images were stacked 
using Zerene Stacker to provide greater image 
depth of field. Drawings were based on individual 
specimens and diagnostic characteristics used were 
compared with collections from other localities. 
In an effort to provide quantitative 
morphological characters, character lengths and 
angles were measured using ImageJ digital 
measurement software (Rasband 2015). Table 1 lists 
the various morphometric characters assessed and 
their abbreviations used in text and figures. APP 
and PPP basal width, median height, and angle of 
pronotal projections (Fig. 1) were measured for all 
eight taxa. Additional analyses of larval abdominal 
projections were conducted on P. biloba, P. proteus 
and P. scotti. These included the length and width of 
abdominal segments and the associated inner and 
outer margins of the abdominal spines (Fig. 2). All 
measurements were taken at a fixed (10X) zoom in
1 
2 
3 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 195 
 
 
 
Fig. 4. APPMH/APPBW pronotal ratios for P. biloba (n=11), P. californica (n=8), P. comstocki (n=6), P. dorsata 
(n=26), P. pictetii (n=16), P. princeps (n=9), P. proteus (n=17) and, P. scotti (n=14). 
 
 
 
 
Fig. 5. PPPMH/PPPBW pronotal ratios for P. biloba (n=11), P. californica (n=8), P. comstocki (n=6), P. dorsata 
(n=26), P. pictetii (n=16) , P. princeps (n=9), P. proteus (n=17) and, P. scotti (n=14). 
4 
5 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 196 
dorsal view. A non-parametric linear regression 
model was used in an attempt to provide 
quantitative boundaries for the separation of the 
three species using the abdominal morphometric 
ratios. This analysis was performed using the “party 
package” in R (Torsten et al. 2006, R Core Team 
2016). 
Additional analysis of the rearward process on 
the ninth sternum of males of P. dorsata and P. 
pictetii was performed by drawing lines caudally 
from the two lateral margins and measuring the 
resulting angle formed at the intersection of the two 
lines (measured in ventral view) (Fig. 3). Last instar 
larvae that could be positively identified or reared 
specimens with associated exuvia, were used in this 
analysis. 
Once data were compiled, means and standard 
deviations were calculated to determine the amount 
of intraspecific and interspecific variability for each 
character. We also used R (R Core Team 2016) to 
perform linear regression trees on morphometric 
ratios from available material to determine the most 
useful character for separating morphologically 
similar species. 
Specimens used in this study were borrowed 
from the following institutional collections: Lake 
Champlain Research Institute, State University of 
New York at Plattsburgh,  New York (LCRI), C. P. 
Gillette Museum of Arthropod Diversity, Colorado 
State University, Fort Collins, Colorado (CSUIC), 
The Illinois Natural History Survey, Champaign, 
Illinois (INHS), The Canadian National Insect 
Collection, Ottawa, Ontario (CNC) the Bill P. Stark 
Collection, Mississippi College, Clinton, Mississippi 
(BPSC), and the North Carolina Department of 
Environmental Protection, Raleigh, North Carolina 
(NCDEP). Specimen data used in this study are 
available at MaterialExaminedPteronarcysKey.csv. 
 
RESULTS AND DISCUSSION 
We used pronotal measurements to provide 
measurable characteristics to aid in the description 
of the pronotal projections traditionally defined in 
the literature with relatively subjective terminology. 
Pteronarcys californica and P. comstocki possessed the 
most well developed pronotal projections with 
acute apices: P. californica (n=8) 
APPMH/APPBW=1.25 ± 0.40 SD (Fig. 4), 
PPPMH/PPPBW=1.14 ± 0.20 (Fig. 5); P. comstocki 
(n=6) APPMH/APPBW=1.29 ± 0.10 (Fig. 4), 
PPPMH/PPPBW=1.05 ± 0.11 (Fig. 5); P. californica 
(n=8) APPA=27.50 ± 9.64 (Fig. 6), PPPA=33.38 ± 8.11 
(Fig. 7), P. comstocki (n=6) APPA=31.83 ± 8.06 (Fig. 6), 
PPPA=31.83 ± 8.42 (Fig. 7). Intermediate 
development of the pronotal projections was noted 
for: P. biloba (n=11) APPMH/APPBW=0.88 ± 0.28 (Fig. 
4), APPA=39.55 ± 13.87 (Fig. 6), PPPMH/PPPBW=0.76 ± 
0.56 (Fig. 5), PPPA=67.55 ± 9.53 (Fig. 7) and P. 
princeps (n=9) APPMH/APPBW=0.64 ± 0.12 (Fig. 4), 
APPA=44.67 ± 14.87 (Fig. 6), PPPMH/PPPBW=0.54 ± 
0.08 (Fig. 5), PPPA=67.44 ± 9.06 (Fig. 7). Intermediate 
development with similar measurements were also 
observed for: P. dorsata (n=26) 
APPMH/APPBW=0.79 ± 0.21 (Fig. 4), APPA=40.96 ± 
20.77 (Fig. 6), PPPMH/PPPBW=0.80 ± 0.22 (Fig. 5), 
PPPA=45.52 ± 12.45 (Fig. 7) and P. pictetii (n=16) 
APPMH/APPBW=0.79 ± 0.10 (Fig. 4), APPA=38.69 ± 
10.04 (Fig. 6), PPPMH/PPPBW=0.81 ± 0.22 (Fig. 5), 
PPPA=47.50 ± 13.08 (Fig. 7) with P. dorsata 
displaying the greatest variability among the two 
species (Figs. 4-7). The pronotal projections of both 
P. proteus and P. scotti displayed relatively weak 
development: P. proteus (n=17) APPMH/APPBW=0.52 ± 0.08 
(Fig. 4), PPPMH/PPPBW=0.44 ± 0.08 (Fig. 5); P. scotti 
(n=14) APPMH/APPBW=0.52 ± 0.08 (Fig. 4), 
PPPMH/PPPBW=0.51 ± 0.40 (Fig. 5) and obtuse 
pronotal projections: P. proteus (n=17) APP 
Angle=71.18 ± 9.28 (Fig. 6), PPP Angle=86.13 ± 9.76 
(Fig. 7), P. scotti (n=14) APP Angle=64.43 ± 9.44 (Fig. 
6), PPP Angle=90.57 ± 9.31 (Fig. 7). 
Several of the diagnostic characters originally 
suggested by Ricker (1952) to separate mature 
larvae of P. proteus and P. scotti were found to be 
unreliable for distinguishing mature larvae of these 
two closely related species. Ricker (1952) reported 
that the longest abdominal projections measured 
along the posterior margin are equal to 1/3 to 1/4 of 
the length of the respective segment for P. proteus. 
For P. scotti, he indicated that the longest projections 
measured along the posterior margin are equal to 
1/5 to 1/6 of the length of the respective segment. 
Although not specifically stated by Ricker (1952), 
we interpreted these ratios as representing our 
proposed PIM/SL morphometric ratio (Fig. 2). 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 197 
 
 
 
Fig. 6. APP Angle pronotal ratios for P. biloba (n=11), P. californica (n=8), P. comstocki (n=6), P. dorsata (n=26), 
P. pictetii (n=16) , P. princeps (n=9), P. proteus (n=17) and, P. scotti (n=14). 
 
 
 
Fig. 7. PPP Angle pronotal ratios for P. biloba (n=11), P. californica (n=8), P. comstocki (n=6), P. dorsata (n=26), 
P. pictetii (n=16) , P. princeps (n=9), P. proteus (n=17) and, P. scotti (n=14). 
6 
7 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 198 
 There was considerable overlap on all abdominal 
segments for PIM/SL (Fig. 8), PIM/SW (Fig. 9) and 
POM/SL (Fig. 11) between P. scotti and P. proteus, 
which accounts for much of the difficulty in 
separating these two species using the abdominal 
projection characteristics described by Ricker (1952). 
Results of a linear tree regression found marginally 
significant differences in the POM/SW on AB4: P. 
proteus (n=17) =0.21 ± 0.04 vs. P. scotti (n=14)=0.15 ± 
0.03 (Figs. 10, 12). AB6 measurements showed 
similar differences: P. proteus=0.17 ± 0.04 vs. P. 
scotti=0.12 ± 0.03 (Figs. 10, 12). 
Additional differences were observed in the 
shape and orientation of the spines in lateral view of 
AB5-7. In P. proteus the spines of AB5-7 when 
viewed laterally are well defined rugose triangular 
projections, the dorsal edge projecting caudally 
perpendicular to the posterior margin of the 
segment (Figs. 50, 53). In P. scotti, the projections on 
AB5-7 viewed laterally appear as low rugose ridges 
that project minimally from the posterior margin on 
AB5 and are nearly imperceptible on AB6-7 (Figs. 
58, 61). 
The antennae of P. proteus have a distinctive 
medial pale band of varying length (Figs. 56, 57), 
this band was absent on all of the P. scotti examined 
(Figs. 64, 66). This character should be used with 
caution because the yellow band can become faded 
and difficult to observe in the exuvia and older 
preserved specimens. Pteronarcys biloba also has a 
yellow medial band on the antennae but can be 
easily differentiated based on the lateral abdominal 
projections. 
While examining material for this study, a large 
proportion of P. dorsata larvae exhibited a convex 
lateral bulge on the pronotum that was clearly 
illustrated in Smith’s (1917) illustration of this 
species. A subsequent illustration of this species in 
Stewart & Stark (2002) did not show this feature. 
This convex bulge is large enough to exceed the 
level of the pronotal projections in most specimens 
examined from the eastern range of this species 
(New York, Virginia to Mississippi) (Figs. 32, 33). 
The bulge was less developed but present in 
populations of the Midwest (Michigan, Minnesota, 
Wisconsin) (Fig. 31) and undeveloped in more 
northern portions of its range (northwestern 
Canada and the Hudson Bay area of northern
 
 
 
 
Fig. 8. PIM/SL ratio AB2-AB8 for P. biloba (n=11), P. proteus (n=17) and P. scotti (n=14) 
8 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 199 
 
 
 
Fig. 9. PIM/SW ratio AB2-AB8 for P. biloba (n=11), P. proteus (n=17) and P. scotti (n=14) 
 
 
 
 
 
Fig. 10. POM/SW ratio AB2-AB8 for P. biloba (n=11), P. proteus (n=17) and P. scotti (n=14) 
 
9 
10 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 200 
Quebec) (Fig. 34). Poulton & Stewart (1991) 
illustrated the pronotum of P. pictetii with a convex 
bulge on the lateral margin that appears to exceed 
the level of the pronotal projections. During our 
examination of P. pictetii specimens, we did observe 
a small convex lateral bulge on the pronotum but no 
specimens were examined where the bulge 
exceeded the level of the pronotal projections (Figs. 
40, 41, 42). 
A couplet in the key to the Pteronarcys of the 
Southeast U.S.A. by Stark (2017) separated P. dorsata 
from P. pictetii by the orientation of the anterior 
pronotal projections. These projections are oriented 
posterolaterally at the tip in P. dorsata; whereas, in 
P. pictetii, are pointed anterolaterally. There is 
variation in this character in both P. dorsata and P. 
pictetii. For P. dorsata, the orientation of the apex of 
the anterior pronotal projection ranged from 
posterolateral for specimens from Alberta, the 
Northwest Territories, Michigan, Virginia, and New 
York to anterolateral for specimens examined from 
Mississippi and Virginia. The apex of the anterior 
pronotal projection of P. pictetii examined during 
this study ranged from lateral to posterolateral 
orientation. 
Harden & Mickel (1952) separated mature male 
larvae of P. dorsata from P. pictetii in Minnesota 
based on the shape of the rearward directed process 
on sternum 9. They noted that on male larvae of P. 
pictetii this process was triangular with the sides 
converging markedly caudad. In P. dorsata, the 
process is more rectangular with the sides being 
nearly parallel. Through our analysis clear 
differences were noted in the angle of this process 
between P. dorsata (n=9)at 33.11° ± 5.16 (Figs. 29, 35) 
and P. pictetii (n=11) at 63.91° ± 13.16 (Figs. 38, 43). 
Unpaired t-test results between these two species 
indicated statistically significant differences in the 
angle of this process (p < 0.0001). Our examination 
of available material of P. pictetii and P. dorsata 
confirmed the ability to separate mature males 
using the shape of the rearward projecting process 
on male larvae. Using the above morphological 
characteristics combined with the known 
geographic ranges of the two species, mature larvae 
may be reliably separated.
 
 
 
 
Fig. 11. POM/SL ratio AB2-AB8 for P. biloba (n=11), P. proteus (n=17) and P. scotti (n=14). 
 
11 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 201 
 
 
 
Fig. 12. Non-parametric linear regression tree comparing abdominal ratios for P. biloba (n=11), P. proteus 
(n=17) and P. scotti (n=14). 
 
 
KEY TO MATURE LARVAE 
 
1 Abdominal segments with conspicuous lateral 
projections (Figs. 13, 14, 17, 25, 27, 50, 51, 53, 54, 
58, 59, 61, 62, 66) ………………………………… 5 
1’ Abdominal segments without lateral projections 
(Figs. 18, 23, 28, 36, 37, 44, 45, 49) ……………... 2 
2 Supra-antennal plate developed into a semi-
circular ridge (Figs. 16, 30, 39, 46, 48, 55, 60, 63); 
APP weakly developed, APPMH/APPBW < 1, 
and with rounded apices (Figs. 31, 36, 40, 44, 47, 
49) ………………………………………………… 3 
2’ Supra-antennal plate developed into a pointed 
semi-lunate conical projection visible dorsally 
(Figs. 19, 22); APP strongly developed 
APPMH/APPBW > 1, with pointed apices (Figs. 
20, 21, 23) …………………………… P. californica 
3 APP oriented laterally occasionally appearing 
reflexed distally (Figs. 31, 32, 33, 34, 36, 40, 41, 42, 
44) ……………………………………………........ 4 
3’ APP oriented anterolateral at a 45° angle (Figs. 
47, 49) ……………………………….….. P. princeps 
4 Convex bulge on lateral margin of pronotum 
sometimes present but does not exceed the level 
of pronotal projections (Figs. 40, 41, 44); male 
larvae with rearward projecting process on 
abdominal sternum 9 triangular in shape and 
with sides nearly converging, resulting angle > 
50° (63.91 ± 13.16) (Figs. 38, 43) ……..…. P. pictetii 
4’ Convex bulge on lateral margin of pronotum 
produced often exceeding level of pronotal 
projections (Figs. 32, 36); male larvae with 
rearward projecting process on abdominal 
sternite 9 rectangular and with nearly parallel 
sides, resulting angle < 39 (33.11 ± 5.16) (Figs. 29, 
35) ………………………………….......... P. dorsata 
5 Prominent acute process on each mesothoracic 
wingpad lateral margin (Figs. 24, 27); supra-
antennal plate developed into pointed, semi-
lunate conical projections (Fig. 26); abdominal 
12 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
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Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 202 
projections with curved inner margins strongly 
developed on segments 1-8 (Figs. 25, 27), 
pronotal projections well developed with curved 
margins and sharp apices.; APPMH/APPBW ≥ 
1.19 (Figs. 24, 27) 
……………………………...…………. P. comstocki 
5’ Prominent acute process absent from each 
mesothoracic wingpad, lateral margins smooth 
(Figs. 15, 17, 57, 66); supra-antennal plate 
developed into a low semi-circular ridge (Figs. 
16, 55, 60, 63); APP not strongly developed, 
apices bluntly rounded; APPMH/APPBW ≤ 1.16 
(Figs. 15, 17, 52, 57, 65, 66) ………………….…... 6 
6 Lateral abdominal projections on AB7 and AB8 
reduced and inconspicuous, AB8 POM/SW < 
0.13; APPA relatively obtuse ≥ 55° and with 
rounded corners (Figs. 52, 57, 65, 66) …….……. 7 
6’ Lateral abdominal projections conspicuous and 
well developed on AB1-8, AB8 POM/SW ≥ 0.13 
(Figs. 1, 13, 14); APPA more acute < 55° (Figs., 15, 
17) ……………………….……….......……. P. biloba 
7 Measured in dorsal view AB4 POM/SW < 0.15; 
AB6 POM/SW < 0.12, lateral abdominal 
projections of AB5-7 nearly indiscernible, 
projecting past the posterior margin of the 
segment minimally in lateral view (Figs. 58, 61) 
………………………………….... P. scotti (in part) 
7’ Measured in dorsal view AB4 POM/SW ≥ 0.15; 
AB6 POM/SW > 0.12, lateral abdominal 
projections of AB5-7 well defined, rugose and 
triangular shaped in lateral view, the dorsal edge 
projecting caudally parallel to the lateral surface 
of the segment (Figs. 50, 53) or lateral abdominal 
projections of AB5-7 nearly indiscernible, 
projecting past the posterior margin of the 
segment minimally in lateral view (Figs. 58, 61) 
………………………….……….......................….. 8 
8 In dorsal view abdominal projections of AB1-7 
well developed with the largest projections 
possessing a convex outer margin, appearing 
hook-shaped in dorsal view (Figs. 51, 54, 57); 
AB5-7 well defined, rugose and triangular 
shaped in lateral view, the dorsal edge projecting 
caudally parallel to the lateral surface of the 
segment (Figs. 50, 53), antennae with a 
distinctive medial yellow band (Figs. 56, 57) 
…………………………………………… P. proteus 
8’ In dorsal view lateral abdominal projections of 
AB1-7 less developed, outer margins of largest 
projections nearly straight (Figs. 59, 62, 66), 
lateral abdominal projections of AB5-7 nearly 
indiscernible, projecting past the posterior 
margin of the segment minimally in lateral view 
(Figs. 58, 61), antennae lacking distinctive yellow 
band  (Figs. 64, 66) …...............…. P. scotti (in part) 
 
Species Accounts and Material Examined 
 
Pteronarcys biloba Newman, 1838 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470669 
(Figs. 13–17) 
 
Remarks and Distribution. Previous descriptions 
of the larva of this species are adequate. The first 
description was provided by Smith (1917). Claassen 
(1931) later provided a more detailed description. 
This species can be reliably separated from other 
members of the genus based on the presence of the 
conspicuous abdominal projections on abdominal 
segments 1-8 (Figs. 13, 14), the lack of a prominent 
acute lateral projection on the mesothorax (Fig. 15), 
and the acute angle (APP > 55) of the anterior 
pronotal projection (Figs. 15, 17). The distribution of 
this species extends from Canadian Maritime 
Provinces south to Alabama (Nelson 2000). 
Material examined. Canada, New Brunswick: 
Gloucester Co., Tobusintac River, off Hwy 8, 17 
June 1993, 4♀, 4L, B.C. Kondratieff & R.W. Baumann 
(CSUIC); Restigouche Co., Trout Brook, 5 June 1956, 
♂, Exv., F.P. Ide & J.K. Malia (CNC); Prince Edward 
Island: Prince Co., Mill River, Hwy 148 L. of 
Howlan, 19 June 1993, 2Exv, B.C. Kondratieff & 
R.W. Baumann (CSUIC); USA, Georgia: Gilmer 
Co., Mountaintown Creek, State Hwy 2152, L 
34.7500, W 85.5544, 9 February 2017, 4L, B.C. 
Kondratieff & C. Verdone (CSUIC); New York: 
Albany Co., Catskill Creek, Rt. 145 nr. Cooksburgh, 
N 42.4183, W 74.2054, 21 May 2008, Exv, L. Myers, 
B.C. Kondratieff & R.W. Baumann (LCRI); 
Cattaraugus Co., Connoisarauley Creek, East Otto 
Rd., Ashford, N 42.4247, W 78.6976, 24 June 2009, 
Exv, E.L. White (LCRI); Clinton Co., Trib. to 
Canfield Brook, jct. Kennedy & Bart Merrill Rd.,
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 203 
 
 
 
Fig. 13-16. Pteronarcys biloba. 13. Abdomen lateral view; New York: Franklin Co., N. B. Saranac River. 14. 
Abdomen dorsal view; New York: Franklin Co., N. B. Saranac River. 15. Head and Pronotum dorsal view; 
New York: Franklin Co., N. B. Saranac River. 16 Supra-antennal process dorsal view; New York: Franklin 
Co., N. B. Saranac River. 
 
 
N 44.7216, W 73.6687, 14 March 2009, L, L. Myers 
(LCRI); True Brook, Bridge on True Brook Rd., N 
44.6470, W 73.7990, 12 November 2009, L, C. 
Binggeli & A. Grooves (LCRI); Essex Co., North 
Branch Boquet River, Moss Rd., Lewis, N 44.3144, W 
73.5176, 6 April 2010, ♀, Exv, L. Myers, reared 25 
April 2010 (LCRI); Roaring Brook, River Rd. nr. 
Lake Placid, N 44.1503, W 73.7674, 22 May 2007, ♂, 
Exv, L. Myers, reared 27 May 2007 (LCRI); West 
Branch Ausable River, Notch on Rt. 86 in 
Wilmington, 10 May 2006, ♀, Exv, L. Myers (LCRI); 
Franklin Co., North Branch Saranac River, Loon 
Lake Rd., N 44.5392, W 74.0515, 31 May 2006, Exv, 
L. Myers, reared 27 May 2006 (LCRI); Lewis Co., 
Indian River, Cemetery Rd. off Rt. 812, N 43.9753, W 
75.3701, 5 June 2008, Exv, L. Myers & R. Mowrey 
(LCRI); Independence River, Pine Grove Rd. N 
43.7301, W 75.3655, 5 June 2008, ♀, Exv, L. Myers & 
R. Mowrey (LCRI); Saint Lawrence Co., Trib. to 
Little River, Rt. 3 nr. Star Lake, N 44.1905, 
13 14 
15 16 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 204 
 
 
 
Fig. 17. Pteronarcys biloba - Larval habitus; New York: Clinton Co., Tributary to Canfield Brook. 
 
 
W 75.0633, 2Exv, 5 June 2008, L. Myers & R. 
Mowrey (LCRI); Warren Co., Hudson River, Glen 
Athol Rd., N 43.5786, W 73.8598, 30 June 2007, L, 
L. Myers & B.C. Kondratieff (LCRI); Vly Brook, Rt. 
8 nr. Wevertown, N 43.6529, W 73.9228, 2 May 
2008, ♂, Exv, L. Myers & R. Mowrey, reared 2 June 
2008 (LCRI); Tennessee: Carter Co., Doe River ca. 
0.5 mi. below Roan Mountain State Park Welcome 
Center, 12 April 1998, ♂, Exv, D.A. & E.L. Etnier, 
reared 17 April 1998 (CSUIC); West Virginia: 
Pocahontas Co., East Fork Greenbrier River, 
Forest Rd. 51 off Rt. 28, 26 April 1987, 3L, B.C. 
Kondratieff & R.F. Kirchner (CSUIC); Randolph 
Co., Gandy Creek, Rt. 40, 15 May 1990, 2L, B.C. 
Kondratieff, J. Welch & R.F. Kirchner (CSUIC). 
 
Pteronarcys californica Newport, 1848 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470665 
(Figs. 18–23) 
 
 
17 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 205 
 
 
Fig. 18-21. Pteronarcys californica – 18. Abdomen dorsal; Colorado: Larimer Co., Poudre River. 19. Supra-
antennal process dorsal view; Colorado: Larimer Co., Poudre River. 20. Head and pronotum dorsal view; 
Colorado: Larimer Co., Poudre River. 21. Head and thorax dorsal view; Colorado: Larimer Co., Poudre 
River. 
 
 
Remarks and Distribution. Previous larval 
descriptions and illustrations are available for this 
species (Smith 1917, Claassen 1931, McCafferty 
1981). It can be separated from larvae of other 
Pteronarcys based on the absence of lateral 
abdominal projections (Figs. 18, 23), well-
developed, acutely pointed pronotal projections 
(Figs. 20, 23), pointed triangular wing pads (Figs. 21, 
23), and a pointed, semi-lunate, conical supra-
antennal process (Figs. 19, 22). This widespread and 
often abundant species is distributed throughout 
the Canadian Rockies from Alaska south to Arizona 
and west to the Coastal Ranges of far western North 
America (Jewett 1959, Baumann et al. 1977, Stewart 
& Oswood 2006). 
Material Examined. Canada, British Columbia: 
Summerland, 12 May 1959, ♀, Exv, R.E. Leech 
(CNC); USA, Colorado: Larimer Co., Big Thompson 
River, Viestenz Smith Park, 9 June 2005, ♀, 3Exv, R. 
Thorp (CSUIC); Big Thompson River, 1 mile N 
Drake, 7 October 2005, 3L, L. Myers (LCRI); Poudre 
River, Fort Collins, Lyons Park, 22 May 1987, ♀, Exv,
18 19 
20 21 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 206 
 
 
 
Fig. 22. Pteronarcys californica. Supra-antennal process; Colorado, Grand Co., Colorado River. 
 
 
 
 
Fig. 23. Pteronarcys californica. Larval habitus; Colorado, Grand Co., Colorado River. 
22 
23 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 207 
B.C. Kondratieff (CSUIC); Grand Co., Colorado 
River, US 40 below Byers Canyon, 4L, 28 April 2010, 
B. Heinold (CSUIC); Oregon: Clackamas Co., 
Salmon River, submerged wood, 1 mi. W of Zig Zag, 
27 December 1995, L, R. Durfee (CSUIC). 
 
Pteronarcys comstocki Smith, 1917 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470670 
(Figs. 24–27) 
 
Remarks and Distribution. This is the most 
infrequently encountered member of this genus. 
Scattered records are available from New 
Brunswick, Maine, New York, Pennsylvania, 
Virginia, West Virginia, and Kentucky (Nelson 
2000). Larvae of P. comstocki are easily separated 
based on the well-developed curved abdominal 
projections on AB1-8 (Fig. 25), pointed, semi-lunate 
supra-antennal process (Fig. 26), acute, well 
developed pronotal projections, and the presence of  
 
 
 
Fig. 24-26. Pteronarcys comstocki. 24. Pronotum and thorax dorsal view; New York: Essex Co., Tributary of 
Ausable River. 25. Abdomen dorsal view; New York: Essex Co., Tributary of Ausable River. 26. Supra-
antennal process; New York: Essex Co., Tributary of Ausable River. 
 
24 25 
26 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 208 
 
 
Fig. 27. Pteronarcys comstocki. Larval habitus; Virginia: Highland Co., Back Creek. 
 
a sharply pointed process on the mesothorax (Fig. 
24, Fig. 27). 
Material Examined. USA, New York: Essex Co., 
Boquet River, Rt. 9N South of Elizabethtown, N 
44.2125, W 73.5839, 23 May 2008, 2Exv, L. Myers, 
B.C. Kondratieff & R.W. Baumann (LCRI); East 
Branch Ausable River, Andrus Rd. Upper Jay, N 
44.3139, W 73.7856, 25 June 2008, Exv, L. Myers 
(CSUIC); East Branch Ausable River, Rt. 9 Ausable 
Forks, N 44.4379, W 73.6795, 8 April 2015, 3L, L. 
Myers (LCRI); Keene NY, Tributary of Sable River, 
20 June 1941, Exv, T.H. Frison & H.H. Ross (CNC); 
Vermont: Windsor Co., White River, Crossing Rt. 
100 S. Quarry Hill Rd., N 43.8976, W 72.8217, 2 April 
2014, L, L. Myers (LCRI); Virginia: Highland Co., 
Back Creek, CR 600/603 Bridge, 29 December 1981, 
♂, ♀, 2Exv, reared 4 April 1982, B.C. Kondratieff & 
R.F. Kirchner (CSUIC). 
 
Pteronarcys dorsata (Say, 1823) 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470662 
(Figs. 28–36) 
 
Remarks and Distribution. The geographical range 
of this widespread species extends from eastern 
North America south to Florida and Louisiana and 
across Canada to Alaska (Earle 2009, Nelson 2000,
27 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 209 
 
 
 
Fig. 28-31. Pteronarcys dorsata. 28. Abdomen dorsal view; Michigan: Otsego Co., West Branch Sturgeon 
River. 29. Ventral view of process on AB 9 of male; Michigan: Otsego Co., West Branch Sturgeon River. 30. 
Supra-antennal process; Michigan: Otsego Co., West Branch Sturgeon River. 31. Pronotum dorsal view; 
Michigan: Otsego Co., West Branch Sturgeon River. 
 
 
Stewart & Oswood 2006). Mature larvae of this 
species can be separated from others based on the 
lack of abdominal projections (Figs. 28, 36), supra-
antennal plate developed into a semi-circular ridge 
(Fig. 30), the presence of a convex lateral bulge on 
the pronotum that often exceeds the level of 
pronotal projections (Figs. 32, 36), and a broadly 
rectangular process on sternum 9 of male larvae, the 
resulting angle=33.11 ± 5.16 (Figs. 29, 35). 
Material Examined. Canada, Alberta: Wood 
Buffalo Co., McMurray, 1 June 1953, 2♀, Exv, S.E. 
Ball (CNC); Northwest Territories: Johnny Hoe 
River, 2 August 1971, L, J.F. Flanagan (CNC); Trail 
River, NWT, 19 September 1971, L, FWI pipeline 
project (CNC); Hearne Channel, Great Slave Lake, 
18-20 July 1947, Exv, W.E. Ricker (CNC); Source of 
Mackenzie River, Great Slave Lake, Mackenzie 
Canada, 31 July 1965, 2♀, Exv, W.E. Ricker (CNC); 
Quebec: River Malian, 1968, L (CNC); Lac Aigneau, 
4 July 1955, 7N, W.E. Ricker (CNC); USA, Michigan: 
Benzie Co., Platte River: Honor at Zimmerman Rd., 
N 44.6602, W 86.0186, 26 March 2011, ♀, Exv, R.E. 
DeWalt & M. Pessino, reared 17 May 2011 (INHS); 
Charlevoix Co., North Branch Boyne River: 1.4 km
28 29 
30 31 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 210 
 
 
 
Fig 32-34. Pteronarcys dorsata. 32. Pronotum dorsal view; Virginia: Montgomery Co., Little River. 33. 
Pronotum dorsal view; New York: Sullivan Co., Delaware River. 34. Pronotum dorsal view; Canada: 
Northwest Territories, Source of Mackenzie River, Great Slave Lake. 
 
 
 
 
Fig. 35. Pteronarcys dorsata. Process of male sternum 9; Mississippi: Simpson Co., Westville Creek. 
 
32 33 
34 
35 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 211 
 
 
 
Fig. 36. Pteronarcys dorsata. Larval habitus; Mississippi: Simpson Co., Westville Creek. 
 
 
N Boyne Falls at US 131, N 45.1802, W 84.9141, 25 
March 2011, ♂, Exv, R.E. DeWalt & M. Pessino, 
reared 20 April 2011 (INHS); Jordan’s River, 2 km S 
East Jordan at Rogers Rd., N 45.1325 W 85.1240, 25 
March 2011, ♀, Exv R.E. DeWalt & M. Pessino, 
reared 6 May 2011 (INHS); Mason Co., Big Sable 
River, 2 km NNW Free-soil at Custer Rd., N 44.1897, 
W 86.2210, 26 March 2011, ♀, Exv, R.E. DeWalt & M. 
Pessino (INHS); Same as previous, ♀, Exv, R.E. 
DeWalt & M. Pessino, reared 19 May 2011 (INHS); 
Osceola Co., Pine River, 2.5 km NE Deguts at 
Norman Rd., N 44.0634, W 85.5482, 26 March 2011, 
♀, Exv, R.E. DeWalt & M. Pessino, reared 16 May 
2011 (INHS); Same as previous, ♂, Exv (INHS); 
Otsego Co., West Branch Sturgeon River, 5.5 km 
NNW Vanderbilt at Thumb Lake Rd., N 45.1859 W 
84.6976, 25 March 2011, ♂, Exv, R.E. DeWalt & M. 
Pessino, reared 4 May 2011 (INHS); Minnesota: 
Cook Co., Cascade River, 13 km W Grand Marias at 
CR-45, N 47.74667, W 90.5255, 4 March 2012, ♀, Exv, 
R.E. DeWalt & M. Pessino, reared 24 April 2011, 
(INHS); Mississippi: Simpson Co., Westville 
36 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
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Fig. 37-40. Pteronarcys pictetii. 37. Abdomen dorsal view; Nebraska: Hall Co., Platte River. 38. Abdomen 
ventral view; Wisconsin: Dunn Co., Red Cedar River. 39. Supra-antennal process; Nebraska: Hall Co., Platte 
River. 40. Pronotum dorsal view; Nebraska: Hall Co., Platte River. 
 
 
Creek, 21 January 1999, L, B. Stark (BPSC); Mill 
Creek, Hwy 472, 16 February 1979, 2L , B. Stark 
(BPSC); Same data as previous, 25 January 1993, 2L, 
B. Stark (BPSC); Westville Creek, Pinole, 12 
February 1998, L, J. Wise (BPSC); New York: 
Sullivan Co., Delaware River, Rt. 55, Berryville, 
Highland, N 41.4764 W 74.9129, 28 May 2009, Exv, 
E. White, P. Novak & G. Kenney (LCRI); Virginia: 
Greene Co., Rapidan River, Ruckersville, 21 March 
1940, ♀, Exv, Frison, Mohr, & Hawkins (CNC); 
Montgomery Co., Little River, Rt. 787, 8 November 
1980, 6♂, 4♀, 6L, 22Exv, B.C. Kondratieff (CSUIC). 
Pteronarcys pictetii Hagen, 1873 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470663 
(Figs. 37–44) 
 
Remarks and Distribution. This species mostly 
occurs in the upper Midwest, however scattered 
records are available from Pennsylvania west to 
Nebraska and Colorado south to Arkansas and 
Kentucky (Nelson 2000, Zuellig et al. 2006). Mature 
larvae of this species can be separated from the 
closely related P. dorsata by the presence of a convex 
lateral bulge that does not exceed the level of the 
37 38 
39 
 
40 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
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Fig. 41-42. Pteronarcys pictetii. 41. Pronotum dorsal view; Nebraska: Red Willow Co., Republican River. 42. 
Pronotum dorsal view, Wisconsin: Dunn Co., Red Cedar River. 
 
 
 
 
Fig. 43. Pteronarcys pictetii. Process of male sternum 9; Nebraska, Deuel Co., South Platte River. 
 
 
pronotal projections (Figs. 40, 41, 44) and the 
approximate triangular shape of the rearward 
projecting process on sternum 9 of male larvae, the 
resulting angle=63.91 ± 13.16 (Figs. 38, 43). 
Material Examined. USA, Colorado: Sedgwick Co., 
South Platte River, Julesburg, N 40.9542, W 
102.3872, 7 March 2008, 2L, B. Heinold & E. Peachey 
(CSUIC); Nebraska: Cherry Co., Niobrara River, Rt. 
83, 9 April 1996, 2L, B.C. Kondratieff & R.W. 
Baumann (CSUIC); Deuel Co., South Platte River, 
Big Springs, Hwy 38, N 41.3889, W 102.0750, 7 
March 2008, ♀, 3L, Exv, B. Heinold & E. Peachey 
(CSUIC); Hall Co., Platte River, Grand Island, 
Mormon State Park, 10 April 1996, ♂, ♀, 2Exv, B.C. 
Kondratieff & R.W. Baumann, reared 20 April 1996 
(CSUIC); Hayes Co., Frenchman River, Rt. 6 W of 
Hamlet, 19 February 1994, 2♂, 2Exv, reared 26 
March 1994, B.C. Kondratieff & H. Rhodes (CSUIC); 
Same as previous, ♂, ♀, 2Exv (CSUIC); Red Willow 
Co., Republican River, 1 mi. S of Perry, 19 February 
1994, ♂, Exv, reared 26 March 1994, B.C. Kondratieff 
& H. Rhodes (CSUIC); Same data as previous, ♂, ♀,
41 42 
43 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 214 
 
 
Fig. 44. Pteronarcys pictetii.  Larval habitus; Nebraska: Deuel Co., South Platte River. 
 
 
Exv, reared 28 March 1994 (CSUIC); Thomas Co., 
Dismal River, Rt. 83, 9 April 1996, ♂, ♀, L, 2Exv, B.C. 
Kondratieff & R.W. Baumann (CSUIC); Middle 
Loup River, Jct. Rts. 2 and 83, 9 April 1996, ♂, Exv, 
B.C. Kondratieff & R.W. Baumann (CSUIC); 
Wisconsin: Dunn Co., Red Cedar River, Sand Creek 
at CR-V, N 45.1683, W 91.6874, 21 March 2011, ♂, 
Exv, R.E. DeWalt & M. Pessino, reared 6 May 2011 
(INHS); Same as previous, ♂, Exv, reared 9 May 
2011 (INHS). 
 
Pteronarcys princeps Banks, 1907 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470666 
(Figs. 45–49) 
Remarks and Distribution. This species is common 
in the Pacific Northwest with scattered reports from 
British Columbia, Idaho, Nevada, and Utah (Jewett 
1959, Baumann et al. 1977, Stewart & Oswood 2006). 
It is sympatric with P. californica, but can be 
differentiated by the anterolateral orientation of the 
APP (Figs. 47, 49) and the semi-circular ridge 
shaped supra-antennal plate (Figs. 46, 48). 
Material Examined. Canada, British Columbia: 
Cultus Lake, 13 March 1937, 3L, W.E. Ricker (CNC); 
Hatchery Creek, Cultus Lake BC, 7 May 1937, ♀, 
Exv, W.E. Ricker (CNC). USA, California: Butte 
Co., Butte Creek, Humboldt Rd., above Colby 
Creek, N 40.1138 W 121.4952, 25 May 2014, 2♂, 3L,
44 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 215 
 
 
 
Fig. 45-47. Pteronarcys princeps. 45. Abdomen dorsal view; California: Butte Co., Butte Creek. 46. Supra-
antennal process; California: Butte Co., Butte Creek. 47. Head and pronotum dorsal view; California: Butte 
Co., Butte Creek. 
 
 
 
Fig. 48. Pteronarcys princeps. Supra-antennal process; California: Siskiyou Co., Big Springs Creek. 
45 46 
47 
48 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
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Fig. 49. Pteronarcys princeps - Larval habitus; California: Siskiyou Co., Big Springs Creek. 
 
 
B.C. Kondratieff & C. Verdone (CSUIC); Butte 
Creek, Humboldt Rd. NW of Butte Meadows, N 
40.0016, W 121.5364, 25 May 2014, 4♂, Exv, B.C. 
Kondratieff & C. Verdone (CSUIC); Nevada Co., 
Rock Creek, North Bloomfield Rd., N 39.3024, W 
120.9785, 24 May 2014, 4♀, Exv, B.C. Kondratieff & 
C. Verdone (CSUIC); Siskiyou Co., Big Springs 
Creek, Mt. Shasta City Park, N 41.3281, W 122.3269, 
23 May 2014, 6♂, 2♀, 2L, Exv, B.C. Kondratieff & C. 
Verdone (CSUIC); Oregon: Clackamas Co., Salmon 
River, submerged wood, 1 mi. W of Zig Zag, 27 
December 1995, L, R. Durfee (CSUIC). 
 
Pteronarcys proteus Newman, 1838 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470671 
(Figs. 50–57) 
Remarks and Distribution. This species ranges 
from Quebec south along the Appalachians to West 
Virginia and Kentucky (Nelson 2000). Larvae of this 
species are most similar to P. scotti but can be 
distinguished by the spines on the abdominal 
segments. In P. proteus, spines of AB 1-7 are well 
developed with the largest projections possessing a 
convex outer margin and appearing hook-shaped in 
dorsal view (Figs. 51, 54, 57). When viewed laterally, 
the spines of AB5-7 are well defined, rugose, 
triangular shaped projections, the dorsal edge 
projecting caudally, parallel to the lateral surface of 
the segment (Figs. 50, 53). Measured in dorsal view 
AB-4 POM/SW ≥ 0.15; AB-6 POM/SW > 0.12 (Care 
should be used in the use of these ratios due to 
marginal overlap in observed measurements with
49 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
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Fig. 50-53. Pteronarcys proteus. 50. Abdomen lateral view; New York: Essex Co., Roaring Brook. 51. Abdomen 
dorsal view; New York: Essex Co., Roaring Brook. 52. Head and pronotum dorsal view; New York: Essex 
Co., Roaring Brook. 53. Abdomen lateral view; Virginia: Botetourt Co., Bear Wallow Creek. 
 
 
P. scotti) (Figs. 10, 51, 54). Live and recently 
preserved specimens also possesses a distinctive 
median yellow antennal band that is absent in P. 
scotti (Figs. 56, 57). 
Material Examined. USA, Massachusetts: 
Berkshire Co., Wright Brook, Falls Rd. nr. NY 
border, N 42.1084, W 73.4807, 4 May 2013, L, L. 
Myers (LCRI); New York: Cattaraugus Co., small 
stream, Limestone Run Rd., 0.3 mi. S Ridge Run Rd., 
N 42.0270, W 78.7010, 11 February 2012, ♀, Exv, L. 
Myers & B.C. Kondratieff, reared 16 April 2012 
(LCRI); Columbia Co., Tributary Green River, SE 
Austerlitz at Harvey Mountain Rd., N 42.3063, W 
73.4667, 26 May 2011, ♀, Exv, R.E. DeWalt & M. 
Pessino, reared 16 May 2011 (INHS); Delaware Co., 
Spring-fed trib. to Emory Brook, Rt. 28 
Fleischmanns, N 42.1511, W 74.5224, 27 May 2009, 
♂, Exv, L. Myers & B.C. Kondratieff, reared 10 June 
2009 (LCRI); Essex Co., Cold Brook, Rt. 9, S. Poke-
O-Moonshine, N 44.3869, W 73.5081, 6 April 2010, 
♂, Exv, L. Myers, reared, 25 May 2010 (LCRI); Cold 
River, High Peaks Wilderness Area, N 44.0886, W 
74.2653, 11 May, 2006, ♀, Exv, L. Myers (LCRI); 
Pettigrew Brook, Bonnieview Rd. nr. Wilmington, N 
44.4302, W 73.8003, 15 March 2006, 3L, L. Myers & 
R. Younghanz (LCRI); Roaring Brook, River Rd. 
50 51 
52 53 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 218 
 
 
 
Fig. 54-56. Pteronarcys proteus. 54. Abdomen dorsal view; Virginia: Botetourt Co., Bear Wallow Creek. 55. 
Supra-antennal process; New York: Essex Co., Roaring Brook. 56. Antennae; New York: Delaware Co., 
Spring-fed tributary to Emory Brook. 
 
 
nr. Lake Placid, N 44.2866, W 73.9324, 22 May 2007, 
♂, Exv, L. Myers, reared 27 May 2007 (LCRI); 
Greene Co., Batavia Kill headwaters, end of Rt. 56, 
Windham, N 42.2888, W 74.1141, 18 June 2008, Exv, 
L. Myers (LCRI); Warren Co., Small trib. to Lake 
George, Rt. 9N nr. Cotton Point, N 43.5183, W 
73.6740, 1 May 2008, ♂, Exv, L. Myers, reared 25 
May 2008 (LCRI); Washington Co., Shelving Rock 
Brook, Shelving Rock Rd., N 43.5519, W 73.5963, 30 
April 2008, ♂, ♀, 2Exv, L. Myers, reared 18 May 2008 
(LCRI); Same data as previous, ♂, ♀, 2Exv, reared 12 
May 2008 (LCRI); North Carolina: Ashe Co., Three 
top Creek, off SR 1100 Below Long Hope Creek, N 
36.4107, W 81.6196, 28 August 2013, L, S. Beaty, E. 
Fleek & D. Black (NCDEP); Caldwell Co., Jackson 
Camp Creek, SR 1372, N 36.0694, W 81.5925, 23 May 
2011, 10L, S. Beaty, E. Fleek & D. Black (NCDEP); 
Caswell Co., Hogans Creek, SR 1301, N 36.4392, W 
79.5153, 11 August 2010, 2L, B. Crouch, S. Beaty & 
V. Holland (NCDEP); Chatham Co., East Branch 
Price Creek, 14 November 1997, L, S. Beaty, E. Fleek 
& D. Black (NCDEP); Guilford Co., Reedy Fork, SR 
2128, N 36.1728, W 79.9533, 10 September 2013, 2L, 
S. Beaty, E. Fleek & D. Black (NCDEP); McDowell 
Co., South Muddy Creek, SR 1764, N 35.6500, W 
81.8550, 9 July 2012, 2L, M. Walters, E. Fleek & S.
54 55 
56 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 219 
 
 
 
Fig. 57. Pteronarcys proteus. Larval habitus; Massachusetts: Berkshire Co., Wright Brook. 
 
 
Beaty (NCDEP); Crooked Creek, SR 1135, N 35.6055, 
W 82.1169, 10 July 2012, 7N, M. Walters, E. Fleek & 
T. Mormon (NCDEP); Stokes Co., Crooked Creek, 
off SR 1626, N 36.5356, W 80.0767, 1 November 2011, 
L, S. Beaty, V. Holland & M. Walters (NCDEP); 
Wilkes Co., Roaring River, SR 1990, N 36.2497, W 
81.0442, 10 August 2011, 2L, M. Walters, E. Fleek & 
M. Hale (NCDEP); Fishing Creek, SR 2381, N 
36.1833, W 81.0506, 3 November 2011, 3L, S. Beaty 
(NCDEP); Shell Creek, SR 2318, N 36.1327 W 
81.3966, 19 April 2012, 3L, S. Beaty, M. Walters & V. 
Holland (NCDEP); Garden Creek, SR 1739 , N 
36.3908, W 81.0711, 8 March 2014, 7L, M. Walters, V. 
Holland & E. Fleek (NCDEP); Watauga Co., Norris 
Fork, SR 1337, N 36.2800, W 81.6766W, 27 August 
2013, 2L, S. Beaty, E. Fleek & D. Black (NCDEP); 
Vermont: Washington Co., Mill Brook, Jct. Rt. 100 
and Fiddlers Green, N 44.1792, W 72.8375, 10 April 
2013, L, L. Myers (LCRI); Virginia: Bath Co., Wilson 
Creek, Rt. 629, 6 April 1979, ♂, Exv, S. Hiner 
(CSUIC); Botetourt Co., Bear Wallow Creek, 0.5 
miles S. Rt. 43,11 April 1979, ♀, Exv, B.C. 
Kondratieff, reared 23 April 1979 (CSUIC); Giles 
Co., Small stream into Dismal Creek at Rt. 42, 15 
March 1980, ♂, Exv, B.C. Kondratieff, reared 1 April 
1980 (CSUIC); Montgomery Co., Craig Creek, Rt. 
621, 3 May 1980, ♂, Exv, B.C. Kondratieff, reared 12 
May 1980 (CSUIC); Nelson Co., South Fork Tye 
River, Rt. 56, 3 May 1980, ♂, Exv, B.C. Kondratieff 
(CSUIC); Small trib. of South Fork Tye River, Rt. 687,
57 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 220 
 
 
Fig. 58.-61. Pteronarcys scotti. 58 Abdomen lateral; Tennessee: Sevier Co., Little River. 59. Abdomen dorsal 
view; Tennessee: Sevier Co., Little River. 60. Supra-antennal process; Tennessee: Sevier Co., Little River. 61. 
Abdomen lateral view; Tennessee: Carter Co., Cades Cove Creek. 
 
 
3 May 1980, ♂, Exv, B.C. Kondratieff, reared 11 May 
1980 (CSUIC); Patrick Co., Rock Castle Creek, Rt. 
605 off Rt. 8, 14 April 1979, ♂, Exv, B. C. Kondratieff, 
reared 26 April 1979 (CSUIC); Rockingham Co., 
Home Quarry Run, Rt. 257, 16 May 1971, ♂, Exv, S. 
Kirkland (CSUIC); Tazewell Co., Cove Creek, Rt. 
622, 1.5 miles NW of Cove Cr., 16 May 1994, 2L, B.C. 
Kondratieff & R.F. Kirchner (CSUIC). 
 
Pteronarcys scotti Ricker, 1952 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:470668 
(Figs. 58–66) 
Remarks and Distribution. When viewed dorsally 
the projections of AB1-7 in P. scotti are less 
developed than those observed in P. proteus, outer 
margins of largest projections appearing nearly 
straight in dorsal view (Figs. 59, 62, 66). Viewed 
laterally the spines of AB5-7 appear as low rugose 
ridges that are nearly indiscernible, projecting past 
the posterior margin of the segment minimally in 
lateral view (Figs. 58, 61). Measured in dorsal view 
AB4 POM/SW <0.15; AB6 ≤ 0.12 (Figs. 10, 59, 62). 
Additionally, this species lacks the distinctive 
medial yellow antennal band that is present in P. 
 
58 59 
60 61 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 221 
 
 
Fig. 62-65. Pteronarcys scotti. 62. Abdomen dorsal view; Tennessee: Carter Co., Cades Cove Creek. 63. Supra-
antennal process; Tennessee: Carter Co., Cades Cove Creek. 64. Antennae; Tennessee: Sevier Co., Little 
River. 65. Pronotum; Tennessee: Sevier Co., Little River. 
 
 
proteus (Figs. 64, 66). This species is restricted to the 
Appalachians with records available from 
Pennsylvania, Virginia, North Carolina, Georgia, 
South Carolina, and Tennessee (Nelson 2000). 
Material Examined. USA, Georgia: Murray Co., 
Jigger Creek, East Cowpen Rd., N 34.9626, W 
84.6386, 9 February 2017, 2L, C. Verdone & B.C. 
Kondratieff (CSUIC); Rabun Co., Trib. to Overflow 
Creek, Overflow Creek Rd., N 34.9539, W 83.2216, 
10 February 2017, 2L, C. Verdone & B.C. Kondratieff 
(CSUIC); North Carolina: Alleghany Co., Little 
River, SR 1128, N 36.4677, W 81.1333, 5 September 
2013, 3L, S. Beaty, V. Holland & M. Walters 
(NCDEP); Avery Co., Horse Creek, US 19E, N 
36.1016, W 81.9864, 7 August 2012, 2L, M. Walters, 
E. Fleek & M. Hale (NCDEP); Buncombe Co., S. 
Hominy Creek, Dale Pen Rd., N 35.4882, W 82.7425, 
1 June 2010, 10L, V. Holland & S. Beaty (NCDEP); 
Caldwell Co., Jackson Camp Creek, SR 1372, N 
36.0694, W 81.5925, 23 May 2011, 10L, S. Beaty, E. 
Fleek & D. Black (NCDEP); Cherokee Co., Worm 
Creek, off US 19, N 35.1959, W 83.7893, 11 February 
2015, 3L, E. Fleek, V. Holland & M. Walters 
(NCDEP); Graham Co., Tulula Creek, SR 1275, N 
35.3205, W 83.8025, 2 September 2014, 3L, S. Beaty, 
M. Walters & V. Holland (NCDEP); Haywood Co., 
East Fork Pigeon River, Off US 276, Hungry Creek, 
N 35.4105, W 82.8100, 12 July 2006, 2L, S. Beaty 
62 63 
64 65 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
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Fig. 66. Pteronarcys scotti. Larval habitus; North Carolina: Macon Co., Cullasaja River. 
 
 
(NCDEP); Cove Creek, nr. Maggie Valley, 22 May 
1970, ♂, Exv, B.C. Kondratieff & R.F. Kirchner 
(CSUIC); Henderson Co., Mills River, SR 1337, N 
35.3986, W 82.5950, 15 August 2007, 3L, S. Beaty, M. 
Walters & D. Black (NCDEP); Mills River, NC 191-
280, N 35.3913, W 82.5680, 27 August 2010, 10L, M. 
Walters, E. Fleek & T. Mormon (NCDEP); Jackson 
Co., Moses Creek, SR 1739, N 35.3147, W 83.1255, 16 
September 2015, 10L, S. Beaty (NCDEP); Macon Co., 
Clear Creek, SR 1618, N 35.0044, W 83.2113W, 15 
July 2014, 4L, S. Beaty, E. Fleek & V. Holland 
(NCDEP); Cullasaja River, Rt. 28 N of Falls, 24 May 
1993, ♂, 3Exv, B.C. Kondratieff & R.F. Kirchner 
(CSUIC); McDowell Co., Catawba River, SR 1274, N 
35.6144, W 82.2300, 10 July 2012, 4L, M. Walter, C. 
Tyndall, E. Fleek & T. Morman (NCDEP); 
Transylvania Co., North Fork French Broad River, 
SR 1326, N 35.2013, W 82.8566, 27 July 2010, 10L, S. 
Beaty, E. Fleek & V. Holland (NCDEP); Indian 
Creek, US 64, N 35.1266, W 82.9147, 14 July 2014, 5L, 
S. Beaty, E. Fleek & V. Holland (NCDEP); Watauga 
Co., Norris Fork, SR 1337, N 36.2800, W 81.6766, 27 
August 2013, 3L, S. Beaty, E. Fleek & D. Black 
(NCDEP); Wilkes Co., Garden Creek, SR 1739, N 
36.3903, W 81.0711, 8 March 2014, 2L, V. Holland, M. 
Walters & E. Fleek (NCDEP); Tennessee: Carter 
66 
 
 
Myers, L.W. and B.C. Kondratieff. 2017. Larvae of North American species of Pteronarcys (Plecoptera: Pteronarcyidae). 
Illiesia, 13(16):192-224.  https://doi.org/10.25031/2017/13.16 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 13 – Number 16 – Page 223 
Co., Doe River ca. 0.5 mi. below Roan Mt. St. Park 
Welcome Center, 12 April 1998, ♂, Exv, D.A. & E.L. 
Etnier (CSUIC). Sevier Co., Little River, Elkmont, 14 
May 1939, ♀, Exv, Frison & Ross (CNC Paratypes); 
West Prong Pigeon River, at park headquarters, 
GSMNP, 21 April 1949, L, 3Exv, W.E. Ricker (CNC). 
Cades Cove Creek, 3.2 mi E Cable Mill, GSMNP, ♂, 
♀, Exv, B.C. Kondratieff (CSUIC); Virginia: Floyd 
Co., Dodd Creek, Rt. 8, 20 March 1980, ♂, Exv, B.C. 
Kondratieff, reared 28 March 1980 (CSUIC); 
Grayson Co., Laurel Creek, Rt. 603, 22 April 1979, 
♀, Exv, R. Firth, reared 9 May 1979 (CSUIC) 
Pittsylvania Co., Sandy Creek, Rt. 58, 11 April 1981, 
♀, Exv, B.C. Kondratieff, reared 20 April 1981 
(CSUIC); Washington Co., Big Laurel Creek, Rt. 
603, Konnarock, 19 April 1980, 3♀, 3Exv, B.C. 
Kondratieff, reared 15 May 1980 (CSUIC). 
 
ACKNOWLEDGEMENTS 
We would like to thank the following 
individuals for their assistance in various aspects of 
this paper, Ed DeWalt, Illinois Natural History 
Survey, Champaign, Illinois; Chris Verdone, 
Colorado State University, Fort Collins, Colorado, 
Lindsey Pett, University of Vermont, Burlington 
Vermont; Tim Mihuc, Lake Champlain Research 
Institute, State University of New York, Plattsburgh, 
New York; Norman Taber, Art Department, State 
University of New York, Plattsburgh, New York; 
Owen Lonsdale, Canadian National Collection, 
Ottawa, Ontario; Steven Beaty, North Carolina 
Division of Water Resources, Raleigh North 
Carolina; Bill Stark, Mississippi College, Clinton, 
Mississippi; Illustrations were provided by Kaitlyn 
Mroczka, Kaitlyn Images, Plattsburgh, New York. 
 
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Submitted 16 October 2017, Accepted 13 December 2017, 
Published 26 December 2017 
Hosted and published at the University of Illinois, Illinois 
Natural History Survey, Champaign, Illinois, U.S.A.