Research Article |
Corresponding author: Eric J. South ( ejsouth@illinois.edu ) Academic editor: Marco Gottardo
© 2019 Eric J. South, R. Edward DeWalt, Mark A. Davis, Michael Jared Thomas.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
South EJ, DeWalt RE, Davis MA, Thomas MJ (2019) A new stonefly species (Plecoptera, Perlidae) from the Interior Highlands USA, with morphological and molecular comparison to other congeneric species. ZooKeys 858: 45-70. https://doi.org/10.3897/zookeys.858.33818
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Thirty-one species of Nearctic Perlesta Banks, 1906 (Plecoptera: Perlidae) are recognized. A new species is described from western Arkansas and eastern Oklahoma, USA, Perlesta sublobata South & DeWalt, sp. nov., from the adult male, adult female, and egg. Perlesta sublobata males are differentiated from other congeners by a combination of a prominent ventral caecum and a distinct dorsal extension of the lateral sclerites of the aedeagus. A preliminary molecular phylogenetic hypothesis is proposed for Perlesta based on 17 congeners and three outgroup taxa using partial mitochondrial cytochrome c oxidase subunit I sequence data. Illustrations, stereomicroscope images, and scanning electron micrographs support the description and comparison to other Perlesta.
Arkansas, Nearctic, new species, Oklahoma, Perlesta sublobata South & DeWalt, stonefly
Two species are recognized from China (
At least eight species of Perlesta co-occur in the United States Interior Highlands, a mountainous region defined by the United States Geological Survey as encompassing southern Missouri, western Arkansas, eastern Oklahoma, and extreme southeastern Kansas (
The holotype male and all paratypes are deposited in the Illinois Natural History Survey (
Terminology of all stages follows
Genomic DNA from 20 P. sublobata specimens, 17 congeners, and three outgroup taxa was extracted using the Qiagen DNeasy Kit, amplified for a fragment of the mitochondrial gene encoding for the COI subunit via polymerase chain reaction using either primers LCO1490 and HCO2198 (
Haplotypes. Description of COI haplotypes for 18 Perlesta species and three outgroup taxa. Number of specimens (N) and GenBank accession number are listed for each corresponding haplotype. Multiple specimens sharing the same haplotype are listed consecutively. All specimens collected from the USA except P. nelsoni Stark, 1989 (Canada). State or province is listed by standard postal abbreviation. Sequences obtained from GenBank are denoted with *.
Species | Sex | GenBank | N |
|
Lat./ Long. | Stream | State/ Prov. | Collector(s) |
---|---|---|---|---|---|---|---|---|
Beloneuria georgiana (Banks, 1914) | ♂ | MH778486 | 1 | 909265 | 34.69804N, -83.78149W | tributary of Dukes Creek | GA | E. J. South |
Perlesta adena Stark, 1989 | ♂ | MH778426 | 1 | 793345 | 36.39021N, -86.25096W | Rocky Creek | TN | S. A. Grubbs |
Perlesta armitagei Grubbs & DeWalt, 2018 | ♂ | MH778427 | 1 | 457510 | 39.0342N, -86.16788W | Little Salt Creek | IN | R. E. DeWalt |
Perlesta bjostadi Kondratieff & Lenat, 2006 | ♂ | MH778428 | 1 | 793346 | 36.84673N, -77.56095W | Nottoway River | VA | B. C. Kondratieff |
Perlesta browni Stark, 1989 | ♀ | MH778429 | 1 | 658464 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South |
Perlesta cinctipes (Banks, 1905) | ♂ | MH778430 | 1 | 658465 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South |
♂ | MH778431 | 1 | 658466 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♀ | MH778432 | 1 | 658467 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778433 | 1 | 658468 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
Perlesta decipiens (Walsh, 1862) | ♂ | MH778434 | 1 | 658778 | 41.3337N, -88.18761W | Kankakee River | IL | A. Yanahan |
♀ | MH778435 | 1 | 658777 | 41.3337N, -88.18761W | Kankakee River | IL | A. Yanahan | |
Perlesta ephelida Grubbs & DeWalt, 2012 | ♂ | MH778436 | 1 | 658780 | 44.72652N, -86.14303W | Platte River | MI | R. E. DeWalt, S. K. Ferguson |
♀ | MH778437 | 1 | 658781 | 44.72652N, -86.14303W | Platte River | MI | R. E. DeWalt, S. K. Ferguson | |
♂ | MH778438 | 1 | 658469 | 37.49727N, -92.63033W | Osage Fork of Gasconade River | MO | E. J. South | |
♂ | MH778439 | 1 | 658470 | 37.49727N, -92.63033W | Osage Fork of Gasconade River | MO | E. J. South | |
♀ | MH778440 | 1 | 658477 | 37.49727N, -92.63033W | Osage Fork of Gasconade River | MO | E. J. South | |
Perlesta frisoni Banks, 1948 | ♂ | *HQ568861 | 2 | NA | 35.62276N, -83.44288W | West Prong Little Pigeon River | TN | R. E. DeWalt |
♂ | *JF884174 | 2 | NA | 35.4968N, -83.8337W | Twentymile Creek | NC | R. E. DeWalt | |
Perlesta golconda DeWalt & Stark, 1998 | ♂ | MH778441 | 1 | 550392 | 41.67397N, -91.56452W | Iowa River | IA | M. Kippenhon |
Perlesta lagoi Stark, 1989 | ♂ | MH778442 | 9 | 658456 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South |
Perlesta lagoi Stark, 1989 | ♂ | MH778443 | 9 | 658457 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South |
♂ | MH778444 | 9 | 658460 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778445 | 9 | 658461 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778446 | 9 | 658462 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♀ | MH778447 | 9 | 658471 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♀ | MH778448 | 9 | 658472 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♀ | MH778449 | 9 | 658473 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778450 | 9 | 658474 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778451 | 1 | 658459 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778452 | 1 | 658475 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778453 | 1 | 658476 | 38.45202N, -92.48643W | South Moreau Creek | MO | E. J. South | |
♂ | MH778454 | 1 | 658776 | 41.66065N, -81.11747W | Bates Creek | OH | E. J. South, R. E. DeWalt | |
Perlesta mihucorum Kondratieff & Myers, 2011 | ♂ | MH778455 | 1 | 793347 | 42.4401N, -73.8137W | Hannacroix Creek | NY | L. Myers, J. Myers |
Perlesta nelsoni Stark, 1989 | ♂ | *KR144298 | 1 | NA | 45.976N, -66.719W | St. John River | NB | K. Heard et al. |
Perlesta ouabache Grubbs & DeWalt, 2011 | ♂ | MH778456 | 1 | 516699 | 42.45994N, -89.23985W | Sugar River | IL | R. E. DeWalt et al. |
Perlesta roblei Kondratieff & Kirchner, 2003 | ♂ | MH778457 | 1 | 793348 | 36.4684N, -77.1443W | Kirbys Creek | NC | B. C. Kondratieff et al. |
Perlesta sublobata sp. nov. | ♂ | MH778458 | 8 | 793207 | 36.12052N, -93.69319W | War Eagle Creek | AR | E. J. South |
♂ | MH778459 | 8 | 793212 | 34.03869N, -93.41752W | Antoine River | AR | E. J. South | |
♂ | MH778460 | 8 | 793218 | 34.17985N, -93.07021W | Caddo River | AR | E. J. South | |
♂ | MH778461 | 8 | 793230 | 36.04161N, -93.70482W | War Eagle Creek | AR | E. J. South | |
♂ | MH778462 | 8 | 793266 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♂ | MH778463 | 8 | 793271 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♂ | MH778464 | 8 | 793273 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
Perlesta sublobata sp. nov. | ♂ | MH778465 | 8 | 793288 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South |
♂ | MH778466 | 1 | 793208 | 36.12052N, -93.69319W | War Eagle Creek | AR | E. J. South | |
♂ | MH778467 | 3 | 793209 | 35.66925N, -93.83033W | Mulberry River | AR | E. J. South | |
♂ | MH778468 | 3 | 793233 | 33.97121N, -94.22292W | Cossatot River | AR | E. J. South | |
♀ | MH778469 | 3 | 793324 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♂ | MH778470 | 2 | 793211 | 35.66925N, -93.83033W | Mulberry River | AR | E. J. South | |
♂ | MH778471 | 2 | 793224 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♂ | MH778472 | 1 | 793214 | 34.17985N, -93.07021W | Caddo River | AR | E. J. South | |
♂ | MH778473 | 1 | 793234 | 33.97121N, -94.22292W | Cossatot River | AR | E. J. South | |
♂ | MH778474 | 1 | 793299 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♂ | MH778475 | 1 | 793312 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♀ | MH778476 | 1 | 793328 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
♀ | MH778477 | 1 | 793331 | 33.95608N, -93.44362W | Little Missouri River | AR | E. J. South | |
Perlesta teaysia Kirchner & Kondratieff, 1997 | ♂ | MH778478 | 1 | 515560 | 39.1355N, -86.1601W | tributary of Middle Fork Salt Creek | IN | R. E. DeWalt |
♀ | MH778479 | 1 | 457522 | 39.1355N, -86.1601W | tributary of Middle Fork Salt Creek | IN | R. E. DeWalt | |
Perlesta WI-1 (undescribed) | ♂ | MH778480 | 1 | 516410 | 46.07721N, -92.24608W | St. Croix River | WI | R. E. DeWalt et al. |
♀ | MH778481 | 1 | 552631 | 46.07721N, -92.24608W | St. Croix River | WI | R. E. DeWalt, S. K. Ferguson | |
♂ | MH778482 | 1 | 576963 | 45.57953N, -87.78796W | Menominee River | WI | R. E. DeWalt et al. | |
♂ | MH778483 | 1 | 658779 | 45.77348N, -92.78164W | St. Croix River | MN | R. E. DeWalt | |
♀ | MH778484 | 1 | 583370 | 45.82306N, -92.77001W | Snake River | MN | R. E. DeWalt | |
Perlesta xube Stark & Rhodes, 1997 | ♂ | MH778485 | 1 | 790543 | 39.40911N, -88.89952W | Mud Creek | IL | E. J. South, R. E. DeWalt |
Perlinella drymo (Newman, 1839) | ♂ | MH778487 | 1 | 514716 | 40.2942N, -87.2546W | Wabash River | IN | R. E. DeWalt, M. Pessino |
Perlinella ephyre (Newman, 1839) | ♂ | MH778488 | 1 | 548835 | 42.32815N, -83.8595W | Huron River | MI | R. E. DeWalt et al. |
We generated a maximum likelihood tree using MEGA 7.0 (
Males are distinguished by a combination of a prominent ventral caecum with a broad ventral setal patch and a distinct dorsal extension of the lateral sclerites of the aedeagus. Females possess a subgenital plate with a deep V-shaped notch and truncate lobes. Eggs have a smooth chorion and a well-developed, distally flanged collar.
Male. Habitus moderately dark (Fig.
Perlesta sublobata sp. nov., paratype males from Little Missouri River, Arkansas A habitus, dorsal view (
Perlesta sublobata sp. nov., terminalia of paratype males from Little Missouri River, Arkansas A tenth tergite and paraprocts, dorsal view (
Perlesta sublobata sp. nov., aedeagus of paratype male from Little Missouri River, Arkansas (
Female. Female habitus similar to male, but of larger size and wings of lighter pigmentation (Fig.
Perlesta sublobata sp. nov., paratype females from Little Missouri River, Arkansas A habitus, dorsal view (scale bar) B head and pronotum (
Egg. Length ca. 360 µm, width ca. 280 µm. Chorion smooth with fine pitting (Fig.
Perlesta sublobata formed a monophyletic group with strong support (ML bootstrap support = 97%, Bayesian posterior probability = 92%). The nearest neighbor species to P. sublobata was P. decipiens (Walsh, 1862) at 1.8% sequence divergence. Maximum intraspecific COI genetic distances were less than minimum interspecific distances within all tested Perlesta (Table
Intra and interspecific distance. Maximum intraspecific and minimum interspecific (nearest neighbor) Kimura 2-parameter values for COI within Perlesta. Key: N = number of specimens, P. WI-1 is an undescribed species from Wisconsin.
Species | N | Maximum intraspecific distance (%) | Nearest neighbor | Nearest neighbor distance (%) |
---|---|---|---|---|
P. adena | 1 | 0 | P. nelsoni | 19.3 |
P. armitagei | 1 | 0 | P. xube | 7.2 |
P. bjostadi | 1 | 0 | P. decipiens | 4.5 |
P. browni | 1 | 0 | P. armitagei | 15.1 |
P. cinctipes | 4 | 0.7 | P. armitagei | 10.4 |
P. decipiens | 2 | 1.2 | P. lagoi | 1.5 |
P. ephelida | 5 | 0.7 | P. ouabache | 4.8 |
P. frisoni | 2 | 0 | P. nelsoni | 14.9 |
P. golconda | 1 | 0 | P. WI-1 | 17.8 |
P. lagoi | 13 | 0.8 | P. decipiens | 1.5 |
P. mihucorum | 1 | 0 | P. sublobata | 4.8 |
P. nelsoni | 1 | 0 | P. teaysia | 9.4 |
P. ouabache | 1 | 0 | P. sublobata | 2 |
P. roblei | 1 | 0 | P. ephelida | 15.6 |
P. sublobata | 20 | 0.8 | P. decipiens | 1.8 |
P. teaysia | 2 | 0.2 | P. frisoni | 15.9 |
P. WI-1 | 5 | 2.4 | P. ouabache | 3.9 |
P. xube | 1 | 0 | P. armitagei | 7.2 |
Maximum Likelihood phylogenetic reconstruction of 44 unique Perlesta CO1 haplotypes using the GTR+I+G nucleotide substitution model. Haplotypes represented by more than one specimen are indicated in parentheses beside corresponding GenBank accession numbers. Outgroup taxa: Beloneuria georgiana, Perlinella drymo, and Perlinella ephyre. Bootstrap scores from 1,000 replicates are displayed at nodes. Scale bar represents the estimated number of nucleotide substitutions per site.
Bayesian phylogenetic reconstruction of 44 unique Perlesta CO1 haplotypes using the GTR+I+G nucleotide substitution model. Haplotypes represented by more than one specimen are indicated in parentheses beside corresponding GenBank accession numbers. Outgroup taxa: Beloneuria georgiana, Perlinella drymo, and Perlinella ephyre. Posterior probabilities are indicated at nodes. Scale bar represents the estimated number of nucleotide substitutions per site.
The shape and armature of the aedeagus are the most distinct morphological features of P. sublobata.
Perlesta golconda, aedeagi of males from Missouri River, Nebraska A lateral view showing dorsal caecum (DC) and ventral caecum (VC) (
The known distribution of P. golconda, originally limited to Illinois (DeWalt and Stark 1998), has expanded to include Iowa, Indiana, Michigan, and Nebraska (
The female habitus of P. sublobata resembles two Interior Highlands congeners, P. decipiens and P. ephelida Grubbs and DeWalt, 2012. However, P. sublobata differs from P. decipiens and P. ephelida by subgenital plate morphology. Perlesta decipiens has a deep U-shaped notch bordered by truncate lobes, typically with darker pigmentation on the posterior margins (Fig.
With the exception of one locality (OK, Washington Co., Caney River), all collection sites for P. sublobata are within or closely adjacent to the Interior Highlands, a region containing four contiguous U. S. Environmental Protection Agency (EPA) Level III Ecoregions: Ozark Highlands, Boston Mountains, Arkansas Valley, and Ouachita Mountains. Collection sites for P. sublobata within the Interior Highlands are partially canopied, hardwood forested, wadeable, low gradient streams (ca. 15–20 m wide) with substrata composed mostly of sand, gravel, and cobble. The type locality is a low gradient run (ca. 25 m wide) of the Little Missouri River (Fig.
The specific epithet is derived from sub, Latin for under, and lobata, the feminine adjectival form of lobus, Latin for a rounded projection or protuberance (
Holotype: ♂, in 95% ethanol, USA: Arkansas: Pike Co., Little Missouri River, 10.0 km SSE Delight at AR-19, 33.95608, -93.44362, 15.vi.2016, E. J. South (
Paratypes. USA: Arkansas: Clark Co., Caddo River, 6.7 km NNW Arkadelphia at Super 8 Motel at US-67, 34.17985, -93.07021, 14.vi.2016, E. J. South, 6♂ (
USA: Arkansas: Franklin Co., Mulberry River, Hwy 23 at Turner’s Bend, 35.66984, -93.82962, 5.vii.1986, B. C. Poulton, 4♂ (
USA: Illinois: Carroll Co., Mississippi River, Savanna, 42.09622, -90.16227, 19.vi.1999, R. E. DeWalt, ♂ (
1 | Fully everted aedeagus with dorsal caecum (fig. 7.273) | 1a |
– | Fully everted aedeagus without dorsal caecum (fig. 7.361) | 17 |
1a | Aedeagus with ventral caecum | 1b |
– | Aedeagus without ventral caecum | 2 |
1b | Ventral caecum prominent, length ca. 2/5 aedeagal sac width, fine ventral seta-like spines present; dorsal caecum moderately developed (Fig. |
P. sublobata |
– | Ventral caecum less prominent, length ca. 1/3 aedeagal sac width, without distinct fine ventral seta-like spines; dorsal caecum poorly developed; dorsal aedeagal patch with lateral margins darker than mesal field, appearing as two tracks (fig.7.364); ocelli usually connected by V-shaped area with pale center | P. golconda |
We thank Dr. Sam W. Heads (