Corresponding author: JoVonn G. Hill (
Academic editor: Zhu-Qing He
Hill JG (2022) Revision of
The North American Coastal Plain was recently designated as the world’s 36th global biodiversity hotspot based on the high levels of biodiversity and endemism of vascular plants and habitat loss greater than 70% in the region (
Habitus drawings of
Geographic distribution of
Most specimens examined in this study were collected by Dr. Theodore Hubble and Dr. Irving Cantrall of the
In order to conduct a thorough study of the genus, the male genitalia, which are typically concealed within the terminalia, were dissected and examined. Habitus and internal genitalia photographs were taken with a Leica Z16 stereoscope equipped with a Leica DFC420 camera at different stages during dissection. Images were automontaged with the Leica Application Suite. For scanning electron micrographs, specimens were mounted on stubs with silver paste and coated with 30 nm of platinum, then imaged with a JEOL–JSM65600F SEM. Measurements were made with a reticle mounted inside a Leica MZ12.5 stereomicroscope in the following ways:
Body Length — Dorsally from the fastigium vertices to the distal end of the genicular lobe of caudal femur in a parallel plane with the abdomen.
Pronotum length — Dorsally, along the median carina.
Cercus Length — Laterally, maximum possible measurement of the left cercus.
Cercus Basal Width — Laterally, along the point of attachment from the dorsal to ventral margin.
Mid Cercus Width — Laterally, at the mid-length of the left cercus.
Cercus Apex Width — Laterally, along the distal end.
Subgential Plate Tubercule Length — Laterally, from the base to the apex.
Subgential Plate Tubercule Width — Posteriorly, at the widest point.
Based on male morphology and distribution,
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Small size (11–22 mm).
Body linear in shape (Fig.
Appearing apterous with wings reduced to a minute, vestigial scale.
Body brownish-green or bronze with a black stripe running from behind the eye to near the end of the abdomen (Fig.
Hind tibia and tarsi dull green.
Large size (17–28 mm).
Body somewhat elongate.
Wings developed into small linear pads.
Body green or brown with white and black striping.
Hind tibia blue, tarsi pink.
Size variable – small (10–20 mm) to larger (18–28 mm).
Body more robust.
Brachypterous but wings obvious.
Body green to bronze with a metallic luster; black postocular stripe.
Hind tibia and tarsi black or red depending on the species.
Larger (18–28 mm).
Body robust.
Wings reduced to small and slender pads.
Body green.
Hind tibia and tarsi pink.
Larger (16–29 mm).
Body robust.
Appearing apterous with wings reduced to a minute, vestigial scale.
Body green with black, white, or orange striping.
Hind tibia blue, tarsi pink.
Lateral lobes of subgenital plate expanded dorsally; tubercule longer than wide (Figs
Cerci generally falcate (Figs
Ventral valves of aedeagus more translucent; flattened (Figs
Lateral lobes of subgenital plate not expanded; tubercule approximately as long as wide (Figs
Cerci triangular (Figs
Ventral valves of aedeagus more opaque; cylindrical (Figs
1 | Male cerci generally falcate with the apex nearly as wide as long at their bases (Figs |
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– | Male cerci generally triangular with the apex much narrower than the base (Figs |
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2 | Tubercule of subgenital plate broader (Figs |
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– | Tubercule of subgenital plate narrower (Figs |
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3 | Dorsal valves of the aedeagus shorter than the ventral valves (Figs |
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– | Dorsal valves of the aedeagus equal in length or nearly so to that of the ventral valves; not peninsular Florida |
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4 | Dorsal valves of the male aedeagus rounded apically and expanded laterally such that they appear lobate (Figs |
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– | Dorsal valves of the aedeagus truncated or slightly angular apically and not expanded laterally, usually parallel sided (Figs |
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5 | Dorsal valves of the aedeagus more truncated apically and not twisted (Figs |
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– | Dorsal valves of the aedeagus more angled apically (Fig. |
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Species of small size (M: 11.8–17 mm, F: 17.5–22.2 mm). Body somewhat gracile and subcylindrical.
The dorsal valves are translucent to semi-translucent lobes that are flat, truncate, shortened to elongate depending on the species. The ventral valves are opaque and more strongly sclerotized than the dorsal valves, caudally projecting cylindrical lobes of various shapes depending on the species (Figs
Overall dull greenish brown to yellow, sometimes with bronze highlights. Antenna yellowish basally, remainder ferruginous. Antennal crescent complete. Head, thorax, and abdomen pale yellow, infuscated dorsally, especially along the midline. A lateral, well-defined, piceous, post-ocular stripe extends from the caudal margin of the eye through the thorax and towards the end of the abdomen; lateral area of head and thorax below post-ocular stripe creamy-yellow. Hind femora luteous. Hind tibia, pale dull green, often dulled basally; with black or black tipped spines (Figs
Naked leapers.
Most easily differentiated from the other species in the group based on the shape of the male cerci, which in
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Florida, Walton County, Defuniak Springs, 30 August 1915, Rehn and Hebard (1♂).
Dorsal and lateral SEM of
Partial silhouettes of
Caudal view of
Mobile Bay (Baldwin County, AL) east through the panhandle of Florida to Bay and Jackson counties (Fig.
Differs from
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1♂, FLA., Bay Co., Ecofina Creek WMA,
Ecofina Creek WMA,
Occurs in a narrow portion of the eastern Florida panhandle. At present, it is known only from Bay, Calhoun, Franklin, Gulf, and Liberty counties (Fig.
From the Latin
This species can be found in much drier conditions that other members of the genus. At the type locality this species inhabited fine grasses in a sandy upland with
Differs from other species in the group based the shape of the internal male genitalia. In dorsal view, the dorsal valves are lightly sclerotized and semi-translucent, have apices that are rounded to sub-truncate, and shorter than the ventral valves (Fig.
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Florida [Duval Co.,] Jacksonville. Aug. [18]85.
Peninsular Florida from the northeast boarder with Georgia along the Atlantic Ocean west to eastern bank of the Suwannee River and south to the southern borders of De Soto, Highlands, and Okeechobee Counties (Fig.
Found in a variety of grassland situations from seasonal ponds (Fig.
Differing from other species in the group based on the shape of the internal male genitalia (Fig.
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6 mi S Old Town, 29.5156769, -83.0002496, 28 Sept. 2017, J.G. Hill, Collected from roadside sandhill and ditch (1♂). Deposited in the Mississippi Entomological Museum.
Same data as type (2♂, 2♀).
Florida: Dixie Co., 4 mi N Shamrock, 14 August 1947, T.H. Hubbell (1♂); 6 mi S. Steinhatchee R[iver], 5 August 1925, T.H. Hubbell (4♂); Cross City, 21 November 1925, T.H. Hubbell (1♂). Jefferson Co., 0.4 mi N Lamont, 16 August, 1947, T.H. Hubbell (5♂); 0.4 mi NE Fanlew, 16 August 1947, T.H. Hubbell (1♂); 0.7 mi N Jct. US 90 and Fla 257, 17 August 1947, T.H. Hubbell (1♂); 0.9 mi E Thomas City, 16 August, 1947, T.H. Hubbell (1♂); 4.4 mi NE Fanlew, 16 August 1947, T.H. Hubbell (1♂); 4.6 mi E Monticello, 17 August 1947 (1♂); near Covington, 31 Oct, 1942, T.H. Hubbell (5♂); Lloyd, 20 August 1938 (2♂, 4♀). Lafayette Co., 2 mi W Taylor County Line, 9 June 1941, Friauf and Hubbell (1♂); 12 mi W Mayo, 9 November 1941, Friauf & Hubbell (1♂). Leon Co., Chaires, 4 August1925, T.H. Hubbell (4♂). Taylor Co., 4.7 mi N Salem, 7 October 1945, T.H. Hubbell (1♂); Boyd, 15 October 1942, T.H. Hubbell (1♂); Perry, 5 August 1925, T.H. Hubbell (2♂); Hampton Springs, 31 October 1947, T.H. Hubbell (2♂). Madison Co. 2 mi E Aucilla River on US 90, 17 August 1947, T.H. Hubbell (1♂); 1.7 mi N Shady Creek, 16 Sept. 1942, T.H. Hubbell (1♂). Wakulla Co., 1.5 mi NW St. Marks, 15 August 1947, T.H. Hubbell (2♂).
“Big Bend” region of Florida from Leon and Wakulla counties, south through the flatwoods to the western banks of the Suwannee River in Dixie County (Fig.
Flatwoods and grassy sandhills (Fig.
Named in honor of Bettie Mae Page, an iconic American photo model and former resident of Florida, who rose from a background of poverty and abuse to become a symbol of self-expression and body positivity.
Differing from other species in the group based on the shape of the internal male genitalia (Fig.
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GA., Thomas Co., Wade Tract,
Same data as type (6♀).
Found in southern Georgia and north Florida, from Berrien County, GA west to the Chattahoochee River, and south to Liberty and Baker Counties, FL (Figs
Flatwoods and pitcher plant bogs. I observed this species feeding on
Named in honor of the Wade Family who, in 1979, placed an 85-ha tract of old growth longleaf pine savanna into a perpetual conservation easement. Today, the “Wade Tract”, is one of the most important remaining examples of the long leaf pine ecosystem in existence and is also the type locality of this species.
Differing from other species in the group based on the shape of the internal male genitalia (Fig.
Distribution of
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GA., Appling Co. Moody Forest N.A.,
Plant communities at the collection localities of
Same data as type, except BOLD DNA JGH 0066,
All known locations occur on the lower Coastal Plain of Georgia Bulloch County south to Ware and Charlton Counties (Fig.
Flatwoods and pitcher plant bogs.
Named after the state of Georgia, from which this species is apparently endemic.
The four new species of
A complete phylogeny of the North American
Despite being relatively secure in terms of conservation at present,
This publication is a contribution of the Mississippi Agriculture and Forestry Experiment Station and was partially supported by funding from the National Institute of Food and Agriculture, the National Science Foundation OPUS (2043909), the U.S. Fish and Wildlife Service, and Tall Timber Research Station. Some of the specimens used in this study were collected under permits from Florida State Parks, Florida Fish and Wildlife Commission, the U.S. Fish and Wildlife Service, Tall Timbers, The Georgia Department of Natural Resources, and The Nature Conservancy. I thank Irvin Cantrall, Ted Hubbell, and J.J. Friauf for collecting many of the historic specimens used in this study, and Matt Thorn for assisting with collecting some of the modern specimens. Additionally, I thank Steve Orzell for facilitating access to the wonderful natural areas on Avon Park Air Force Base and to Ashley Baker for her assistance producing the figures for this article.