Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and 
Tyrrhenoleuctra Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
http://zoobank.org/ urn:lsid:zoobank.org:pub:E891BDE6-1253-4633-9120-AC0BAAA2E627 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 6 – Page 98 
 
 
 
FIRST SEM MICROGRAPHS OF REPRESENTATIVES OF PACHYLEUCTRA 
DESPAX, 1929 AND TYRRHENOLEUCTRA CONSIGLIO, 1957  
(PLECOPTERA: LEUCTRIDAE) 
 
Bill P. Stark1 and Charles H. Nelson2 
 
1 Department of Biology, Box 4045, Mississippi College, Clinton, Mississippi, 39058, U.S.A. 
E-mail: stark@mc.edu 
 
2 Department of Biology, Geology and Environmental Science, 615 McCallie Avenue, The University of 
Tennessee-Chattanooga, Chattanooga, Tennessee, 37403, U.S.A. 
E-mail: charles-nelson@utc.edu 
 
 
 
ABSTRACT 
Adult terminalia of two populations of Pachyleuctra Despax, 1929 and three populations of Tyrrhenoleuctra 
Consiglio, 1957 were examined with scanning electron microscopy in order to provide details of various 
anatomical features exhibited by species in these genera. 
 
Keywords: Plecoptera, Leuctridae, Pachyleuctra, Tyrrhenoleuctra, scanning electron microscopy 
 
 
 
 
INTRODUCTION 
From a phylogenetic perspective, the small 
stonefly genera Pachyleuctra Despax, 1929 and 
Tyrrhenoleuctra Consiglio, 1957 merit interest. 
The proposed phylogeny of the subfamily 
Leuctrinae (Nelson & Hanson 1973, Shepard & 
Baumann 1995), hypothesizes that 
Tyrrhenoleuctra is a basal lineage and the sister 
group of the clade comprising the remaining 
included genera of the subfamily. Pachyleuctra, 
however, is a member of an apical clade and the 
sister group of the species-rich genus Leuctra 
Stephens, 1835. This genus is a Pyrenean group 
with three recognized species after the studies of 
Aubert (1952) and Despax (1951). 
Tyrrhenoleuctra, a stonefly genus known from the 
Iberian Peninsula, North Africa and several 
Mediterranean islands, was proposed by 
Consiglio (1957) as a replacement name for 
Strobliella Klapálek, 1903, after the latter generic 
name was discovered to be a junior homonym of 
Strobliella Kieffer, 1898. This genus currently 
includes five species (DeWalt et al. 2019); two of 
these are morphologically indistinguishable and 
were recently recognized on the basis of 
molecular data (Fochetti & Tierno de Figueroa 
2009, Tierno de Figueroa & Fochetti 2014). 
Beginning in the 1970’s the scanning electron 
microscope (SEM), by providing high resolution 
and increased magnification while retaining great 
depth of field, has been a useful tool in 
morphological studies requiring fine structural 
detail of the phenotype (Wipfler et al. 2016). This 
present study is based on a SEM analysis of a 
small sample of Pachyleuctra and Tyrrhenoleuctra 
specimens and seeks to provide baseline 
information on specific surface structures of these 
insects and to augment previous morphology 
based taxonomic studies as revealed by line 
drawings using the light microscope. 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 99 
MATERIALS AND METHODS 
Pachyleuctra and Tyrrhenoleuctra specimens 
were provided by Dr. Romolo Fochetti, 
Universitá degli Studi della Tuscia, Viterbo, Italy 
(UST), Dr. M.A. Puig Collection, Girona, Spain 
(MAPC), and Dr. Gilles –Vinçon Collection, 
Grenoble, France (VIN). To optimize SEM 
examination of body anatomical structures, 
wings were removed and the torsos placed in an 
ultrasonic cleaner and vibrated for 15 seconds 
before they were dehydrated in solutions of 90%, 
95% and 100% ethanol for 10 minutes in each 
solution. Following this treatment, the 
specimens were placed in hexamethyldisilizane 
for 1 hour, mounted on aluminum stubs with 
double stick copper tape, coated with a 
gold/palladium alloy in a Hummer coater and 
examined with an Amray 1810d scanning 
electron microscope. 
 
RESULTS  
 
Pachyleuctra Despax, 1929 
(Figs. 1–8) 
 
Leuctra (Pachyleuctra) Despax, 1929: 298, type 
species Leuctra (Pachyleuctra) montana Despax = 
Pachyleuctra benllochi (Navás, 1917) 
 
Current species and type localities: 
P. benllochi (Navás, 1917), San Juan del Erm, 
Lérida, Spain 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4308 
 
P. bertrandi Aubert, 1952, Etang de Puig, Pyrenees 
Mountains, France 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4307 
 
P. ribauti Despax, 1929, Saint-Béat, Haute-
Garonne, France 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4305 
 
Material examined: P. benllochi: FRANCE: Ger 
Tributary, Pyrenees Mountains, Mente Pass, 
1300m, 27 May 1993, G. Vinçon, 5♂ 4♀ (VIN). 
SPAIN: Lleida, Pyrenees Mountains, Escrita River 
basin spring, 1809m, 2 July 2008, M.A. Puig, 1♂ 
(MAPC). 
Adult habitus. Slightly brachypterous to 
macropterous in available specimens. General 
color of body and wings dark brown to black 
without distinctive contrasting pigment. 
Male genitalia (n = 3). Epiproct strongly reduced 
(Vinçon & Ravizza 2001). A moderately slender 
pair of long, closely appressed paraprocts 
(specilla) form a recurved probe that extends 
from below the cerci to above the apex of the 
abdomen (Fig. 3); apex of paraprocts recurved 
over abdomen, tips flattened and bearing a small 
circular orifice near the inner subapical margins 
(Figs. 2–4). Apical half of paraprocts bearing long 
slender setae along caudal surface and lateral 
surfaces of the apex (Figs. 3–4). Abdominal terga 
5, 7–9 bear a median pair of spine-like lobes on 
the anterior segmental margins. Cerci somewhat 
conical in shape and bearing a small apical 
nipple-like process (Figs. 1–2). Sternum 9 bearing 
a circular, densely hairy vesicle (Figs. 5–6). 
Female genitalia (n = 2). Subgenital plate on 
sternum 8 consists primarily of a pair of ear-like 
lobes that extend from the basoposterior margin 
of sternum 8 across most of sternum 9 (Fig. 7). 
Lobes and much of sternum 8 clothed with long, 
slender setae. 
Mesosternum (n = 1). Mesosternal furcasternum 
circular in outline (Fig. 8); mesosternal 
basisternum bearing a narrow, dark, hairless 
median line that extends across at least half of the 
basisternum surface (Fig. 8). Mesosternal 
spinasternum relatively long, slender and 
distinctly separated from mesosternal 
postfurcasterna. Metasternal prefurcasterna 
united (Fig. 8). 
 
Tyrrhenoleuctra Consiglio, 1957 
(Figs. 9-–22) 
 
Strobliella Klapálek, 1903: 13, Preoccupied (Kieffer, 
1898) 
Tyrrhenoleuctra Consiglio, 1957: 1, novum nomen, 
type species, Strobliella minuta Klapálek = 
Tyrrhenoleuctra minuta (Klapálek) 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 100 
 
 
 
Figs. 1–6. SEM micrographs of Pachyleuctra benllochi male structures (Escrita River basin, Pyrenees 
Mountains, Spain). 1. Terminalia, dorsal. 2. Details of paraprocts and cerci, dorsal. 3. Terminalia, 
caudolateral. 4. Paraproct apex caudolateral (arrow indicates orifice). 5. Terminalia, ventral. 6. Details of 
vesicle. (ce = cerci; or = orifice; pp = paraproct; ve = vesicle) 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 101 
 
 
Figs. 7–10. SEM micrographs of Pachyleuctra benllochi (Escrita River basin, Pyrenees Mountains, Spain) and 
Tyrrhenoleuctra zavattarii (Rio Pirastzu, Sardinia, Italy) adult structures. 7. P. benllochi female terminalia, 
ventral. 8. P. benllochi, adult mesosternum, ventral. 9. T. zavattarii, female terminalia, ventral. 10. T. 
zavattarii, female subgenital plate. (bs2 = mesobasisternum; cx2 = mesocoxa; fs2 = mesofurcasternum; ml = 
median line; pfs2 = mesopostfurcasternum; sgp = subgenital plate; ss2 = mesospinasternum) 
 
 
Current species and type localities: 
T. antoninoi Fochetti & Tierno de Figueroa, 2009, 
Mallorca Island, Spain 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4121 
 
T. lusohispanica Tierno de Figueroa & Fochetti, 
2014, Algarve, Portugal 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:464237 
 
 
T. minuta (Klapálek, 1903), Sierra Morena, Spain 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4127 
 
T. tangerina (Navás, 1922), Fondak, Morocco 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4125 
 
T. zavattarii (Consiglio, 1956), Monte Limbara, 
Sardinia, Italy 
http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfile.org:
TaxonName:4123 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 102 
 
 
Figs. 11–16. SEM micrographs of Tyrrhenoleuctra zavattarii (Rio Pirastzu, Sardinia, Italy) and 
Tyrrhenoleuctra sp. (Rio Andila, Estrema Dura, Spain). 11. T. zavattarii, male terminalia, dorsal. 12. T. 
zavattarii, details male epiproct, dorsal. 13. T. zavattarii, male cerci and base of paraprocts, caudodorsal. 14. 
T. zavattarii, apex of epiproct and paraprocts, dorsal. 15. T. sp., head and pronotum, dorsal. 16. T. sp. basal 
antennal segments and palpus. (ce = cercus; ep = epiproct; pp = paraprocts) 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 103 
 
 
 
Figs. 17–22. SEM micrographs of Tyrrhenoleuctra sp. (Rio Andila, Estrema Dura, Spain) male structures. 17. 
Male terminalia, caudodorsal. 18. Male epiproct and paraprocts dorsal. 19. Male epiproct and paraprocts, 
dorsolateral. 20. Epiproct, lateral. 21. Epiproct and paraprocts, dorsoapical. 22. Vesicle, ventral. (ce = 
cercus; ep = epiproct; pp = paraprocts; ve = vesicle) 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 104 
Material examined: T. c.f. minuta: SPAIN: 
Sevilla, Arroyo del Moro, Sierra Morena, 38˚ 02’ 
23” N, 5˚ 56’ 10” W, 347m, 18 February 2015, J.M. 
Luzón-Ortega, 2♂ 2♀ 1 larva, specimens damaged 
(UST). T. zavattarii: ITALY: Sardinia, Sardegus, 
Rio Pirastzu, Strada, S. Teresa- Costelsordo Ponte, 
100m, 6 February 2000, 10♂ 3♀ (UST). 
Tyrrhenoleuctra sp.: SPAIN: Estrema Dura, 
Calera De Leon, Rio Andila, 650m, 24 February 
2001, 5♂ 2♀ (UST). 
 
Adult habitus. Brachypterous or macropterous 
among male specimens examined (3–6 mm), 
forewings usually reaching midlength of 
abdomen; females fully winged, forewing length 
(7.5–8.0 mm). General color of wings and body 
pale brown, without distinctive contrasting 
pigment pattern; head with numerous (~30) 
occipital rugosities (Fig. 15), at least among 
specimens of Tyrrhenoleuctra sp. listed above. 
Pronotum with a small number of rugosities 
(~10). Antennae and palpi hairy (Fig. 16). 
Male genitalia (n = 6). Epiproct pronounced; in 
dorsal or caudal view divided by a median 
groove (Figs. 11, 12 & 17, 18). Tyrrhenoleuctra 
zavattarii epiproct dorsally or caudally appears 
triangular (Fig. 11); in profile posteriorly 
hemispherically-shaped, abruptly becoming 
flattened to apex. Tyrrhenoleuctra sp. epiproct 
dorsally or caudally appears caret-shaped (Fig. 
17); in profile posteriorly hemispherically-shaped 
sloping gradually to apex (Fig. 20). Epiproct 
surface of both species covered by uniformly 
distributed short peg-like sensilla bearing 
approximately 80–160 sensilla per epiproct (Figs. 
12 & 18). Epiproct sensilla of T. zavattarii in dorsal 
view (Fig. 12) appear slightly more closely packed 
and of a more uniform length than those 
comprising the sparse patch of Tyrrhenoleuctra sp. 
(Fig. 18). In both T. zavattarii and Tyrrhenoleuctra 
sp. a single pair of long slender paraprocts 
(specilla) extend from a ventrobasal plate on the 
abdominal apex (Fig. 11); paraprocts are closely 
fused and extend through the median groove of 
the epiproct as a spear-like structure (Figs. 11–12 
& 17–19). Apex of paraprocts emerge from 
subapical notch in epiproct and the emergent 
section of the paraprocts bear 5–6 partial, circular 
annuli formed from densely packed single rows 
of short setae (Figs. 12, 14 & 18–21). Paraproct tips 
of T. zavattarii broadly rounded (Figs. 12, 14), 
those of Tyrrhenoleuctra sp. bluntly pointed (Figs. 
20, 21). Cerci oriented vertically to the body axis, 
and shaped somewhat cylindrically with truncate 
apices (Figs. 11, 13 & 17, 19). Vesicle small, 
circular and covered with long hairs throughout 
(Fig. 22). 
Female genitalia (n = 2). Abdominal sternum 7 
larger than other segments, overlapping and 
covering much of sternum 8; sternum 8 with 
extensive membranous areas on the 
posteromedian region of the segment (Fig. 9). 
Median section of posterior margin of segment 7 
truncate for a short section of the segment (Fig. 
10). 
 
DISCUSSION 
Based on the postulated apical systematic 
position of Pachyleuctra in Leuctrinae phylogeny 
several features of the genitalia examined in this 
study are likely apomorphies: male epiproct 
reduced; male paraprocts separated but closely 
appressed; and female subgenital plate with lobes 
extending from the basoposterior margin. 
Recently, as part of a study on Pyrenees 
Leuctridae, Vinçon & Ravizza (2001) reviewed the 
species belonging to this genus and observed that 
P. benllochi closely resembles P. bertrandi. Owing 
to this close similarity these researchers depicted 
the lateral view of the male terminalia of both 
species with one illustration (their fig. 14e, p. 113). 
Added fine details of the male and female 
genitalia of P. benllochi are provided by SEM 
micrographs of this study, but specimens of P. 
bertrandi were not available for examination. 
Vinçon & Ravizza (2001), however, determined 
that these two species can be differentiated by the 
shape of the tergal process found on male 
abdominal segments 5, 7–9. 
In distinguishing between the three traditional 
species of Tyrrhenoleuctra, Aubert (1963) 
considered several characters: body size, wing 
length, leg color, male epiproct length, male 
cercal length, color of female sternites 8 and 9 and 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 105 
shape of the female subgenital plate posterior 
border. Puig et al. (1990), however, described 
morphological variability in T. minuta adult male 
and female wing length and the female tenth 
sternite. Fochetti & Tierno de Figueroa (2009) and 
Tierno de Figueroa & Fochetti (2014) concluded 
that “…morphological characters cannot be 
confidently and consistently used for 
identification of species of Tyrrhenoleuctra”. 
Additionally, they suggested that new species 
descriptions based entirely on morphology are 
not reliable because of variation. Instead they 
relied on differences in the mitochondrial genes 
cytochrome oxidase subunit-1 (COI) and 12s 
ribosomal gene to recognize two additional 
cryptic species belonging to Tyrrhenoleuctra. 
Nonetheless, Fochetti et al. (2009) did observe that 
T. zavattari is both genetically as well as 
morphologically homogenous. SEM images in 
this study indicate that some ultrastructural and 
structural features of the epiproct and paraprocts 
of this species while similar to those of 
Tyrrhenoleuctra sp. are distinct. Various 
researchers [e.g. Wiens (2004), Okiwelu & 
Noutcha (2014), Wanninger (2015)], have 
suggested that morphology is important and that 
an integrated or combined approach 
incorporating both morphological and molecular 
data is necessary in systematic studies. 
In summary, this initial SEM analysis of 
populations of Pachyleuctra and Tyrrhenoleuctra 
has identified a few morphological features, 
especially those involving male and female 
genitalia that might potentially be useful in 
subsequent taxonomic investigations. This result, 
however, is based on limited material and the 
systematic utility of these characters will need to 
be corroborated by additional SEM studies of the 
species of both genera over their known 
geographic range. 
 
ACKNOWLEDGEMENTS 
Specimens used in this study were collected 
by Romolo Fochetti, Julio Miguel Luzón-Ortega, 
Maria Angeles Puig and Gilles Vinçon. We are 
grateful to each of these colleagues for their 
generosity. 
REFERENCES 
Aubert, J. 1952. Plécoptères pyrénéens. Notes sur 
le genera Pachyleuctra Despax et description de 
quelques larves nouvelles. Bulletin de la 
Société Entomologique de France, 57:121–-126. 
Aubert, J. 1963. Les Plécoptères des cours d’eau 
temporaires de la péninsule ibérique. 
Mitteilungen der Schweizerischen Entomologishen 
Gesellschaft, 35:301–315. 
Consiglio, C. 1957. Tyrrhenoleuctra Nomen 
Novum. Bollettino Società entomologica 
italiana, 87:1–2. 
Despax, R. 1929. Description sommaire d'un sous-
genre nouveau et de deux espèces nouvelles 
de genre Leuctra Steph (Plecoptera). Bulletin de 
la Societe Entomologique de France. 298–301. 
Despax, R. 1951. Faune de France. 55. Plécoptères. 
P. Lechevalier, Paris. 280 pp. 
DeWalt, R.E., M.D. Maehr, H. Hopkins, U. Neu-
Becker, & G. Stueber. 2019. Plecoptera Species 
File Online. Version 5.0/5.0. [Retrieved 12 July 
2019]. http://Plecoptera.SpeciesFile.org 
Fochetti, R. & J.M. Tierno de Figueroa. 2009. A 
new species of Leuctridae discovered by 
means of molecular and biochemical 
approaches: Tyrrhenoleuctra antoninoi n. sp. 
(Insecta: Plecoptera). Zootaxa, 2112:41–46. 
Fochetti, R., E. Sezzi, J.M. Tierno de Figueroa, 
M.V. Modica, & M. Oliverio. 2009. Molecular 
systematics and biogeography of the western 
Mediterranean stonefly genus Tyrrhenoleuctra 
(Insecta, Plecoptera). Journal of Zoological 
Systematics and Evolutionary Research, 
47:328–336. 
Kieffer, J.J. 1898. Synopse des Cécidomyies 
d’Europe et d’Algérie décrites jusqu’á ce jour. 
Bulletin de la Société d’Histoire Naturelle de 
Metz. 8:1–64. 
Klapálek, F. 1903. Über neue und wenig bekannte 
Arten der paläarktischen Neuropteroiden. 
Bulletin International de l'Académie des 
Sciences de Bohême, 7:1–14. 
Nelson, C.H. & J.F. Hanson. 1973. The genus 
Perlomyia (Plecoptera: Leuctridae). Journal of 
the Kansas Entomological Society, 16:187–199. 
Okiwelu, S.N. & M.A.E. Noutcha. 2014. The 
evolution of integrative insect systematics. 
 
 
Stark, B.P. and C.H. Nelson. 2019. First SEM micrographs of representatives of Pachyleuctra Despax, 1929 and Tyrrhenoleuctra 
Consiglio, 1957 (Plecoptera: Leuctridae).   Illiesia, 15(06):98-106. https://doi.org/10.25031/2019/15.06 
 
 
Illiesia – http://illiesia.speciesfile.org      Volume 15 – Number 2 – Page 106 
Annual Research & Review in Biology, 4:2301–
2317. 
Puig, M.A., M. Ferreras-Romero, & A. Garcia-
Rojas. 1990. Morphological variability of 
Tyrrhenoleuctra minuta (Klapálek, 1903). Pp. 
357–360. In: Mayflies and Stoneflies: Life 
Histories and Biology. Campbell, I.C. (editor). 
Kluwer Academic Publishers, P.O. Box 17, 
3300 AH Dordrecht, The Netherlands. 366 pp. 
Shepard, W.D. & R.W. Baumann. 1995. Calileuctra, 
a new genus and two new species of stoneflies 
from California (Plecoptera: Leuctridae). Great 
Basin Naturalist, 55:124–134. 
Stephens, J. F. 1836. Illustrations of British 
entomology; or a synopsis of indigenous 
insects: containing their generic and specific 
distinctions with an account of their 
metamorphoses, times of appearance, 
localities, food and economy, as far as 
practicable, with coloured figures (from 
Westwood) of the rare and more interesting 
species. 6:134–145. 
Tierno de Figueroa, J.M. & R. Fochetti. 2014. A 
second new species of Tyrrhenoleuctra 
discovered by means of molecular data: 
Tyrrhenoleuctra lusohispanica n. sp. (Plecoptera: 
Leuctridae). Zootaxa, 3764:587–593. 
http://dx.doi.org/10.11646/zootaxa.3764.5.7 
Vinçon, G. & C. Ravizza. 2001. Leuctridae 
(Plecoptera) of the Pyrenees. Annales de 
Limnologie, 37:293–322. 
Wanninger, A. 2015. Morphology is dead - long 
live morphology! Integrating MorphoEvoDevo 
into molecular EvoDevo and phylogenomics. 
Frontiers in Ecology and Evolution, 3:54–62. 
https://doi.org/10.3389/fevo.2015.00054 
Wiens, J.J. 2004. The role of morphological data in 
phylogeny reconstruction. Systematic Biology, 
53:653–661. 
Wipfler, B., H. Pohl, M.I. Yavorskaya, & R.G. 
Beutel. 2016. A review of methods for 
analyzing insect structures - the role of 
morphology in the age of phylogenomics. 
Current Opinion in Insect Science, 18:60–68. 
 
 
 
 
Submitted 13 August 2019, Accepted 20 September 2019, 
Published 9 October 2019 
Hosted and published at the University of Illinois, Illinois 
Natural History Survey, Champaign, Illinois, U.S.A.