Corresponding author: R. Edward DeWalt (
Academic editor: C. Geraci
Ohio is an eastern USA state that historically was >70% covered in upland and mixed coniferous forest; about 60% of it glaciated by the Wisconsinan glacial episode. Its stonefly fauna has been studied in piecemeal fashion until now. The assemblage of Ohio stoneflies was assessed from over 4,000 records accumulated from 18 institutions, new collections, and trusted literature sources. Species richness totaled 102 with estimators Chao2 and ICE Mean predicting 105.6 and 106.4, respectively. Singletons and doubletons totaled 18 species. All North American families were represented with
Regional biodiversity studies are of great importance for setting conservation priorities, in determining conservation status of species, and in examining factors that govern diversity (
Ohio is an eastern state of the USA with a total area of 105,910 km2. It is bound on the south and east by the Ohio River and drained by 10 United States Geological Survey six digit scale Hydrologic Drainage Units (USDA 2009, HUC6s) (
HUC6 drainages and point locations for Ohio
Pre-European settlement vegetation percentage cover for Ohio (from
Ohio’s stonefly fauna has been studied in a piecemeal fashion.
The coauthors have embarked on a study of the stonefly fauna of the Midwest, including distribution modeling of up to 160 species known from Illinois, Indiana, Michigan, Ohio, Ontario, and Wisconsin in order to reconstruct pre-European settlement range. Given that there has been no comprehensive assessment of the stonefly assemblage in Ohio we have elected to prepare one. This analysis is based on the accumulated specimen records from our own efforts, the efforts of colleagues over several decades, the examination of nearly 30,000 specimens borrowed from regional museums, and reliable literature records. We ask several questions of these data:
• How many stonefly species inhabit Ohio?
• How completely has the fauna been sampled?
• How are functional niche traits distributed across the assemblage?
• Does drainage affiliation affect assemblage composition and species richness?
Specimens are the only resource where identifications may be verified, so the study was based on an abundance of specimens examined from 18 regional museums (
Specimen origin, institutional coden, number of specimen records, and number of specimens examined.
Institution | Coden | #Records | Specimens |
Brigham Young University | BYU | 1167 | 18811 |
B. P. Stark Collection | BPSC | 6 | 81 |
Canadian National Collection | CNC | 46 | 252 |
Cincinnati Museum of Natural History | CMNH | 2 | 2 |
Cleveland Museum Natural History | CLEV | 66 | 171 |
Field Museum Natural History, Chicago | FMNH | 13 | 40 |
Illinois Natural History Survey | INHS | 639 | 2839 |
Michigan State University | MSUC | 11 | 63 |
Ohio Biological Survey | OBS | 573 | 2690 |
Ohio Environmental Protection Agency | OEPA | 83 | 142 |
Ohio Historical Society | OHSC | 17 | 17 |
Ohio State University | OSU | 468 | 668 |
Purdue University | PURC | 7 | 18 |
R. Fred Kirchner Collection | RFKC | 164 | 857 |
Royal Ontario Museum | ROME | 3 | 15 |
Southern Illinois University Carbondale | SIUC | 1 | 5 |
University of Michigan | UMMZ | 3 | 3 |
Western Kentucky University | WKU | 170 | 873 |
Literature | Author Year | 641 | 4940 |
Total | 4,080 | 32,487 |
New specimens were collected using sweep nets, beating sheets, hand picking, and dipnetting throughout the state. Many nymphs were reared in Styrofoam cups or in a Frigid Units Living Stream at the University of Illinois. Illinois Natural History Survey (INHS) specimens collected after 2007 were preserved in 95% EtOH and stored in a –20C freezer for future molecular studies.
Locations for all specimens were georeferenced to the finest scale permitted by the label data. Coordinate precision for each record was marked as a radius about the location: from GPS = code 1(10 m); post–processed with small town or road crossing and stream name = code 2 (1,000 m); town name only or large town with stream name = code 3(10,000 m), county level record = code 4 (100,000 m); state level record = code 5 (1,000,000) m. Only records with codes 1–3 were used for species accumulation curves and nonparametric multidimensional scaling (NMDS) analyses.
Ohio stoneflies. Stream widths inhabiting and functional niche traits. Width 1=seep, 2=1–2 m, 3=3–10 m, 4=10–30 m, 5=30–60 m, 6=>60 m, 7=Lake Erie. Voltinism 1, 2 or 3 yr; development 1=fast, 2=slow seasonal. Diapause 1=present, 2=absent. Dispersal Season W=winter, Sp=spring, Su=summer. Feeding O=omnivore, P=predator, S=shredder. Female Mobility L=low, M=moderate, H=high. Nymphal Growth=months of growth, Respiration 1=no gills, 2=with gills. Size at maturity 1=<9 mm, 2=9–16 mm, 3=>16 mm. Emergence Synchrony 1=>1 mo., 2=<1 mo. Thermal preference 1=coldwater, 2=coolwater, 3=warmwater. Active hyperlinks are embedded LSIDs linking to species pages in the
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1–5 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
2–3 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
3–6 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 3 | |
1–4 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
3–4 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
2–4 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
2–5 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
1–4 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
3 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
3 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
2–5 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
2–5 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
2 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
1–6 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 3 | |
3 | 1 | 1 | 1 | W | S | L | 6 | 1 | 1 | 1 | 2 | |
1–4 | 1 | 2 | 2 | Sp | S | M | 11 | 1 | 1 | 2 | 1 | |
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1–2 | 1 | 2 | 2 | Su | S | M | 11 | 1 | 1 | 1 | 2 | |
3 | 1 | 2 | 2 | Su | S | M | 11 | 1 | 1 | 2 | 2 | |
2–4 | 1 | 2 | 2 | Su | S | M | 11 | 1 | 1 | 2 | 2 | |
1–4 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 1 | 2 | |
2–4 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 1 | 2 | |
2 | 1 | 2 | 2 | Su | S | M | 11 | 1 | 1 | 2 | 2 | |
2–4 | 1 | 1 | 2 | Su | S | M | 6 | 1 | 1 | 1 | 2 | |
1–4 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 2 | 2 | |
2–3 | 1 | 1 | 1 | W | S | M | 6 | 1 | 1 | 1 | 2 | |
2–3 | 1 | 1 | 1 | W | S | M | 6 | 1 | 1 | 1 | 2 | |
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1–5 | 1 | 1 | 1 | Sp | S | M | 6 | 2 | 1 | 1 | 2 | |
2–4 | 1 | 2 | 1 | Su | S | M | 9 | 2 | 1 | 1 | 1 | |
1–6 | 1 | 1 | 1 | Sp | S | M | 6 | 2 | 1 | 1 | 2 | |
1–3 | 1 | 2 | 2 | Su | S | M | 11 | 1 | 1 | 1 | 1 | |
2–3 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 1 | 1 | |
2–3 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 1 | 1 | |
2–3 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 2 | 2 | |
2–3 | 1 | 1 | 1 | Sp | S | M | 6 | 1 | 1 | 2 | 2 | |
1–3 | 1 | 1 | 1 | W | S | M | 11 | 1 | 1 | 1 | 1 | |
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3–6 | 1 | 1 | 1 | W | S | M | 6 | 1 | 2 | 1 | 2 | |
2–6 | 1 | 1 | 1 | W | S | M | 6 | 2 | 2 | 1 | 3 | |
6 | 1 | 1 | 1 | W | S | M | 6 | 2 | 2 | 1 | 3 | |
2–5 | 1 | 1 | 1 | W | S | M | 6 | 2 | 2 | 1 | 3 | |
3–5 | 1 | 1 | 1 | W | S | M | 6 | 2 | 2 | 1 | 2 | |
3–5 | 1 | 1 | 1 | W | S | M | 6 | 2 | 2 | 1 | 2 | |
3–5 | 1 | 1 | 1 | W | S | M | 6 | 2 | 2 | 1 | 2 | |
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2–4 | 1 | 1 | 1 | Sp | P | M | 6 | 1 | 1 | 2 | 2 | |
2–4 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
3 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
2–3 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
3 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
2–3 | 1 | 1 | 2 | Sp | P | M | 6 | 1 | 1 | 2 | 2 | |
2–4 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
1–3 | 1 | 1 | 1 | Sp | P | M | 6 | 1 | 1 | 2 | 2 | |
1–5 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
3 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 1 | 2 | 1 | |
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3–6 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
2–5 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
5 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
4–6 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 3 | |
2–6 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
2–7 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 3 | 2 | 2 | |
4–6 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
3 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
2 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
3–6 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 3 | |
4 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
2–5 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
2–6 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
4 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 3 | |
2 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 1 | 1 | |
3–5 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 2 | |
3–5 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 3 | |
3–5 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 2 | |
3–6 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 2 | |
4–7 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 3 | |
3–6 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 2 | |
3–5 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 2 | |
2–6 | 1 | 2 | 2 | Su | P | H | 11 | 2 | 2 | 2 | 2 | |
3–5 | 2 | 2 | 2 | Su | P | H | 23 | 2 | 3 | 2 | 2 | |
2–6 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 2 | |
3–6 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 3 | |
3 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 3 | |
3–6 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 3 | |
3–6 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 2 | |
2–5 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 2 | |
4 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 2 | |
3-6 | 1 | 1 | 1 | Su | P | H | 4 | 2 | 2 | 2 | 2 | |
3–5 | 1 | 1 | 1 | Su | P | H | 9 | 2 | 3 | 2 | 2 | |
3–6 | 1 | 1 | 1 | Su | P | H | 9 | 2 | 2 | 2 | 3 | |
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1–5 | 1 | 1 | 1 | Sp | P | H | 9 | 1 | 2 | 2 | 2 | |
2–3 | 1 | 2 | 2 | Sp | P | H | 11 | 1 | 2 | 2 | 1 | |
1–5 | 1 | 1 | 1 | Sp | P | M | 6 | 1 | 2 | 2 | 2 | |
3–6 | 1 | 1 | 1 | Su | P | H | 6 | 1 | 2 | 2 | 3 | |
2 | 1 | 1 | 1 | Sp | P | M | 6 | 1 | 2 | 2 | 2 | |
2–4 | 1 | 1 | 1 | Sp | O | M | 6 | 1 | 2 | 2 | 2 | |
2 | 1 | 2 | 2 | Su | P | H | 11 | 1 | 2 | 2 | 2 | |
3 | 1 | 2 | 2 | Su | P | H | 11 | 1 | 2 | 2 | 1 | |
2–4 | 1 | 2 | 2 | Sp | P | M | 11 | 1 | 2 | 2 | 2 | |
2–5 | 1 | 1 | 1 | Sp | P | M | 6 | 1 | 1 | 2 | 3 | |
2–4 | 1 | 2 | 2 | Su | P | H | 11 | 1 | 2 | 2 | 2 | |
2–4 | 1 | 2 | 2 | Sp | P | H | 11 | 1 | 2 | 2 | 2 | |
3 | 1 | 2 | 2 | Su | P | H | 11 | 1 | 3 | 2 | 1 | |
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2–3 | 1 | 2 | 2 | Su | S | H | 11 | 2 | 2 | 2 | 1 | |
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3 | 3 | 1 | 1 | Su | S | H | 35 | 2 | 3 | 2 | 1 | |
5–6 | 3 | 1 | 1 | Su | S | H | 35 | 2 | 3 | 2 | 2 |
The NMDS analysis was conducted using PC–ORD Ver. 5 (
Nine Ohio HUC6 drainages, number of unique locations, species richness and 16 environmental variables used in NMDS analysis. Pre-European settlement vegetation is percentage cover. PEMM = Portage Escarpment Mesophytic Forest, Forest_UL =Upland forest, Forest_MX = Mixed deciduous/coniferous forest, WL_NW = nonwooded wetland, WL_W = wooded wetland, RR_Mean = Relief Ratio mean.
% Pre-European Settlement Vegetation | Elevation (m) | Relief Ratio | ||||||||||||||||
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USGS<br/> HUC6s | Sites | Species Richness | DRAIN_km2X1000 | PEMM | PRAIRIE | FOREST_UL | FOREST_MX | SAVANNA | WL_NW | WL_W | ELEV_MEAN | ELEV_STD | ELEV_MAX | ELEV_MIN | ELEV_MEDIAN | RR_MEAN | RR_MEDIAN | SLOPE % |
WL_Erie | 95 | 25 | 23.31 | 0.0 | 2.3 | 59.3 | 0.7 | 2.0 | 1.6 | 34.1 | 239 | 35.8 | 411 | 147 | 233 | 0.35 | 0.33 | 0.8 |
SE_LErie | 133 | 65 | 8.25 | 1.2 | 0.1 | 68.8 | 24.0 | 0.0 | 1.0 | 4.9 | 285 | 53.9 | 420 | 173 | 285 | 0.45 | 0.45 | 2.2 |
UOH_Bvr | 50 | 44 | 8.62 | 0.0 | 0.0 | 80.2 | 17.2 | 0.0 | 0.2 | 2.4 | 330 | 38.6 | 436 | 182 | 336 | 0.58 | 0.61 | 6.2 |
UOH_LKan | 80 | 48 | 7.93 | 0.0 | 0.2 | 80.7 | 15.9 | 0.0 | 0.3 | 2.9 | 263 | 45.8 | 433 | 151 | 261 | 0.40 | 0.39 | 10.3 |
Musk | 116 | 56 | 20.85 | 0.0 | 0.0 | 76.2 | 20.6 | 0.0 | 0.4 | 2.8 | 316 | 43.0 | 460 | 174 | 317 | 0.50 | 0.50 | 6.8 |
Scioto | 201 | 71 | 16.87 | 0.0 | 3.9 | 69.9 | 12.1 | 0.0 | 0.3 | 13.7 | 283 | 43.2 | 454 | 142 | 289 | 0.45 | 0.47 | 3.9 |
GrMiami | 109 | 34 | 10.70 | 0.0 | 1.3 | 79.4 | 5.1 | 0.0 | 0.4 | 13.8 | 299 | 44.7 | 469 | 138 | 306 | 0.49 | 0.51 | 1.9 |
MOH_Rac | 39 | 35 | 5.06 | 0.0 | 0.0 | 84.9 | 10.0 | 0.0 | 0.0 | 5.1 | 233 | 34.1 | 363 | 136 | 234 | 0.43 | 0.43 | 9.8 |
MOH_LMia | 119 | 57 | 9.37 | 0.0 | 1.8 | 60.8 | 16.4 | 0.0 | 0.0 | 21.0 | 269 | 45.5 | 404 | 131 | 275 | 0.50 | 0.53 | 4.6 |
A total of 4,051 database records accounting for 32,487 specimens were accumulated for this project (
Species richness estimators predicted slightly higher values (
Ohio
Singleton and doubleton species richness.
Species richness of Ohio
The stonefly fauna of Ohio was represented by all nine families known to inhabit the North American continent (
Species richness of Ohio
Stream size is often an important determinant of stonefly communities. This dataset demonstrated that most species inhabited a range of stream sizes (
The vast majority of stonefly species inhabiting Ohio have single year life cycles; only 17 (16.7%) had multiyear life cycles (
The number of months of nymphal growth had the largest number of trait states of all species trait categories. There were 9 species with an exceedingly short growth period of four months. These included all
Species traits distributions for the Ohio stonefly assemblage. Traits from Table 2.
Volt. | Devel. | Diap. | Dispers. | Feed. | Mobil. | Grow. | Respir. | Size (mm) | Synch. (mo.) | Therm. | |||||||||||
1 | 85 | 1 | 57 | 1 | 56 | W | 25 | O | 1 | L | 15 | 4 | 8 | 1 | 56 | <9 | 46 | >1 | 36 | 1 | 19 |
2 | 15 | 2 | 45 | 2 | 46 | Sp | 28 | P | 56 | M | 42 | 6 | 42 | 2 | 46 | 9–16 | 35 | <1 | 66 | 2 | 67 |
3 | 2 | Su | 49 | S | 45 | H | 45 | 9 | 4 | >16 | 21 | 3 | 16 | ||||||||
11 | 31 | ||||||||||||||||||||
23 | 15 | ||||||||||||||||||||
35 | 2 |
More research on the feeding of
Female mobility is an important trait that confers ability to colonize and recolonize after local extinction. The vast majority of species exhibited medium to high female mobility. Low female dispersal ability was exhibited by 14.6% of Ohio species. Low mobility is a complex character state that is exhibited mostly by winter emerging species that emigrate by crawling, skating (wings held up to breeze and skating on tarsi), or floating on logs or ice floes.
The presence or absence of gills is often thought of as indicative of a species’ ability to tolerate warmer waters and lower oxygen concentrations. No formal analysis has been conducted of the association of gilled stoneflies with water temperature preference, but informal studies in Europe indicate that many gilled species inhabit mountainous areas with cold and cool water temperatures (M. Tierno de Figueroa pers. comm.). In eastern North America there are over 50 species of perlids (
The majority of Ohio species (54.9%) did not have gills (some
Size at maturity is a trait that has direct bearing on risk to survival. Larger species are usually longer lived and exposed to risks for a longer period of time and may make more attractive prey items than some smaller species. Small species (<9 mm total length) were much more frequent in the list than large species (>16 mm). Smaller species are more likely to have short growth periods and diapause. They are also more likely to have fast cycles and disperse in the winter and spring than larger species.
Species with synchronous (<1 month) emergence periods were more frequent in Ohio than asynchronous (>1 month) species. Winter emerging species tend to have less synchronous emergence periods due to fluctuating winter temperatures from freezing and thawing. Those that emerge in spring and summer have sharper seasonal cues that lead to nymphal development being more synchronous, leading to emergence of adults over a shorter period of time. Spring and summer emerging species were three times more frequent in Ohio than winter-emerging ones; hence, the great disparity in synchronous over asynchronous emergence is easily understood.
Thermal requirements are not well understood in stoneflies or other aquatic insects and the terms coldwater, coolwater, and warmwater are relative when it comes to defining a temperature requirement. Trait state assignment here is based more on professional experience than for any other set of traits. Coldwater species contribute only 18.6% to the total number of species found in OH. Most of Ohio is or was heavily wooded, a feature usually related to coolwater conditions. This is by far the most frequent thermal tolerance state for stoneflies in Ohio. Another 15.7% of species can truly be categorized as warmwater species. This would include several perlid species.
Randomization tests in the NMDS analysis recommended a three dimensional solution. An overall stress value for the three dimensional analysis was low at 1.53. A plot of Axis 1 vs Axis 3 separated the communities of Ohio stoneflies best (
Non–parametric Multi–Dimensional Scaling of Ohio
Ohio is a state that has its eastern flank in the Allegheny Plateau, an extension of the Appalachian Mountains. Ohio’s western flank is mostly till plain resulting from the Wisconsinan glaciation. The stonefly fauna found in the state is a mixture of species requiring cooler waters and deep forest and those that have evolved with warmwater streams and even intermittency of flow. The number of species occurring in Ohio is indicative of being between these two extremes. Ohio supports at least 102 species, maybe as high as 119 (e.g., Chao 2 upper 95 percentile) (
Comparison of Ohio
Sørensen Index of Similarity between
IL | IN | MI | OH | ON | WI | PA | WV | KY | |
IL | 0 | ||||||||
IN | 0.814 | 0 | |||||||
MI | 0.444 | 0.417 | 0 | ||||||
OH | 0.659 | 0.743 | 0.415 | 0 | |||||
ON | 0.416 | 0.429 | 0.656 | 0.551 | 0 | ||||
WI | 0.569 | 0.523 | 0.744 | 0.548 | 0.686 | 0 | |||
PA | 0.425 | 0.470 | 0.394 | 0.629 | 0.535 | 0.512 | 0 | ||
WV | 0.440 | 0.485 | 0.349 | 0.628 | 0.467 | 0.461 | 0.826 | 0 | |
KY | 0.588 | 0.684 | 0.354 | 0.749 | 0.437 | 0.474 | 0.648 | 0.721 | 0 |
Ohio had 18 species that were collected from just one or two locations, 17.6% of all species found. A discussion for a limited number of these species is presented below to identify for state, federal, and non–profit conservation organizations that these species, and the streams in which they reside, should be considered for protected status. Some streams are already in the public trust.
Several streams across the state are exceedingly rich in species and have been well sampled. A tributary of the East Branch of the Chagrin River in Stebbins Gulch of Holden Arboretum (Geauga Co.) produced 28 species including several coldwater species. A tributary of the East Fork Queer Creek at Ash Cave (Hocking Co.) has produced 23 species. The Olentangy River near Columbus (Franklin Co.) has produced 17 species historically. Upstream of the city near Highbank Metropark a diverse assemblage still persists, although it may not hold the full complement of species once found in the river. The Clear Fork of the Mohican River within the ravine area of Mohican State Park (Ashland Co.) has produced 14 species and probably still supports most of them. Big Lyons Falls Creek, also in Mohican State Park; a tributary of the North Fork Little Beaver River, 5 km S Negley, in Columbiana Co.; and Mill Creek at Doty Road (Lake Co.) all produced 13 species. Gray’s Run at Lowellville (Mahoning Co.) produced a total of 12 species, many of which were Appalachian coldwater species. Most of these locations are protected by public or private means. These, and several others too numerous to list here, are important for protecting the lotic diversity of aquatic organisms in the Ohio.
HUC6 drainages explain some of the variation in stonefly communities across Ohio (
The data suggested that smaller drainages of the eastern and southern part of the state followed a pattern of increasing richness with drainage area, but that the largest drainage did not (
Some of the most species rich HUC6 assemblages were located in a band of upland forest and higher gradient streams that straddled drift plain and unglaciated terrain from Cincinnati to Ashtabula. These assemblages were dominated by widespread species that typically inhabit cool and warmwater streams. There was also a small component of coldwater Appalachian fauna including species of
Two other drainages, the Upper Ohio Little Kanawha and the Middle Ohio Raccoon, had assemblages that were defined by mixed deciduous and coniferous forests and higher slope values in southern, unglaciated Ohio. These are relatively small drainages with short streams, many of which become intermittent in the summer. Consequently, many species with egg or nymphal diapause are found here. This trend also occurs in southern Illinois and Indiana (
The
A large dataset from 18 regional museums, new collecting, and trusted literature sources demonstrated that the
Ohio species are mostly univoltine-fast with egg or nymphal diapause and the largest proportion of them are summer emerging (
It appears that Ohio has been well sampled; species estimators (Chao2 and ICE Mean) suggested that 3 or 4 more species could be found. Given that neighboring Pennsylvania and West Virginia have 142 and 139 species, respectively, it is likely that species shared with them will eventually be found in Ohio (
A great number of species in Ohio were rare, being known from only 1–5 locations (
This paper lays a foundation for planned future work, including natural range modeling of species within the larger framework of the Midwest USA and Canada. This will allow our research team to reconstruct pre-European settlement ranges for most species in the region. We are also focusing considerable effort to use these baseline distributions against which to measure climate related changes in distribution by modifying climate variables in light of predicted CO2 emissions scenarios.
The authors thank several individuals for loan of specimens: Bill Stark (BPSC), Richard Baumann and Shawn Clark (BYU), Ian Smith (CNC), Gregory Dahlem (CINC), Joe Keiper (formerly CMNH), Daniel Summers (FMNH), Gary Parsons (MSUC), Brian Armitage (OBS), Michael Bolton (OEPA), Bob Glotzhober (OHSC), Norm Johnson (OSUC), Arwin Provonsha (PERC), Fred Kirchner (RFKC), Antonia Guidotti (ROM), Jay McPherson (SIUC), and Mark O’Brien (UMC). We also wish to thank Donald W. Webb for his contributions at the beginning of this effort. This work was partially supported by the National Science Foundation DEB 09–18805 ARRA.
Raw data and data matrices. (doi: