Revision of the Melanoplus Scudderi (Orthoptera: Acrididae: Melanoplinae) Species

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Revision of the Melanoplus scudderi (Orthoptera: Acrididae: Melanoplinae) species

group and a preliminary investigation into the grasshopper fauna of the grasslands

of the southeastern United States

JoVonn Grady Hill

A Dissertation Submitted to the Faculty of Mississippi State University in Partial Fulfillment of the Requirements for a degree in Doctor of Philosophy in Entomology in the Department of Biochemistry, Molecular Biology, Entomology, and Plant Pathology

Mississippi State, Mississippi

May 2015

JoVonn Grady Hill

2015

Revision of the Melanoplus scudderi (Orthoptera: Acrididae: Melanoplinae) species

group and a preliminary investigation into the grasshopper fauna of the grasslands

of the southeastern United States

JoVonn Grady Hill

Approved:

______Richard L. Brown (Major Professor)

______Gerald T. Baker (Committee Member)

______Matt Eitel Dakin (Committee Member)

______Daniel Otte (Committee Member)

______Lisa Wallace (Committee Member)

______Michael A. Caprio (Graduate Coordinator)

______George M. Hopper Dean College of Agriculture and Life Science

Name: JoVonn Grady Hill

Date of Degree: May 8, 2015

Institution: Mississippi State University

Major Field: Entomology

Major Professor: Dr. Richard Brown

Title of Study: Revision of the Melanoplus scudderi (Orthoptera: Acrididae: Melanoplinae) species group and a preliminary investigation into the grasshopper fauna of the grasslands of the southeastern United States

Pages in Study: 213

Candidate for Doctor of Philosophy

The Melanoplus scudderi species group, when erected by Blatchley, contained six

species. The revision of the group presented here describes 21 new species and

establishes the Melanoplus carnegiei group, all of which are associated with grasslands in

the Southeast. Analysis of DNA barcoding data, suggest that the technique does not

adequately delineate species based on the morphological concepts presented here, but did

produce three broad mitochondrial DNA clades that correspond to the Apalachicola and

Mississippi River discontinuity phylogeographic pattern displayed by several other

organisms in the Southeastern United States. This pattern is attributed to vicariant events

resulting from Pleistocene glaciation. A hypothetical evolutionary history of the Scudderi

and Carnegiei groups is presented.

Natural grasslands historically covered a significant portion of the southeastern

United States. These grasslands contribute significantly to the biodiversity of the region

with many endemic or rare species inhabiting them. Grasshoppers are important

components of temperate grassland ecosystems, but no comprehensive review of the

grasshopper fauna of southeastern grasslands had previously been conducted. Here a review of the regional fauna was conducted. In total, 211 grasshopper species have been documented in the Southeast, with 111 species (53%) being endemic to natural grasslands. Of particular note, 74 species (34% of the regional fauna) are endemic to the longleaf pine ecosystem. Many of the non-endemic grasshopper species inhabiting grasslands in the region are disjunct from larger populations west of the Mississippi

River, presenting interesting biogeographic distributions. Additionally, a cluster analysis of grasshopper community composition from several types of grasslands in the region supports the faunal distinctiveness of the different types grasslands in the region.

ACKNOWLEDGEMENTS

I thank Dr. Richard Brown for serving as my major professor and providing insightful taxonomic assistance, Dr. Dan Otte for graciously handing this project to me,

Dr. Lisa Wallace for her assistance with the molecular aspects of the study, Dr. Matt

Dakin and Dr. Gerry Baker for their guidance as committee members and comments on this manuscript. Additionally, I thank the late Dr. Irving Cantrall who collected many of the specimens that this revision was based on. Specimens collected by JGH were obtained under permits from the Arkansas Natural Heritage Commission, Florida State

Parks, Georgia Department of Natural Resources, Mississippi Department of Wildlife

Fisheries and Parks, North Carolina Division of Parks and Recreation, The National Park

Service, The Nature Conservancy, South Carolina State Parks, Tennessee Division of

Natural Areas, and Texas Parks and Wildlife. I thank Mark O’Brien of the University of

Michigan Museum of Zoology, Jason Weintraub of the Academy of Natural Sciences of

Drexel University, Steven Krauth, of the Wisconsin Insect Research Collection, Matt

Dakin of Auburn University Museum of Natural History, Paul Skelley of the Florida

State Collection of Arthropods, Bob Blinn of the North Carolina State Insect Collection for preparing loans of material examined in this study. Additionally, I thank Justin Luke,

Jordan Gesell, and Charlotte Chesenut for assisting with specimen preparation and locality data entry. This research received financial support from the Orthopterists’

Society, the William H. Cross Expedition Endowment in the Mississippi State University

Development Foundation, the Georgia Department of Natural Resources, and the

Mississippi Agricultural and Forestry Experiment Station. This research was supported by the National Institute of Food and Agriculture, under Project No. MIS-012040.

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TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... ii

LIST OF TABLES ...... xii

LIST OF FIGURES ...... xiii

CHAPTER

I. INTRODUCTION ...... 1

Introduction ...... 1 Background on the genus Melanoplus ...... 2 Literature Cited ...... 4

II. REVISION OF THE MELANOPLUS SCUDDERI (ORTHOPTERA:ACRIDIDADE) SPECIES GROUP ...... 5

Introduction ...... 5 Methods...... 7 Morphological study ...... 7 Species delimitation ...... 9 Molecular techniques ...... 9 Results ...... 12 Comparison of the groups ...... 12 Diagnosis of the Melanoplus scudderi species group ...... 13 Structural Characters ...... 13 Phallic Structures ...... 14 Coloration ...... 14 Species accounts ...... 15 Melanoplus scudderi Uhler (Figs. 1A-J, 2A-B, 3, 4A-B) ...... 15 Male Measurements (mm) ...... 15 Female Measurements (mm)...... 15 Diagnosis...... 15 Lectotype here designated...... 16 Other specimens examined ...... 16 Distribution ...... 32 Habitat ...... 32 Remarks ...... 32

Melanoplus folkertsi new species (Figs. 5A-J, 8) ...... 37 Male Measurements (mm) ...... 37 Female Measurements (mm)...... 37 Diagnosis...... 38 Holotype ...... 38 Paratype...... 38 Other specimens examined ...... 38 Etymology ...... 39 Distribution ...... 40 Habitat ...... 40 Melanoplus relictus new species (Figs. 6A-J, 8) ...... 42 Male Measurements (mm) ...... 42 Female Measurements (mm)...... 42 Diagnosis...... 42 Holotype ...... 42 Paratypes ...... 42 Other specimens examined ...... 43 Etymology ...... 43 Distribution ...... 43 Habitat ...... 43 Melanoplus coreyi new species (Fig., 7A-J, 8) ...... 45 Male Measurements (mm) ...... 45 Female Measurements (mm)...... 45 Diagnosis...... 45 Holotype ...... 45 Paratypes ...... 46 Other specimens examined ...... 46 Etymology ...... 46 Distribution ...... 46 Habitat ...... 46 Melanoplus muscogee new species (Fig. 8, 9A-J) ...... 49 Male Measurements (mm) ...... 49 Female Measurements (mm)...... 49 Diagnosis...... 49 Holotype ...... 49 Paratypes ...... 50 Other specimens examined ...... 50 Etymology ...... 50 Distribution ...... 50 Habitat ...... 50 Melanoplus quercicola (Hebard) (Fig. 8, 10A-J) ...... 52 Male Measurements (mm) ...... 53 Female Measurements (mm)...... 53 Diagnosis...... 53 Holotype ...... 55

Specimens examined ...... 55 Distribution ...... 55 Habitat ...... 56 Remarks ...... 56 Melanoplus davisi (Hebard) (Fig. 8, 11A-J) ...... 56 Male Measurements (mm) ...... 56 Female Measurements (mm)...... 56 Diagnosis...... 56 Holotype ...... 57 Other Specimens Examined ...... 57 Distribution ...... 59 Habitat ...... 59 Remarks ...... 59 Melanoplus mississippi new species (Figs. 12A-J, 13)...... 59 Measurements (mm) ...... 59 Female Measurements (mm)...... 59 Diagnosis...... 60 Holotype ...... 60 Paratypes ...... 60 Other specimens examined ...... 60 Etymology ...... 61 Distribution ...... 62 Habitat ...... 62 Remarks ...... 62 Melanoplus arkansas new species (Fig. 14A-J, 17) ...... 64 Male Measurements (mm) ...... 64 Female Measurements (mm)...... 66 Diagnosis...... 66 Holotype ...... 66 Paratypes ...... 66 Other specimens examined ...... 66 Etymology ...... 67 Distribution ...... 67 Melanoplus ouachita new species (15A-J, 17) ...... 67 Male Measurements ...... 67 Female Measurements (mm)...... 68 Diagnosis...... 68 Holotype ...... 68 Paratypes ...... 68 Other specimens examined ...... 69 Etymology ...... 70 Distribution ...... 71 Habitat ...... 71 Melanoplus cohni new species (Figures 16A-J, 17) ...... 71 Male Measurements ...... 71

Female Measurements (mm)...... 72 Diagnosis...... 72 Holotype ...... 72 Paratypes ...... 72 Other specimens examined ...... 73 Etymology ...... 73 Distribution ...... 73 Habitat ...... 73 Melanoplus seltzerae new species (Figs 17, 18A-J) ...... 75 Male Measurements ...... 75 Female Measurements (mm)...... 75 Diagnosis...... 76 Holotype ...... 76 Paratypes ...... 76 Etymology ...... 77 Distribution ...... 77 Habitat ...... 77 Melanoplus baronei new species (17, 19A-J) ...... 79 Male Measurements ...... 79 Female Measurements (mm)...... 79 Diagnosis...... 79 Holotype ...... 81 Paratypes ...... 81 Other specimens examined ...... 81 Etymology ...... 81 Distribution ...... 81 Habitat ...... 81 Melanoplus latus Morse New Status (Figs. 20A-J, 21A-D, 22J) ...... 81 Male Measurements (mm) ...... 82 Female Measurements (mm)...... 82 Female Measurements (mm)...... 82 Diagnosis...... 84 Lectotype...... 84 Other specimens examined ...... 85 Distribution ...... 86 Habitat ...... 86 Melanoplus decurvus new species (Figs. 17, 23A-J) ...... 87 Male Measurements (mm) ...... 87 Female Measurements (mm)...... 88 Diagnosis...... 88 Holotype ...... 88 Paratypes ...... 88 Other specimens examined ...... 88 Etymology ...... 89 Distribution ...... 89

vii

Habitat ...... 89 Melanoplus ozarkensis new species (Fig. 24A-J, 23) ...... 91 Male Measurements ...... 91 Female Measurements (mm)...... 91 Diagnosis...... 91 Holotype ...... 91 Paratypes ...... 91 Other specimens examined ...... 93 Etymology ...... 95 Distribution ...... 95 Habitat ...... 95 Melanoplus francoisensis new species (Figs. 25, 26A-J) ...... 96 Male Measurements (mm) ...... 96 Female Measurements (mm)...... 97 Diagnosis...... 97 Holotype ...... 97 Paratypes ...... 97 Other specimens examined ...... 98 Etymology ...... 98 Distribution ...... 98 Habitat ...... 98 Melanoplus texarkana new species (Figs. 27A-J, 28) ...... 100 Male Measurements ...... 100 Female Measurements ...... 100 Diagnosis...... 100 Holotype ...... 100 Paratype...... 101 Other specimens examined ...... 101 Etymology ...... 102 Distribution ...... 102 Habitat ...... 103 Melanoplus taurus new species (Figs. 29A-J, 30) ...... 105 Male measurements (mm) ...... 105 Female measurements (mm) ...... 106 Diagnosis...... 106 Holotype ...... 106 Paratypes ...... 108 Other specimens examined ...... 108 Etymology ...... 108 Distribution ...... 108 Habitat ...... 108 Melanoplus optimus new species (Fig. 30, 31A-J) ...... 109 Male measurements ...... 109 Female measurements (mm) ...... 110 Diagnosis...... 110

viii

Holotype ...... 110 Paratypes ...... 110 Other specimens examined ...... 110 Etymology ...... 111 Distribution ...... 111 Habitat ...... 111 Melanoplus ottei new species (Fig. 30, 32A-J) ...... 113 Male Measurements ...... 113 Female Measurements (mm)...... 113 Diagnosis...... 113 Holotype ...... 113 Paratypes ...... 113 Other specimens examined ...... 114 Etymology ...... 114 Distribution ...... 116 Habitat ...... 116 Melanoplus irwinorum new species (Fig. 30, 33A-J) ...... 116 Male Measurements (mm) ...... 116 Female Measurements (mm)...... 116 Diagnosis...... 116 Holotype ...... 117 Paratypes ...... 117 Etymology ...... 117 Distribution ...... 117 Habitat ...... 117 Melanoplus texensis Hart New Status (Fig. 30, 34A-J) ...... 119 Male Measurements (mm) ...... 119 Female Measurements (mm)...... 119 Diagnosis...... 119 Holotype ...... 120 Other specimens examined ...... 120 Distribution ...... 120 Habitat ...... 120 Melanoplus dawsoni (Scudder) Incertae sedis ...... 121 The Carnegiei group ...... 123 Diagnosis of species in the Melanoplus carnegiei Group ...... 123 Phallic Structures ...... 124 Coloration ...... 124 Melanoplus carnegiei Morse (Figs. 35A-J, 38) ...... 125 Male Measurements (mm) ...... 125 Female Measurements (mm)...... 127 Diagnosis...... 127 Holotype ...... 127 Other specimens examined ...... 127 Distribution ...... 129

Habitat ...... 129 Melanoplus acidocercus Hebard (Figures 36A-J, 38) ...... 129 Male Measurements (mm) ...... 130 Female Measurements (mm)...... 130 Diagnosis...... 130 Holotype ...... 130 Other specimens examined ...... 131 Distribution ...... 131 Habitat ...... 131 Melanoplus chattahoochee new species (Fig. 37A-J, 38) ...... 133 Male Measurements (mm) ...... 133 Female Measurements (mm)...... 133 Diagnosis...... 133 Holotype ...... 134 Paratypes ...... 134 Other specimens examined ...... 134 Etymology ...... 136 Distribution ...... 136 Melanoplus savannah new species (Figs. 38, 39A-J) ...... 137 Male Measurements (mm) ...... 137 Female Measurements (mm)...... 137 Diagnosis...... 137 Holotype ...... 138 Paratypes ...... 138 Other specimens examined ...... 138 Etymology ...... 138 Distribution ...... 138 Habitat ...... 140 Melanoplus carolinensis new species (Fig. 38, 40A-J) ...... 140 Measurements (mm) ...... 140 Diagnosis...... 140 Holotype ...... 141 Etymology ...... 141 Distribution ...... 141 Habitat ...... 141 Molecular Analysis ...... 143 Discussion ...... 146 Biology ...... 147 Molecular Discussion ...... 149 Eastern Clade ...... 151 Central Clade ...... 152 Western Clade ...... 153 Endemism in the Interior Highlands ...... 153 Events that may have led to speciation of the Scudderi and Carnegiei groups ...... 154

The broad distribution of Melanoplus scudderi ...... 157 Conservation ...... 158 List of Groups and Species ...... 159 Scudderi group ...... 159 Carnegiei group ...... 160 Literature Cited ...... 161

III. THE GRASSHOPPER FAUNA OF SOUTHEASTERN GRASSLANDS: A PRELIMINARY INVESTIGATION ...... 166

Introduction ...... 166 The Southeastern Fauna ...... 170 Affinities With Western Grasslands ...... 175 Grasshopper Composition of Grasslands ...... 180 Calcareous Grasslands and Glades ...... 182 Longeleaf Pine Ecosystem (Fall Line Sandhills, Ohoopee Dunes, Florida Sandhill/Scrub) ...... 184 Species shared between Prairies and Sandhills ...... 184 Cedar Hardwood Forests ...... 185 Southern Appalachian Balds ...... 185 Marshes ...... 186 Conservation/Management ...... 186 Conclusion ...... 189 Literature Cited ...... 199

APPENDIX

A. KEY TO THE SPECIES OF THE MELANOPLUS SCUDDERI AND CARNEGIEI SPEICE GROUPS ...... 203

Key to the Scudderi and Carnegiei species groups ...... 204 Key to species of the Melanoplus scudderi group based on male genitalia and distribution...... 204 Key to species of the Melanoplus carnegiei group ...... 212

LIST OF TABLES

1 Taxa and associated collecting data for specimens used in molecular analysis...... 10

2 Estimates of Inter and Intraclade Evolutionary Divergence over sequence pairs...... 144

3 Proportion of North American and southeastern grasshoppers that occur in southeastern grasslands, are endemic to the Southeast, are endemic to natural grasslands in the Southeast, and are longleaf endemics...... 172

4 List of grasshopper species that occur in the southeastern United States...... 190

5 Grassland-specific grasshopper taxa with notes on their distribution pattern and habitat preferences...... 197

xii

LIST OF FIGURES

1 Melanoplus scudderi...... 34

2 Variant of the male genitalia of M. scudderi found east of the Appalachian Mountains...... 35

3 Locality of specimens of Melanoplus scudderi that were examined in this study...... 36

4 Long-winged forms of Melanoplus scudderi (A-B) and Melanoplus Mississippi (C)...... 37

5 Melanoplus folkertsi...... 41

6 Melanoplus relictus...... 44

7 Melanoplus coreyi...... 48

8 Distribution of M. folkertsi, M. muscogee, M. coreyi, M. relictus, M. quercicola, and M. davisi...... 51

9 Melanoplus muscogee...... 52

10 Melanoplus quercicola...... 54

11 Melanoplus davisi...... 58

12 Melanoplus mississippi...... 63

13 Distribution of Melanoplus mississippi...... 64

14 Melanoplus arkansas...... 65

15 Melanoplus ouachita...... 70

16 Melanoplus cohni...... 74

17 Distribution of M. arkansas, M. ouachita, M. cohni, M. seltzerae, M. baronei, and M. decurvus...... 75

18 Melanoplus seltzerae...... 78 xiii

19 Melanoplus baronei...... 80

20 Melanoplus latus...... 83

21 Variation in the genitalia of M. latus...... 85

22 Distribution of Melanoplus latus...... 87

23 Melanoplus decurvus...... 90

24 Melanoplus ozarkensis...... 92

25 Distribution of M. ozarkensis, and M. francoisensis...... 96

26 Melanoplus francoisensis...... 99

27 Melanoplus texarkana...... 104

28 Distribution of Melanoplus texarkana...... 105

29 Melanoplus tarus...... 107

30 Distribution of Melanoplus optimus, M. latus, M. texarkana, M. texensis, M. irwinorum, M. ottei, and M. tarus...... 109

31 Melanoplus optimus...... 112

32 Melanoplus ottei...... 115

33 Melanoplus iriwinorum...... 118

34 Melanoplus texensis...... 122

35 Melanoplus carnegiei...... 126

36 Melanoplus acidocercus...... 132

37 Melanoplus chattahoochee...... 135

38 Distribution of Melanoplus acidocercus, M. carnegiei, M. carolina, M. chattahoochee, and M. savannah...... 136

39 Melanoplus savannah...... 139

40 Melanoplus carolinensis...... 142

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41 Molecular phylogentetic analysis of species in the Melanoplus scudderi and carnegiei groups by maximum likelihood using Jukes- Cantor model...... 145

42 Distribution of samples from each clade showing the Apalachicola- Chattahoochee River /Mississippi River discontinuity...... 146

43 Landscape photographs of various Southeastern grasslands...... 169

44 Restricted distributions of some flightless grasshoppers in the Southeast...... 174

45 Members of the Melanoplus scudderi group (s.l.) endemic to sandhills along the Fall Line...... 175

46 Some spur-throat grasshopper species that inhabit southeastern grasslands...... 177

47 Some spur-throat grasshopper species that inhabit southeastern grasslands...... 178

48 Some slant-faced and tooth-legged grasshopper species that inhabit southeastern grasslands...... 179

49 Some band-winged grasshopper species that inhabit southeastern grasslands...... 180

50 Cluster analysis results of grasshopper species composition from 15 different grassland types across the southeastern United States and Konza Prairie in Kansas...... 182

CHAPTER I

INTRODUCTION

“.. we continued over these expansive illumined grassy plains, or native fields,

above twenty miles in length, and in width eight or nine …” -Wm. Bartram (1791)

describing the treeless landscape near present-day Montgomery, Alabama

Introduction

It seems obvious that grasshoppers inhabit grasslands and that grasslands would be important to grasshopper diversity. However, in the southeastern United States there has been no explicit study of the importance of grasslands in the region to the grasshopper fauna. This may be due in part to the largely “forgotten” nature of these

grasslands (See Noss 2012). Despite a burst of interest in these systems over the last

several decades and an abundance of evidence, quotes have continued to be made such as

“If we could have flown over the southeastern USA before immigrants from

Europe arrived in large numbers, it would have looked like the Amazon Region of

today – endless stretches of unbroken broadleaf or mixed forest, cut here and

there by rivers and streams. Rarely would we have seen open patches of

habitat….” (Tschinkel 2006).

Indeed, savannahs and prairies in the region historically covered large swaths of land, and were important features of the landscape as attested to by early surveyors notes and traveler’s descriptions through the region (Barone 2005, Bartram 1774).

In this dissertation I investigate whether the grasslands of the southeastern United

States are important to the regional biodiversity of the grasshopper fauna by conducting a

taxonomic revision of the Melanoplus scudderi species group, and then conducting a

review of the regional grassland fauna.

Background on the genus Melanoplus

The Scudderi species group belongs to the subfamily Melanoplinae, which are the

largest North American subfamily of acridid grasshoppers with 79 genera and

approximately 856 species. Over half of the diversity in the subfamily occurs in the genus

Melanoplus with approximately 430 described species (Knowles and Otte 2000, Otte

2012, Otte 2014). Based on molecular clock evidence, most of Melanoplus species are

thought to have originated during a period from the late Pliocene through the Pleistocene.

The large number of species of relatively recent origin indicates a rate of diversification

that rivals other famous radiations such as the African Rift Lake cichlids and Hawaiian

Drosophila (Knowles and Otte 2000).

The majority of North America’s most economically destructive agricultural pest

species are in Melanoplus, including M. bivitattus (Say), M. femurrubrum, M.

sanguinipes (F.), and M. spretus (Walsh), the latter once extremely devastating, but now

possibly extinct. All members of the genus can be identified by having a spine on the

prosternum between the forelegs, the vertex of the head rounded in profile, hyaline hind

wings, lobes of the metasternum wider than long, a pronotum with little sculpture, and a 2

pronotum with rounded lateral margins. Given these similarities, members of the genus can still be quite variable, with some being relatively large, such as Melanoplus. differentialis (Thomas) (28-44 mm in length) or relatively small, such as Melanoplus

pygmaeus Davis (14-23 mm). Coloration, especially that of the hind tibiae, can be

variable between and within Melanoplus species. Wings are always present, but wing

length can be variable with some species being macropterous (long winged), some being

brachytperous (short winged), whereas, others can have variable wing lengths.

Brachypterous species often have local distributions and relatively small geographic

ranges due to the lack of a strong dispersal mechanism

The last major revision Melanoplus was conducted 117 years ago by Samuel

Scudder (Scudder 1897). As a result of the high diversity in the genus, recent revisionary

work has focused on species groups, the most recent of which include the Viridipes group

and the Texanus group (Hilliard 2001, Otte 2004). Otte (2012) described 80 new species

of Melanoplus, from twelve species groups, further demonstrating the spectacular

radiation of the genus; however, no revision of these groups was presented. In the

following chapter, a revision of the Melanoplus scudderi species group is presented.

Literature Cited

Barone, J.A. 2005. Historical presence and distribution of prairies in the Black Belt of Mississippi and Alabama. Castanea 70: 170-183.

Bartram, W. 1791. Travels of William Bartram. F. Harper, ed. Yale University Press (1958 reprint of 1791 original), New Haven, CT. 727 pp.

Hilliard, J.R. 2001. Two new grasshopper species in the Texanus group of the genus Melanoplus (Orthoptera: Acrididae: Melanoplinae) with biological notes on the group. Transactions of the American Entomological Society. 127: 31-68.

Knowles, L.L. and D. Otte 2000. Phylogenetic analysis of montane grasshoppers from the sky islands of western North America (genus Melanoplus). Evolution 54:1337-1348.

Noss, R. 2013. Forgotten Grasslands of the South: Natural History and Conservation. Island Press. Washington. 317 pp.

Otte, D. 2002. Studies of the Melanoplus 1. Review of the viridipes Group (Acrididae: Melanoplinae). Journal of the Orthopterists' Society 11: 91–118.

Otte, D. 2012. Eighty new species of Melanoplus from the United States (Acrididae: Melanoplinae) Transactions of the American Entomological Society 138: 73-167.

Otte, D. 2014. Aptenopedes and two related genera in Florida with a description of fourteen new species (Acrididae: Melanoplinae) Transactions of the American Entomological Society 140: 245-292.

Scudder, S.H. 1897. Revision of the Orthopteran group Melanopli (Acrididae) with special reference to North American forms. Proceedings of the United States National Museum. 20 (1124): 1-421.

Tschinkel W.R. 2006. The Fire Ants. The Belknap Press of Harvard University Press, Cambridge, MA. 723 pp.

CHAPTER II

REVISION OF THE MELANOPLUS SCUDDERI (ORTHOPTERA:ACRIDIDADE)

SPECIES GROUP

Introduction

Members of the Melanoplus scudderi species group are late summer to fall maturing short-winged (brachypterous) grasshoppers that inhabit a wide variety of natural and disturbed habitats across most of the eastern United States. Blatchley (1920) loosely defined the scudderi group and included M. scudderi scudderi (Uhler), Melanoplus

acidocercus Hebard, Melanoplus carnegiei Morse, Melanoplus davisi (Hebard),

Melanoplus dawsoni (Scudder), and Melanoplus quercicola (Hebard) in the group.

These species were included in the group based on the shared characters of the wings

being shorter than or slightly longer than the pronotum with their the dorsal margins

attingent or slightly overlapping and with the tips being subacute to obliquely rounded,

having hind legs with red tibia, the male cerci being wide at the base and tapering to the

apex, and having a low, but distinct carina on the pronotum. Blatchley (1920) only

focused on the taxa of the eastern United States, thus the two western sub-species of M.

scudderi, Melanoplus scudderi texensis Hart and Melanoplus scudderi latus Morse,

should be included in the group as well. Additionally, M. dawsoni, though closely related

to the Scudderi group, likely belongs in its own grouping based on differing phallic and

external characteristics that were not considered by Blatchley, thus this species is not treated here.

The majority of brachypterous grasshopper species inhabiting the eastern United

States often have local distributions and relatively small geographic ranges due to the lack of a strong dispersal mechanism and consequential specialization of a certain habitat type (See Hubbell 1932 for example). Melanoplus s. scudderi is a glaring exception to

this trend. As currently defined, Melanoplus scudderi scudderi has the broadest

distribution of any flightless grasshopper in the eastern United States, occurring from the

Atlantic Ocean to approximately the 100th meridian and southward from the southern

portions of Massachusetts, Michigan, Minnesota and South Dakota to the Gulf of Mexico

(Blatchley 1920). In addition to a wide distribution, M. s. scudderi also has an apparently

wide niche breadth, where the species is capable of inhabiting both natural and disturbed

habitats.

The application of new techniques often results in the recognition of new species

in what was previously considered a single species. In the time since M. scudderi was

described, new characters involving the sclerotized portions of the internal male genitalia

have been found to be diagnostic for identification of species within the genus (Hubbell

1932, Cohn and Cantrall 1974, Otte 2002). Male genitalia in insects (especially in the

Melanoplinae) are posited to be under sexual selection, which can result in rapid reproductive isolation and result in cryptic species with similar external morphology

(Eberhard 1996, Knowles and Otte 2000, Otte 2002). Given the broad distribution and niche breadth of M. s. scudderi, it was suspected that the group, in particular M. s.

scudderi, likely consisted of multiple species. For this reason a study of the male genitalia of the Scudderi group was undertaken.

Methods

Morphological study

A majority of the specimens examined in this study were collected by Dr. Irving

Cantrall of the University of Michigan Museum of Zoology (UMMZ), who made

extensive collections of the Scudderi group in 1954 and 1958 with hopes to carry out a

study of the group. However, no such revision was ever completed. Other specimens

examined in the study were borrowed from the Academy of Natural Sciences of

Philadelphia (ANSP), Auburn University Museum of Natural History (AUMNH), Florida

State Collection of Arthropods (FSCA), Mississippi Entomological Museum (MEM),

United States National Museum (USNM), and the University of Wisconsin Insect

Research Collection. All type specimens of newly described species are deposited in the

USNM, with paratypes being deposited in the ANSP, MEM, and UMMZ.

In order to conduct a thorough study of the Scudderi group, the male genitalia,

which are typically concealed within the terminalia, were examined. In order to reveal

these structures, specimens were relaxed by soaking them in hot water for several hours

or cold water overnight. Once the specimen was sufficiently relaxed, it was positioned

under a Leica MZ 12.5 dissecting stereomicroscope, such that the terminalia were in

view, and using fine tipped forceps the pallium and subgenital plate are rotated ventrally.

Then the tips of the forceps were inserted into the body cavity anterior to the genital mass

until the tip is underneath the mass. The forceps tips are then used to raise the genital

mass out of the body cavity and into position for examination. Most specimens were 7

examined in this matter, but for each species several more detailed examinations were made by cutting the muscles and connecting tissue, removing the genital structures from

the body cavity, and placing these in a 10% solution of potassium hydroxide (KOH) for

approximately 5 minutes to soften the extraneous tissue. This tissue then was brushed

away from the sclerotized genital structures with forceps or stiff brush. After

examination, the cleaned genital mass was then placed in a micro-vial containing glycerin

and stored on the pin below the specimen for future reference.

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. Species distributions maps were produced using Google Earth Pro© software.

Measurements were made with a measuring 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.

Tegmina Length — Laterally from the junction of the ventral margin of the

tegmen and the caudal margin of the pronotum to their apex

Cercus Length — Laterally, maximum possible measurement.

Cercus Basal Width — Laterally, along the point of attachment from the dorsal to

ventral margin.

Mid Cerci Width — Laterally, at the mid-length of the cerci

Cercus Apex Width — Laterally, along the distal end.

Species delimitation

Hubbell (1932) was the first to call attention to the interspecific variation in the

Melanoplinae, and since then characters of the male aedeagus have been used extensively

in defining species in the group (Roberts 1941, Gurney 1951, Otte 2003, Barrientos

Lozano 2011, Otte 2012). It is assumed that variability in the genitalia of Melanoplus function as reproductive isolating mechanism, which in turn leads to genetic variability.

Here, as with other recent revisions within the subfamily, new species designations were based primarily on distinct differences in the male genitalia. This variation is found

primarily in the shape of the valves. Additional characters used for species delimitation

include external characters, such as male cerci shape, furcula length, and discrete, definable distributions.

Molecular techniques

A more thorough molecular study of the group is currently underway. However,

DNA barcode data were available for 39 specimens as part of another project by JGH.

(See Table 1 for sample info.) Sequences from Melanoplus pygmaeus (Davis) and

Chortophaga viridifasciata (DeGeer) were also available and used for near and far

outgroups respectively. DNA barcoding seeks to use a short segment of an organism's

DNA for identification purposes.

Table 1 Taxa and associated collecting data for specimens used in molecular analysis.

Species State County Locality Accession # Melanoplus scudderi MS Monroe Peacely Ferry Rd. Melanoplus scudderi MS Hinds Natchez Trace Prkway JN281401 Melanoplus scudderi MS Choctaw 6 mi S Eupora Melanoplus scudderi MS Itawamba 2 mi E Fulton Melanoplus scudderi MS George 4.5 mi NW Lucedale Melanoplus scudderi MS Calhoun Skuna JN281406 Melanoplus scudderi MS Chickasaw Buena Vista JN281405 Melanoplus scudderi TN Wilson Vesta Glade NA JN281404 Melanoplus scudderi TN Davidson Couchville Glade NA JN281402 Melanoplus scudderi MS Attalla Natchez Trace Prkwy JN281400 Melanoplus scudderi TN Maury Stillhouse Hollows NA JN281413 Melanoplus scudderi AL Bibb Bibb Co. Glades NA JN281407 Melanoplus scudderi AL Dallas 5 mi NW Cahaba Melanoplus texarkana LA Natchitoches Kisatchie NF Melanoplus ouachita AR Conway Petit Jean SP Melanoplus scudderi AR Union Strong JN281398 Melanoplus scudderi LA Union 18 mi SE Farmerville JN281397 Species State County Locality Accession # Melanoplus folkertsi FL Gadsden 2 mi S Chattahoochee JN281421 Melanoplus folkertsi GA Decatur 3.34 mi S Faceville JN281411 Melanoplus folkertsi GA Decatur Silver Lake WMA JN281409 Melanoplus folkertsi AL Escambia W CR 40 X HWY 41 Melanoplus folkertsi GA Decatur Silver Lake WMA JN281412 Melanoplus quercicola FL Liberty 1.1 mi S Torreya SP JN281423 Melanoplus savannah SC Aiken 10 mi NW Aiken Melanoplus savannah SC Aiken 2.7 mi N Montmorenci Melanoplus savannah SC Barnwell Ditch Pond HP JN281422 Melanoplus scudderi GA Twiggs 6.2 mi E Bonaire JN281415 Melanoplus acidocercus GA Bibb Mogul JN281428

Melanoplus carnegiei GA Bryan Groveland JN281425 Melanoplus carnegiei GA Bryan Groveland JN281424 Melanoplus quercicola GA Emanuel 14.5 mi S Swainsboro JN281424 Melanoplus carnegiei GA Burke Yuchi JN281427 Melanoplus acidocercus GA Crawford 2 mi E Knoxville JN281447 Melanoplus scudderi GA Muscogee 6.5 mi E Columbus JN281408 Melanoplus muscogee GA Marion 2 mi SE Juniper JN281418 Melanoplus muscogee GA Taylor Fall Line Sandhills NA JN281420 Melanoplus GA Talbot Black Jack Crossing JN281426 chattahoochee

A 658-bp region of mitochondrial cytochrome c oxidase I (COI) is the most common gene used in animals (Hebert et al. 2003). This technique works well for many taxa, with some studies showing that more than 95% of species in test assemblages possess distinctive COI sequences (Herbert et al. 2003). However for some taxa, particularly the Orthoptera, DNA barcoding either fails to distinguish closely related species or in some cases overestimates the number of species, likely as a result of a large number of nuclear mitochondrial pseudeogenes (numts). (Song et al. 2008, Walker and

Funk 2014). As no other molecular data were yet available, the barcode data were

analyzed not for species delineation, but rather to test whether biogeographic patterns

attributed to Pleistocene glaciation displayed by other organisms in the southeast were present in the Scudderi group.

To obtain genetic material, the right mid-femora from pinned specimens were sent to the Canadian Centre for DNA Barcoding for DNA extraction and, polymerase chain reaction (PCR) and sequencing. PCR was performed using the primers

LCOI1490_t1/HCOI2198 with M13 tails. A maximum of 658 base pairs of the mitochondrial cytochrome oxidase I (COI-5P) gene were sequenced. Sequence data are currently managed under project OMEM at the Barcode of Life Data Systems (BOLD,

Biodiversity Institute of Ontario, University of Guelph, Guelph, Ontario Canada;

boldsystmes.org), and have also been submitted to GenBank (accession numbers are

listed in Table 1.). Sequences were aligned with MUSCLE using default parameters and

Neighbor Joining, Maximum Parsimony and Maximum Likelihood trees were

constructed in Mega6 (Tamura et al. 2013). Resulting trees were imported into

FigTree1.4.2 (http://tree.bio.ed.ac.uk/software/figtree/) and Photoshop CS6® for

presentation.

Results

Examination of the internal genitalia revealed that M. s. scudderi is actually a

complex of 23 species, including the subspecies Melanoplus s. latus and M. s. texensis

being raised to species level. Based primarily on the shared characteristic of having a

reduced aedeagal sheath that only forms a collar around the base of the valves, and

generally shorter tegmina (mean 4.2 mm) species), M. carnegiei and M. acidocercus

along with three new species, Melanoplus chattahoochee, Melanoplus carolinensis, and

Melanoplus savannah, are hereby placed in the newly established Carnegiei species

group. In contrast, species in the scudderi group possess a sheath that is produced and a

mean tegmina length of 4.8 for species in the Scudderi group

Comparison of the groups

Scudderi Group

1. Cerci generally falcate (except M. davisi and M. quercicola)

2. Aedeagal sheath not reduced, covering more of the valves

3. Wings relatively longer (mean 4.8 mm)

Carnegei Group

1. Cerci generally triangular (less so in M. savannah and M. carolina)

2. Aedeagal sheath reduced, coving only the base of the valves.

3. Wings relatively shorter (mean 4.2 mm).

Diagnosis of the Melanoplus scudderi species group

Structural Characters

Species of medium to small size (♂ 14.5 - 24.2 mm, ♀ 16 – 28 mm). Head

slightly wider than pronotum; fastigum steeply declivent, Eyes somewhat prominent,

especially in males. Antennae usually with 22 flagellomeres, but often 23 to 26; nearly

cylindrical; equal in width throughout, except two basal articles. Pronotum broadly

convex with distinct, but broadly rounded shoulders, anterior margin sub-truncate, often

somewhat emarginate; lateral margins of prozona parallel, metazona margins diverging

posteriorly; median carina distinct and equal in height throughout; anterior and median

sulci present laterally, indistinct near the median carina; posterior sulci dissecting the

median carina; prozona mostly smooth, with light punctation on the lobes ventrally;

metazona densely punctate, posterior margin broadly obtuse-angulate, prosternal spine

short, sub-cylindrical, blunt. Tegmina ovate, apices varying from subtruncate to

subacuminate; dorsal margins overlapping; length variable but, typically extending to the

anterior edge of the second abdominal tergite, some individuals with tegmina covering

three-fourths or entire abdomen. Pro- and mesothoracic legs moderately stout, the

femora of the mesothoracic leg slightly more inflated than that of the prothoracic leg; metathoracic legs with the femora greatly swollen, extending just beyond the caudal end of abdomen. Metathoracic tibia with eleven to sixteen pairs of spines, but typically 12-13.

Furcula (Fig. 1a) typically rounded protuberances, projecting slightly beyond the end of the segment from which they originate; bases minutely separated. Supra-anal plate (Fig.

1a) triangular, slightly longer than wide, with the median groove anteriorly distinct with

elevated sides, and diverging and becoming less distinct posteriorly. Cercus (Fig. 1b)

falciform, being broader at base than apex, and typically curving at the mid-point, and then tapering to rounded apex (except in M. davisi and M. quercicola). Subgenital plate

slightly conical (Fig 1a-b).

Phallic Structures

The dorsal valves are conical, tapering to a point, shortened to elongate depending

on the species. Ventral valves typically dorso-ventrally flattened and variously shaped

depending on the species. Aedeagal sheath of various lengths, but typically coving more

than the base of the valves. The epiphallus is of the typical Melanoploid shape, having

lophi, ancorae, and an undivided bridge, but more precisely, species in the Scudderi

group have a concave bridge, broadly rounded lophi, convexly curved lateral plates that

are subdeltate in shape with a rounded anterior lobe and an acuminate caudal tip, and

ancora that are triangular in shape, often tapering to a point (Fig. 1G-I).

Coloration

Males: Antenna ferruginous with a dark band on the ventral surface of the second

basal segment. Antennal crescent complete. A black spot present at the juncture of the

clypeus, gena, and frons. Head, thorax, and abdomen ferrugineou-fuscous, infuscated

dorsally, testaceous ventrally. A lateral, well-defined, piceous, post-ocular stripe extends

from the caudal margin of the eye to the second abdominal tergite, but often enfeebled or

lacking on the metazona; lateral area of head, thorax, and abdomen below post-ocular

stripe greyish-yellow. Tegmina ferrugineo-fuscous with slightly lighter venation. Hind

femora ferrugineo-testaceous dorsally and laterally, bimaculate or often feebly

trimaculate, with fuscous spots often extending halfway across medial area of lateral

surface; castaneous ventrally, femur-tibia connection black. Hind tibia bright red, often dulled basally; with black or black tipped spines. (Fig. 1J)

Species accounts

Melanoplus scudderi Uhler (Figs. 1A-J, 2A-B, 3, 4A-B)

Pezotettix scudderi Uhler 1864: 555

Podisma scudderi, Walker 1870: 718

Pezotettix unicolor Thomas 1873: 151

Melanoplus scudderi, Scudder 1897: 212

Male Measurements (mm)

(n = 60) Body length 14.5 – 24.0 (mean = 19.7); pronotum length 3.5 – 5.5 (mean

= 4.5); tegmen length (brachypterous) 3.1 – 6.0 (mean = 4.8), (macropterous) 12.0; hind

femur length 8.5 – 12.5 (mean = 10.9); cerci length 0.9 - 1.4 (mean = 1.2); basal width of cerci 0.6 - 0.9 (mean = 0.6); mid-cercal width 0.4 - 0.7 (mean = 0.5); cerci apex width

0.2- 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 40) Body length 16.0 – 26.8 (mean = 22.6); pronotum length 4.2 – 7.0 (mean

= 5.6); tegmen length (brachypterous) 4.0 – 8.3 (mean = 6.2), (macropterous) 11.2 – 15.0

(13.0); hind femur length 6.2 – 16.4 (mean = 12.5).

Diagnosis

Differing from other species in the shape of the internal male genitalia (Fig. 1C-

F). In dorsal view, the dorsal valves are curved laterally subapically, and the ventral

valves extend approximately two-thirds length of dorsal valves. In lateral view, the 15

valves are slightly bowed dorsally and with the dorsal valves appearing recurved dorsally. East of the Appalachian mountains, the apically curved portion of the dorsal valves are shorter and less strongly curved (Fig 2 A-B).

Lectotype here designated

♂; Oct 18, Uhler [note- labels do not have locality information].

Other specimens examined

Alabama: Autauga Co., 4.4 mi SSE Billingsly, 32º35’49”N 86º 42’28”W, 18

September 2013, J.G. Hill (1♂, 1♀), Prattville, 17 September 1959, M.E. Dakin (1♂,

3♀). Bibb Co., Bibb Co. Glades Preserve, 33°03’26”N 87°02’02”W, 24 August 2010, J.

G. Hill (2♂). Chilton Co., 12 mi SE Clanton, 13 September 1962, M.E. Dakin (7♂, 6 ♀);

Ala 22 near Coosa River, 15 Sept 1962, M.E. Dakin (8♂, 4♀). Choctaw Co., Butler, 1

October 1959, M.E. Dakin (2♀). Clay Co., 12. 5 mi SE Munford, 12 October 1962, M.E.

Dakin (2♂, 2♀). Cleburne Co., Summit of Mt. Cheaha, 6 September 1959, M.E. Dakin

(4♂); 12 October 1962. (1♂). Colbert Co., 15 mi S Cherokee, 34°35’28”N 87°56’11”W,

13 September 2012, J. G. Hill (1♂, 1♀); 3 mi E Tuscumbia, 15 August 1953, J. A. G.

Rehn (1♂). Dallas Co., Selma, 9 September 1915, Rehn & Hebard (2♂). DeKalb Co.,

DeSoto State Park, 5 September 1959, M.E. Dakin (2 ♂, 2♀); Little River Canyon, 5

September 1959, M.E. Dakin (3 ♂, 2♀); 2 mi SE Mentone, 16 August 1953, J. A. G.

Rehn (1♂). Muscadine, 3 September 1939, Rehn & Rehn (1♂). Greene Co., Black

Warrior River on HWY 14, 12 October 1959, M.E. Dakin (7♂, 2♀); Boligee, 2

September 1939, Rehn & Rehn (1♂). Lee Co., 3.7 mi W Opelika on U.S. 280, 10

September 1954, T.H. Hubbell & I.J. Cantrall (12♂, 5♀) Chewacla State Park, 11 August

1960, M.E Dakin (4♂, 1♀). Macon Co., 2.6 mi S Tuskegee, 4 October 1962, M. E.

Dakin (1♂). Marengo County, 2.8 mi W Dixons Mill on U.S. 43, 14 September 1954,

I.J. Cantrall (7 ♂, 6 ♀). Marion Co., 6 mi SW Hackleburg, 34°14’31”N 87°53’45”W, 13

September 2012, J. G. Hill (1♀). Madison Co., 5 mi W Huntsville, 8 August 1939, Rehn

& Rehn (1♂); Huntsville, 29 August 1962, T.H. Hubbell (1 ♂). Montgomery, 8

September 1915, Rehn & Hebard (1♂). 4 mi E Mississippi state line, 1 September 1955,

Rehn (1♂). Pickens Co., 0.7 mi W Ethelsville, 33°25’21”N 88°13’35”W, 8 August 2010,

J. G. Hill (3♂). Sumter Co., Livingston, 32º36’15”N 88º11'13”W, 19 September 2013,

J.G, Hill (1♂, 5♀). Washington Co., 6.5 mi N Wagerville, on U.S. 43, 12

September1958, I. J. Cantrall (6♂, 5♀); Sunflower, 12 September 1958, I. J. Cantrall

(4♂, 1♀). Wilcox Co., Camden, 10 September 1959, M.E. Dakin (1♂). Arkansas:

Baxter Co., 3 mi W Baxter-Fulton County line on US-62, 26 August 1954, T. H. Hubbell

& I. J. Cantrall (5♂, 11♀). Conway Co., 1 mi W Morrilton on US-64, 8 September 1958,

I. J. Cantrall (7♂). Fulton Co., 0.8 mi E Glencoe, 26 August 1954, T. H. Hubbell & I. J.

Cantrall (4♂, 7♀). Marion Co., 1.2 mi W center of bridge over White River, 27 August

1954, T. H. Hubbell & I. J. Cantrall (13♂, 10♀). Conway Co., 1 mi W Morrilton on US-

64. 8 September 1958, I. J. Cantrall (8♂, 7♀). Dallas Co., 1 mi N Fordyce on US-167, 8

September 1958, I. J. Cantrall (4♂). Grant Co., 1.6 mi S Sheridan on US-167, 8

September 1958, I. J. Cantrall (8♂). Izard Co., Devils Knob N.A., 36º00’21”N 92º

02’44”W, 24 September 2013, J.G. Hill (1♂, 5 ♀). Logan Co., Short Mountain Creek, 1 mi S Paris, 28 August 1954, T. H. Hubbell & I. J. Cantrall (2♀). Marion Co., Crooked

Creek Water Trail, 36°14’47”N 92°50’06”W, 25 September 2013, J. G. Hill (3♂, 9♀).

Newton Co., 5.4 mi S Jasper, 27 August 1954, T. H. Hubbell & I. J. Cantrall (11♂, 3♀).

Pope Co., 0.5 mi N center of Arkansas River bridge on Hwy. 7, 28 August 1954, T. H.

Hubbell & I. J. Cantrall (12♂, 7♀); 0.6 mi E Illinois Bridge on US-64, 7 September 1958,

I. J. Cantrall (4♂, 2♀); west bank of Illinois River on US-64, 7 September 1958, I. J.

Cantrall (4♂, 3♀). Prairie Co., Railroad Prairie N. A. 34°48’50”N 91°31’33”W, 19

September 2012, J. G. Hill (4♂, 4♀). Randolph Co., 8.8 mi SW Pocahontas on US-62,

26 August 1964, T. H. Hubbell & I. J. Cantrall (4♂). Sharp Co., 9.6 mi W Imboden on

US-62 & 63, 26 August 1954 (7♂, 11♀). Stone Co., Ozark N.F., 36º06’05”N 92º

11’08”W, 25 September 2013, J.G. Hill (7♂, 9♀). Connecticut: Middlesex Co., Deep

River, 24 August 1904, A.P. Morse (1♂). New Haven Co., New Haven, 29 August 1891,

A.P. Morse (5 ♂, 5 ♀); 29 August 1892, (2♂). Georgia: Baldwin Co., 2.5 mi S

Milledgeville on U.S. 441, 9 September 1954, T.H. Hubbell & I.J. Cantrall (3♂, 2♀).

Floyd Co., Thomas, 9 September 1924, T. H. Hubbell (6♂, 15♂). Macon, 26 September

1918, H. Fox (1♂); 11-16 October 1918 (2♂, 1♂); 16-23 December 1918 (3♂, 3♀); 30

December 1918 (5♂, 4♀); 19 January 1919 (3♂); 22 February 1919 (1♂); 12 March

1919 (1♂); 24 March 1919 (1♀); 27 September 1919 (1♂,1♀); 9 October 1920 (1♀); 16

October 1920 (1♂); 28 October 1922 (6M, 1♂♀); 20 September 1924 (2♂,1♀);

Lakeside, near Macon, 1 October 1921, H. Fox (1♀). Butts Co., Cedar Rocks, 5 mi N

Jackson, 27 October 1923, H. Fox (3♂, 6♀). Chattooga Co., 3.5 mi E Menlo, 16 August

1953, Rehn (1♂). Clarke Co., Athens, 2 December 1922, (1♂). Cobb Co., Austell, 27

August 1910, (1♂). Dade Co., Trenton, 18 September 1951, M. J. D. White (1♂, 2♀).

Decatur Co., Bainbridge, 6 September 1915, Rehn & Hebard (2♂, 5♀). Forsyth Co.,

Coal Mountain, 4 September 1939, Rehn & Rehn (1♂). Fulton Co., Alpharetta, 4

September 1939, Rehn & Rehn (1♀); Atlanta, 21-22 October 1922, H. Fox, (6♂, 5♀).

Hancock Co., 2.4 mi W Sparta on U.A. 22, 9 September 1954. T.H. Hubbell & I.J.

Cantrall (5♂, 3♀). Henry Co., Locust Grove, 13 October 1923 (4♂, 1♀). Lumpkin Co.,

4 mi S Dahlonega, 4 September 1939, Rehn & Rehn (1♂). McDuffie Co., 6.3 mi N.

McDuffie-Warren Co. line, 17 September 1954, T.H. Hubbell & I.J. Cantrall (4♂, 2♀).

McIntosh Co., 8 mi N Darien, 1 September 1945, Rehn (1♀). Muscogee Co., 6.5 mi E

Columbus, 32°33’16”N 84°45’38”W, 25 August 2010, J. G. Hill (1♂, 1♀); 1.75 mi E

Upatoi, 32°32’51”N 84°42’20”W, 11 October 2010, J. G. Hill (2♂, 2♀). Morgan Co., .3 mi S Apalachee River on U.S. 41, 1 September 1951, I.J. Cantrall (1♂). Rabun Co.,

George Mountain, near Dillard, 4 September 1917, Rehn & Hebard (4♂, 2♀). Union

Co., 5 mi N Blairsville, 4 September 1939, Rehn & Rehn (1♂). Illinois: Cass Co.,

Arenzville, 14 August 1913, (1♂). Champaign Co., Homer, 7 October 1906, Hart &

Hood (4♂); Urbana, 28 August 1906, (2♂). Clark Co., Martinsville, 8 August 1929, T. H.

Hubbell (7♂, 1♀). Clay Co., Clay City, 2 September 1909 (1♂). Cook Co., Willow

Springs, 17 September 1905, W. J. Gerhard (1♂); 17 August 1906, E. B. Chope (1♂).

Gallatin Co., Shawneetown, 12 October 1932, Frison & Ross (1♂). Hamilton Co.,

Dahlgren, 24 September 1952, H. H. Ross (1♂). Hardin Co., Elizabethtown, 12 October

1928, Frison (1♂). Jackson Co., Carbondale, 22-25 September 1909, Gerhard (4♂, 4F);

22 October 1957, D. A. Rossman (1♂); 3 mi E Carbondale on IL-13, 25 August 1954, T.

H. Hubbell & I. J. Cantrall (8♂, 4♀); S bank of Big Muddy River on State Hwy. 3, 31

August 1958, I. J. Cantrall (1♂, 2♀); N bank of Big Muddy River on State Hwy. 3, 31

August 1958, I. J. Cantrall (4♂, 1♀). Jersey Co., Grafton, 8 September 1932, Ross &

Mohr (1♂). Lake Co., Beach, 21 August 1906, (1M); 24-26 August 1932, Ross & Mohr

(2♂); Waukegan, 20 August 1906 (1M). Mason Co., Havana, 11 August 1907, (1♂).

Massac Co., Metropolis, 4 September 1932, Ross & Horsfall (5♂). Peoria Co.,

Bartonville, 29 August 1910 (1♂). Piatt Co., White Heath, 20 October 1906, Hart &

Hood (1♂). Pope Co., Dixon Springs, 5 September 1932, Ross & Horsfall (3♂); 13

October 1932, Frison & Ross (2♂); Golconda, 13 October 1932, Frison & Ross (5♂).

Indiana: Brown Co., Brown County State Park, 8 October 1952, D.C. Eades (1♂, 1♀).

Crawford Co., 3 September 1902, W.S. Blatchley (1 ♂). Greene Co., 2 mi S Switz City,

6 November 1956, F.N. Young (1♂, 4♀); Lattas Creek Prairie, 9 September 1958, F.N.

Young (4♂, 2♀). Lake Co., Hessville, 22 September 1907, W.J. Gerard (1♂, 1♀).

Marion Co., 6 September 1896, W.S. Blatchley (2 ♂). Marshall Co., 11 November 1903,

W. S. Blatchley (1♀). Putnam Co., 13 March 1904, W.S. Blatchley (1♂, 1♀).

Vanderburgh Co., Hwy 57 N Evansville, 29 September 1956, D, C. Eades (3♂, 2♀).

Vigo Co., 18 October 1898, W. S. Blatchley (1♂). Iowa: Hancock Co., Pilot Knob State

Park, 22 August 1958, I.J. Cantrall (5♂, 7♀). Henry Co., Mt. Pleasant, 19 September

1935, Countney (2♂). Ida Co., 20 July 1935, H.C Knutson, (2♂). Johnson Co., Iowa

City, 11 October 1910 (2♂); 29 October 1933, Wickham (2♂, 1♀). Monroe Co., 0.5 mi

N Turin, 24 August 1958, I.J. Cantrall (6♂, 2♀). Plymouth Co., 4.2 mi Westfield, 24

August 1958, I. J. Cantrall (15 ♂, 7♀). Taylor Co., 10 October 1934, Carmen (1 ♂).

Union Co., 11 October 1935, Courtney (1♂). Warren Co., 3 October 1934, Rogers (1♂).

Woodbury Co., 7 September 1935, H.E. Jaqua (1♂). Worth Co., 19 September 1935,

Courtney (1♂). Kentucky: Bath Co., Licking River, 25 August 1956, D.C. Eades & J.

Ldn (3♂). Carroll Co., General Butler State Park, 24 August 1956, D.C. Eades & Lui

(1♂, 1♀). Christian Co., Pennyrile State Forest, 20 August 1956, D.C. Eades (1 ♀).

Fulton Co., Fulton, 15 September 1954, T. H. Hubbell & I.J. Cantrall (10♂). Louisiana:

East Feliciana Par., 0.2 mi S Ethel on LA-19, 10 September 1958, I. J. Cantrall (2♂, 2F).

Grant Par., Kisatchie N. F., 31°33’08”N 92°26’03”W, 1 October 2013, J. G. Hill (1♂,

3♀). Union Par., 18 mi SE Farmerville, 32°42’02”N 92°06’11”W, 2 November 2009, J.

G. Hill & D. Otte (1♀). Washington Par., 0.3 mi N Bogalusa city limits, 11 September

1958, I. J. Cantrall (4♂, 2♀). Maryland: Allegany Co., 4 mi E Belle Grove, 5

September 1933, Rehn (4♂, 2♀). Anne Arundel Co., High Island, 10 November 1901,

A. N. Caudell (1♂). Calvert Co., Kenwood Beach, 14 September 1937, A. B. Gurney

(2♂); Little Cove Point, nr. Solomon’s Island, 15 Oct, 1937, (1♂); Battle Creek, 14

September 1937, A. B. Gurney (1♂); Wide Water, 13 October 1934, H. A. Allard (1♂);

nr. Dare’s Beach, 20 September 1956, A. B. Gurney (2♂, 1♀). Cecil Co., Bald Friar, 17

October 1913, B. Long (1♂, 1♀). Charles Co., Pope’s Creek, 22 October 1937, H. A.

Allard. Montgomery Co., Plummer’s Island, 11 October 1906, A. N. Caudell (1♂, 1♀);

29 August 1971, G. F. Hevel (1♂, 1♀); 11 September 1915, M. Hebard (2♂, 2♀); 13

October 1918, M. Hebard (1♂, 1♀); 27 October 1918, Rehn & Hebard (1♂); 28 October

1967, J. F. G. Clarke (2♂, 1♀); 4 mi S Ashton, 4 August 1982, G. F. & J. F. Hevel (1♂);

15 August 1982, G. F. & J. F. Hevel (1♂); Cabin John, 21 September 1956, A. B. Gurney

(1♂). Minnesota: Ramsey Co., St. Anthony Park, 17 August 1899, O. Lugger (1♀).

Massachusetts: Hampden Co., Springfield, J.A. Allen (1 ♂). Plymouth Co: Wareham, 1

September 1917, A.P. Morse (1 ♂). Michigan: Livingston Co., E.S. George Reserve, 22

September 1953, I.J. Cantrall (2♂, 1♀); 16 September 1954 (2♀); 24 August 1970 (1♀).

Washtenaw Co., Ann Arbor, 19 September 1933, I.J. Cantrall (1♂, 3♀); 22 August 1955,

S. K. Gangware (9♂, 8♀). Ann Arbor Arboretum, 27 September 1954, S. K. Gangware

I. J. (15♂, 18♀); Dixboro, 13 October 1957, T.J. Cohn (1♂, 1♀); Lima Twp, 6

September 1920, T.H. Hubbell (1♀). Wayne Co., Grosse Ile, 23 October 1955, G.

Steyskal (1♂, 1 ♀); 10 September 1958 (3♂, 2♀). Mississippi: Adams Co., Natchez, 13-

14 September 1915, Rehn (1♂); Natchez on U.S. Hwy 84, 23 October 1937, Cantrall,

Cohn, & Eades (1 ♂). Amite Co., E outskirts of Gloster, 10 September 1958, I. J.

Cantrall (2♂). Attalla Co., Natchez Trace mi 165, 33°04’56”N 89°31’33”W, 29

September 2008, J. G. Hill (1♀); Natchez Trace mi 168, 33°07’45”N 89°29’34”W, 28

October 2009, J. G. Hill & D. Otte (1♂). Calhoun Co., Calhoun WMA, 33°52’14”N

89°27’49”W, 10 October 2010, J. G. Hill (1♂, 2♀); Skuna Waterway, 33°57’05”N

89°27’28”W, 1 October 2010, J. G. Hill (1♂). Carroll Co., Whitehead Creek,

33°29’16”N 89°18’27”W, 19 September 2012, J. G. Hill (1♂, 8♀). Chickasaw Co.,

Buena Vista, 33°53’30”N 88°49’32”W, 17 September 2010, J. G. Hill (2♂, 1♀). Natchez

Trace Pkwy, 33°56’48”N 88°55’39”W, 21 October 2010, J. G. Hill (1♂, 1♀); Tombigbee

Natl. Forest, 33°55’39”N 88°51’18”W, 18 October 2001, J. G. Hill (3♀); 19 November

2001, J. G. Hill (1♀); 28 July 2003, J. G. Hill (2♀); 17 September 2010, J. G. Hill (1♂);

33°55’35”N 88°51’13”W, 1 October 2004, J. G. Hill (1♀); 33°55’39”N 88°51’18”W, 4

November 2006, J. G. Hill (1♂). Choctaw Co., 1.3 mi N Ackerman on MS-15, 14

September 1954, T. H. Hubbell and I. J. Cantrall (2♀); ~ 6 mi S Eupora, 33°28’09”N

89°13’32”W, 19 September 2007, J. G. Hill (2♂, 1♀). Natchez Trace mi 198,

33°28’43”N 89°12’43”W, 28 October 2009, J. G. Hill & D. Otte (1♂). Covington Co.,

0.8 mi E Collins on US-84, 25 October 1957, Cantrall, Cohn and Eades (2♀). Franklin

Co., 1.2 mi W Roxie, 25 October 1957, Cantrall, Cohn and Eades (1♀). Jackson Co.,

Sandhill Crane Refuge Preserve, 30°55’40”N 88°35’55”W, 19 September 2011, J. G.

Hill (2♂, 1♀). Hinds Co., Jackson, 12 September 1915, Rehn & Hebard (1♂), 17 August

1929, T. H. Hubbell (2♂); Clinton, 17 August 1929, T. H. Hubbell (2♂); Between Bolton and Clinton, 1 September 1939, Rehn and Rehn (1♂); Natchez Trace mi 85, 32°19’11”N

90°21’41”W, 1 November 2009, J. G. Hill & D. Otte (1♂). Itawamba Co., 2 mi E Fulton

34°14’29”N 88°26’39”W, 22 August 2007, J. G. Hill (3♂, 1♀); Canal Section WMA,

34°15’50”N 88°26’44”W, 22 August 2007, J. G. Hill (3♂). Jasper Co., 2 mi E Rose Hill,

32°07’30”N 89°01’45”W, 9 October 2005, J. G. Hill (1♀); 8 September 2006, J. G. Hill

(1♂, 1♀); 1 mi N Stafford Springs on US-11, 25 October 1957, Cantrall, Cohn and Eades

(2♀). Jefferson Davis Co., Jefferson Davis Lake, 31°33’47”N 89°50’39”W, 8 September

2006, J. G. Hill & J. A. MacGown (1♂). Jones Co., 1 mi W Laurel city limits, 25

October 1957, Cantrall, Cohn and Eades (2♀). Kemper Co., 5 mi N Scooba, 32°52’24”N

88°29’28”W, 3 September 2005, J. G. Hill (1♂, 1♀); 1 mi S Electric Mills, 29 August

2005, J. G. Hill (1♂). Jones Co., 1 mi W Laurel city limits, 25 October 1957, Cantrall,

Cohn and Eades (2♀). Lauderdale Co., Kewanee, Cuba Rd., 15 August 2008, J. G. Hill,

L. C. Majure (1♂); Meehan, Chunky River X Pt. Wanita Rd., 18 August 2005, J. G. Hill

(1♂); Meridian, 32°21’13”N 88°39’36”W, 19 August 2005, J. G. Hill (1♂); 0.5 mi W

Vimville, 14 September 1954, T. H. Hubbell and I. J. Cantrall (1♀); 2.5 mi W Vimville,

14 September 1954, T. H. Hubbell and I. J. Cantrall, (2♀). Lowndes Co., Black Prairie

WMA, 33.33194°N 88.56325°W, 29 September 2011, J. G. Hill, C. E. Chesnut (2♀);

Crawford, 33°18’01”N 88°36’38”W, 8 September 2004, J. G. Hill (3♂, 1♀); 2.06 mi S

Mayhew, 33°28’40”N 88°40’09”W, 10 September 2009, J. G. Hill (1♂, 1♀). Noxubee

Co., Noxubee NWR, 33°12’48”N 88°49’38”W, 3 October 2008, J. G. Hill (1♂, 2♀);

33°13’32”N 88°48’19”W, 22 September 2008, J. G. Hill & J. A. MacGown (3♂);

33°15’08”N 88°46’14”W, 29 August 2008, J. G. Hill (1♂); 33°15’33N 88°46’18”W, J.

G. Hill (1♂); 1 mi N Brooksville, 33°15’32”N 88°33’19”W, 3 September 2005, J. G. Hill

(1♂). Lee Co., Natchez Trace Parkway mi 261.8, 34°15’24”N 88°45’22”W, 13 October

2006, J. G. Hill (1♂); 3 mi E Tupelo, 34°14’26”N 88°48’39”W, 13 October 2006, J. G.

Hill (2♂, 2♀). Lincoln Co., 2.8 mi E Brookhaven on US-84, 25 October, 1957 (2♀).

Monroe Co., Peacely Ferry Rd., 34°00’20”N 88°34’23”, 28 August 2006, J. G. Hill (4♂,

1♀). Montgomery Co., Winona, 15 September 1915, Hebard (3♂). Newton Co.,

Bienville Natl. Forest, 32°15’10”N 89°15’44”W, 9 October 2005, J. G. Hill (1♂);

Junction of Hwy 80 & Interstate 120, 32°20’07”N 89°17’10”W, 19 August 2005, J. G.

Hill (2♂); 9 October 2005, J. G. Hill (1♂). Neshoba Co., N of Burnside on MS-15 (2.6 mi. S Winston Co. line), 14 September 1954, T. H. Hubbell and I. J. Cantrall (2♀).

Oktibbeha Co., Osborn, 33°36’21”N 84°44’09”W, 20 October 2004, J. G. Hill & M. E.

Dakin (2♂). 33°30’41”N 89°27’49”W, 15 September 2007, J. G. Hill (1♀). Pike Co., 0.8 mi E Junction US-51 and US-98, 11 September 1958, I. J. Cantrall (2♂); Summit,

September 1915, K. L. Spurlock (1♂); 1.1 mi E of Walthall-Pike County line, 11

September 1958, I. J. Cantrall (2♂); W bank of Bogue Chitto River on US-98, 11

September 1958, I. J. Cantrall (2♂). Pontotoc Co., Natchez Trace State Park,

34°05’52”N 88°53’12”W, 13 October 2006, J. G. Hill (1♂, 1♀); Natchez Trace mi 247,

34°05’52”N 88°51’37”W, 22 August 2009, J. G. Hill (1♂); Natchez Trace mi 250,

34°07’49”N 88°50’18”W, 22 August 2009, J. G. Hill (1♂); 34°08’04”N 88°50’00”W, 1

October 2009, J. G. Hill (1♂). Scott Co., Bienville Natl. Forest, 32°20’12”N

89°26’21”W, 9 October 2005, J. G. Hill (1♂); 6.5 mi S Forest, 32°16’09”N 89°29’47”W,

28 October 2009, J. G. Hill & D. Otte (1♂); Pinkston Hill, T5N R9E Section 15, 17

January 2005, J. G. Hill (2♀); Pinkston Hill, 32°16’06”N 89°23’33”W, 19 August 2005,

J. G. Hill (1♂); Roosevelt State Park, 32°19’05”N 89°47’48”W, 1 November 2009, J. G.

Hill & D. Otte (1♂, 1♀). Tishomingo Co., Iuka, 8 September 1932, T. H. Hubbell (2♂);

1 mi E Paden, 34°39’18”N 88°15’10”W, 14 August 2012, J. G. Hill (1♀); Tishomingo

State Park, 34°36’20”N 88°11’33”W, 5 November 2005, J. G. Hill (1♀). Rankin Co.,

Strickton, 12 September 1915, Hebard (2♂); vic. Sandhill Rd. x Hwy 25 N, 32°28’40”N

88°53’40”W, 6 September 2007, J. G. Hill (10♂, 1♀); 1.5 mi NW Ludlow, 32°34’54”N

89°44’02”W, 6 September 2007, J. G. Hill (1♂). Walthall Co., 0.5 mi E Tylertown, 11

September 1958, I. J. Cantrall (2♂); 5 mi W center of Tylertown, 11 September 1958, I.

J. Cantrall (2♂). Warren Co., 8 mi E Vicksburg, 17 August 1929, T. H. Hubbell (1♂).

Washington Co., Greenville, 14 September 1915, Hebard (1♂). Webster Co., ~3 mi NW

Stewart, 33°28’43”N 89°25’05”, 19 September 2007, J. G. Hill (5♂). Wilkinson Co., 0.3 mi N State line on MS Hwy 33, 10 September 1958, I. J. Cantrall (2♂). Winston Co.,

Noxubee NWR, 33°13’55”N 88°54’40”W, 28 September 2007, J. G. Hill & J. A.

MacGown (1♂); 33°15’46”N 88°50’11”W, 3 October 2008, J. G. Hill & J. A. MacGown

(1♂). Yalobusha Co., Holly Springs N. F., 33°59’03”N 89°45’48”W, 10 October 2010,

J. G. Hill (1♀). Missouri: Benton Co., 4.3 mi E Cole Camp, 28 August 1958, I. J.

Cantrall (7♂, 6♀). Cass Co., Dent Co., Bunker, 29 August 1958, I. J. Cantrall (5♂, 3♀).

LaClede Co., 3 mi NE Phillipsburg on US-66, 21 August 1937, T. H. Hubbell (1♂).

Maries Co., 3.9 mi S Vienna on US-63, 29 August 1958, I. J. Cantrall (7♂, 16♀). Oregon

Co., 1 mi SE Koshkonong, 36º35'21"N 91º38'23"W, 11 September 2011, J.G. Hill (2 ♂).

Pettis Co., 2 mi SW Clifton City on MO-35, 27 August 1958, I. J. Cantrall. Reynolds

Co., Johnson Shut-in State Park, 29 August 1958 (36♂, 4♀). Saline Co., Arrow Rock

State Park, 27 August 1958, I. J. Cantrall (13M, 15♀). Webster Co., 2.5 mi W Seymour,

2 September 1958, I. J. Cantrall (4♂, 3♀). Nebraska: Cass Co., South Bend, 15

October 1910, Bruner (2♂, 3♀); Weeping Water, 24 September 1909, Bruner (7♂, 1♀).

Cuming Co., West Point, September 1884 (1♀). Douglas Co., Omaha (3♂). New Jersey:

Cape May Co., Bennett, 28 August 1912, H. Fox (1♂); Cape May, 8 September 1933, W.

Stone (1♀); between Woodbine & Belleplaine, 21 August 1912, H. Fox (1♂). North

Carolina: Buncombe Co., Asheville, 6 October 1932, E. Brundage (6♀). Chatham Co., 6

mi SE Hayesville, 4 September 1939, Rehn & Rehn (1♂, 2♀).; nr. Merry Oaks, 4

September 1945, Rehn (2♂). Clay Co., near Chatuge Lake dam, September 6 1954, T. H.

Hubbell & party (40♂, 6♀). Columbus Co., Lake Waccamaw, 8 September 1911, Rehn

& Hebard (1♀). Cumberland Co., Fayetteville, 9 September 1911, Rehn & Hebard (11♂,

2♀). Dare Co., Kitty Hawk, 28 August 1938, Gurney (2♂). Halifax Co., 2 mi S Weldon

on US-301, 13 September 1952, T. H. Hubbell et al. (14♂, 25♀). Henderson Co.,

Etawah, 5 September 1939, Rehn & Rehn (1♂, 2♀). Macon Co., 10 mi S Franklin, 4

September 1939, Rehn & Rehn (3♂, 2♀); 3 mi W Highlands, 8 September 1952, T. H.

Hubbell (2♀); Slopes along Cullasaja River, 1.5 mi below Cullasaja Falls, 16 August

1952, Rehn (1♂). Scotland Co., 5 mi SW Laurinburg, 4 September 1945, Rehn (1♂).

Wake Co., Raleigh, 2-22 1904, (4♂, 4♀); 25 August 1904, (1♀). Ohio: Adams Co.,

Buzzards Roost Rock, 20 September 1929, J.S Hine (3♂, 5♀); Locust Grove Prairie, 11

September 1932, E. & J. Thomas & C. Roth (3♂). Allen Co., Lima, 30 August 1932, E.

Thomas, C. Walker (1♂). Auglaize Co., 8 September 1935, E.S. Thomas (1♂).

Champaign Co., 27 August 1933, R.E. Moodruff (2♂, 1♀); 26 September 1953 (8 ♂, 4

♀). Clermont Co., 20 September 1931, E.T. Thomas (1♂, 2♀). Fairfield Co., Lancaster,

5 October 1930, C.F. Walker (1♀). Fairfield Co., Jacobs Ladder, 3 October 1931, M.D.

Nelswender (1♂); Madison, 11 October 1956, R, D. Alexander (1♂, 2 ♀). Franklin Co.,

Prairie T, 31 October 1933, E.S. & J. S. Thomas (1♂). Fulton Co., 10 October 1929, J.

Hine (1♂, 3♀). Geauga Co., Bainbridge, 16 September 1929, R. Goslin (7♂, 3♀).

Greene Co., Spring Valley, 30 September 1933, E. Thomas & D. Smith (1♂, 1♀).

Hamilton Co., Cincinnati, 5 September 1902, (2♂, 1♀). Highlands Co., 19 October

1929, J.S. Hine (4♂, 3♀), Ft. Hill, 6 November 1922, F. Thomas & E McPherson (2♂).

Hancock Co., Van Buren State Park 19 October 1957, T. E. Moore, (1♂). Hocking Co.,

"Neotoma", 11 September 1956, E.S. Thomas & D.C. Eades (2♂, 2♀); Good Hope, 11

October 1956, R.D. Alexander (5♂, 5♀). Ross Co., 3.5 mi S Clarksburg, 26 August

1962, R. D. Alexander (1♂, 2♀); Colerain, 1 September 1930, (1♂, 1 ♀); Seip Mound,

10 October 1931, M.D. Neiswender (1♂, 1♀). Scioto Co., Nile Twp, 12 September

1931, W.W. Baily (1♂, 2♀). Washington Co., Lowell, 15 September 1931, E. S. Thomas

(1♂, 1♀); Musk, 13-14 August 1932, E.S. & J.S. Thomas (2♂). Wood Co., 21 August

1930, J.S.Hine (6♂, 4 ♀). Pennsylvania: Bedford Co., vic. Breezewood, 25-26

September 1957, A. B. Gurney & H. A. Allard. Delaware Co., Castle Rock, 19

September 1908, Hebard & Rehn (4♂, 4♀). Fulton Co., Dott, 3 September 1938, Rehn

(4♂, 2♀); Warfordsburg, 25-26 September 1957, A. B. Gurney & H. A. Allard (1♂, 1♀);

nr. Lashley, 25-26 September 1957, A. B. Gurney & H. A. Allard (4♂). Lancaster Co., 1

mi N Holtville, 27 September 1937, H. R. Roberts (7♂, 10F); 2 mi S Holtville, 27

September 1937, H. R. Roberts (1♂). Montgomery Co., Ashbourne, 27 October 1906, B.

Long (2♂, 1♀). Philadelphia, Chestnut Hill, 13-18 September 1903, M. Hebard (1♂,

1♀); Mount Airy, 4 August 1911, H. Fox (1♀).

South Carolina: Chesterfield Co., Carolina Sandhills NWR, 34º31'19"N 80º13'27"W,

17 September 2013, J. G. Hill (4♂, 4♀). Florence Co., between Mars Bluff & Peedee

River, 3 September 1945, Rehn (2♂); Florence, 6 September 1911, Rehn & Hebard (7♂,

2♀). Greenville Co., 6 mi S Greenville, 22 August 1939, B.E. Montgomery (1♂).

Greenwood Co., Ware Shoals, 14 October 1932, F. Sherman (1♂, 1♀). Jasper Co., 5 mi

N Ridgeland, 2 September 1945, Rehn (1♀). Lexington Co., Batesburg, 24 August 1930,

L. D. Tuthill (1♀). Oconee Co., SW base Stump House Mountain, 4 mi NW Walhalla,

20 August 1952, Rehn (1♂); CCC Camp F2, 3 October 1936, F. Sherman (1♂). Pickens

Co., Rocky Bottom near Sassafras Mountain, 16 September 1938, F. Sherman (1♂);

Clemson College, 23-24 November 1934, F. Sherman (1♂, 2♀); 13 October 1928, F.

Sherman; 5 October 1932, D. Dunavan (1♀). Saluda Co., 3 mi N Edgefield Co Line, 9

September 1952, T.H. Hubbell (8♂, 3♀). South Dakota: Lincoln Co., Canton, 28

August 1923, H.C. Severin (3♀). Union Co., Elk Point, 15 September 1925, H.C. Severin

(3♀). Yankton Co., Yankton, 8 August 1924, H. C. Severin (3♂, 2♀). Tennessee:

Carroll Co., Huntingdon, 1 September 1924, T. H. Hubbell (1♂). Cumberland Co.,

Crossville, 21 August 1924, T. H. Hubbell (1♂); Dorton, 8 September 1924, T. H.

Hubbell (4♂, 4♀); near Pleasant Hill, 28 August 1962 (8♂, 4♀). Davidson Co.,

Couchville Glade N. A., 36°06’04”N 86°31’46”W, 18 September 2009, J. G. Hill (8♂,

4♀). 20 mi N Nashville, 6 September 1932, T. H. Hubbell (2♂, 13♀). Decatur Co., Bath

Springs, 25 August 1924, T. H. Hubbell (9♂, 7♀); Carroll Cabin Barrens N.A.,

35°27’45”N 88°03’18”, 13 September 2012, J. G. Hill (2♂, 2♀). Dickson Co., 4 mi W

Dickson, 2 September 1924, T. H. Hubbell (5♂, 2♀). Fentress Co., Allardt, 4-6

September 1924, T. H. Hubbell (18♂); 20-23 August 1924, T. H. Hubbell (9♂).

Hamilton Co., Chattanooga, 24 August 1903, A. P. Morse (1♂). Hardin Co., 3.7 mi W

Clifton, 35°21’44”N 88°03’48”, 13 September 2012, J. G. Hill (4♂, 3♀). Knox Co.,

University of Tennessee Farm 46, 6 December 1951, H. B. Reed (1♂). Madison Co.,

Jackson, 26 August 1924, T.H. Hubbell (1♀). Marshall Co., Wilson School Rd. N. A.,

35°39’43”N 86°47’44”W, 1 August 2010, J. G. Hill (1♂); 7 September 2010, J. G. Hill

(2♂, 2♀). Maury Co., Still House Hollows N. A., 35°27’45”N 87°16’05”W, 10

September 2010, J. G. Hill (2♀). Rutherford Co., Flat Rock Cedar Glade, 35°51’31”N

86°17’44”W, 9 September 2010, J. G. Hill (5♂, 2♀). Wilson Co., Cedars of Lebanon

State Park, 36°04’52”N 86°18’55”W, 18-19 September 2009, J. G. Hill (2♂, 1♀); Lane

Farm N. A., 36°01’55”N 88°22’07”, 9 September 2010, J. G. Hill (3♂, 4♀); Vesta Cedar

Glade N. A., 36°04’36”N 86°23’45”W, 18 September 2009, J. G. Hill (1♂, 3♀).

Virginia: Albemarle Co., Charlottesville, 13 August 1945, T. H. Hubbell (1♀).

Alleghany Co., 8 mi N Covington, 8 September 1939, Rehn & Rehn (1♀). Arlington

Co., Rosslyn, 22 October 1910, Rehn (1♂, 1♀); 22 October 1915, A. N. Caudell (1♂);

Lyon Park, 18 September 1933, H. A. Allard (1♂). Bath Co., Flag Rock Pass, Warm

Springs Mt., 14 September 1952, T.H. Hubbell (1♂). Bedford Co., Blue Ridge Parkway,

~8 mi S Snowden, 21 August 1953, J. A. G. Rehn (3♂, 1♀). Botetourt Co., Blue Ridge

Parkway nr. Blue Ridge Springs, 21 August 1953, J. A. G. Rehn (1♂); 3 mi E Buchanan,

2 September 1946, B. Long (2♂). 3 mi N Covington, 16 August 1916, Hebard (1♂); 2 mi

E Covington, 16 August 1916, Hebard (1♂). Charlottesville, 31 October 1915, (1♂).

Craig Co., Craig Creek, N McAfee’s Gap, 1 September 1946, Rehn (1♂); Meadow

Creek, nr. Looney, 3 mi SW Newcastle, 1 September 1946, Rehn (5♂, 2♀). Dinwiddle

Co., E McKenney, 13 October 1941, B. Long (1♂). Essex Co., 1 mi SE Dunnsville,

37°52’N 76°48’W, 23 September-19 October 1993, D. R. Smith (3♂, 1♀);

Tappahannock, 19 September 1915, (1♂); 2 October 1915, (1♂). Fairfax Co., Falls

Church, 3 September 1900, (1♂); 22 October 1901, (2♂, 1F); Great Falls, 27 October

1918, Rehn & Hebard (2♂, 2♀); 12 September 1920, A. N. Caudell (3♂); nr. Mt.

Vernon, 5 September 1920, (1♂); Chain Bridge, 4 September 1921, J. R. Malloch (1♂,

1♀); 17 August 1921, A. N. Caudell (1♂); Difficult Run Mt., 31 November 1933,

Bridwell (1♂); nr. Vienna, 3 October 1937, A. B. Gurney (1♂). Floyd Co., 7 mi NW

Floyd, 31 August 1946, Rehn (2♂, 1♀); Blue Ridge Parkway Intersection, 6 mi S Floyd,

Blue Ridge, 31 August 1946, Rehn (3♂). Frederick Co., Rest, 25 August 1916, Hebard

(2♀); 3 mi NW Nain, 23 August, 1953, Rehn (1♂). Giles Co., Mt. Lake Biological

Station, 26 August 1945, T.H. Hubbell (15♂, 6♀). Greensville Co., NW Taylors Mill

Pond, 21 September 1939, B. Long (1♀). Three Creek, NW Emporia, 21 August 1940, B.

Long (1♂). Isle of Wight Co., Burwell Bay, James River, below Rushmere, 27 August

1940, B. Long (1♂). Henrico Co., Richmond, 12 September 1934, L.K. Gloyd (1♂, 3♀).

Lee Co,. 12 October 1961, W. Tarpley (1♂). Montgomery Co., Riner, 31 August 1946,

Rehn (4♂); Upper Roanoke River, nr. McDonald’s Mill, 9 mi NE Blacksburg, 1

September 1946, Rehn (1♀). Loudoun Co., Ashburn, 12 September 1937, A. B. Gurney

(2♂). New Kent Co., Chickahominy River, Lanexa, 13 October 1939, B. Long (3♀);

York River, N Holly Forks, 13 October 1939, B. Long (1♂, 1♀). Montgomery Co.,

Blacksburg, 15 August 1946, T. H. Hubbell (3♂, 1♀). Northumberland Co., Black Pond,

23 September 1921, A. N. Caudell (2♂). Orange Co., Cedar Mt., 3 September 1949, A.

B. Gurney (2♂, 2♀). Prince William Co., Bull Run Mts., October-November 1934, H. A.

Allard (3♂, 2♀); 14 September 1935, H. A. Allard (1♂); 29 September 1935, H. A.

Allard (1♂); High Point, Bull Run Mt., 3 September 1939, H. A. Allard (1♂); 7

September 1941, H. A. Allard (2♂); 19 October 1941, H. A. Allard (1♂). Princess Anne

Co., Virginia Beach 7 September 1903, A. P. Morse (1♂, 1♀). Richmond Co., Naylors,

15-17 September, Henry Fox (1♂). Roanoke Co., Catawba Ridge, 8 September 1939,

Rehn & Rehn (1♂, 1♀); Roanoke, 6 September 1907, A. P. Morse (1♀). 23 September

1936, (1♂); 5 mi SW Catawba, 1 September 1946. Rockbridge Co., Natural Bridge, 12-

13 September 1907, B. Long (1♀). Rockingham Co., Skyline Drive, 15 mi S Panorama,

13 August 1933, H. A. Allard (1♂); Panorama, 7 October 1934, H. A. Allard (1♂); 4 mi

W Broadway, 22 August 1953, J. A. G. Rehn (1♂, 1♀). Shenandoah Co., Woodstock, 25

August 1916, Hebard (2♂, 3F); Pugh’s Run, 2 September 1940, A. B. Gurney (1♂, 1♀);

Short Mt. nr. Mt. Jackson, Massanutten Mts., 10-12 October 1940, Allard & Gurney

(4♂). Staunton, 16-24 August 1918, A. N. Caudell (1♂). Suffolk Co., N Whitemarsh

School, 12 October 1939, B. Long (2♂, 3♀); SE Cherry Grove Rd., 15 October 1939, B.

Long (4♂, 4♀). Surrey Co., W Claremont, 7 September 1941, B. Long (1♀); Hog Island,

James River, B. Long (1♀); Virginia Beach, 7 September 1903, A. P. Morse (5♂, 3♀).

Sussex Co., Jarratt, 21 August 1940, B. Long (1♂). Wythe Co., Wytheville, 5 September

1903, A. P. Morse (1♂). Washington D. C.: Dalecarlia Reservoir & Outlet, 22 August

1915, W. L. McAtee (1♂); Soldier Home, 28 October 1917, (1♂); Washington D.C, 15

September 1883, C. V. Riley (1♂); August 1911, L. Bruner (3♂, 2♀); September 1929,

A. N. Caudell (1♂, 1♀). West Virginia: Hampshire Co., 2.2 mi E Capon Bridge, 27

August 1969, I.J. Cantrall (1♂, 1♀); Ice Mt., 14-15 September 1935, H. A. Allard (1♂).

Hardy Co., Moorfield, 26 September 1937, H. A. Allard & A. B. Gurney (4♂, 2♀).

Wisconsin: Grant Co., 6 Serptember 1962, O. Warnke (1♂, 2♀); Wyaluing State Park,

T6N R6W Sec 17, 20-26 August 1975 (2♂).

Distribution

Melanoplus scudderi is distributed over a large portion of the eastern U.S., from eastern South Dakota east to the Atlantic coast and south to the Gulf Coast. The species is absent from the extreme southeastern coastal plain and the Mississippi River Valley

(Fig 3A).

Habitat

Melanoplus scudderi prefers open weedy or brushy habitats. Originally, these

grasshoppers were probably denizens of barrens, glades, forest openings, prairies, and

savannahs across its range. Human disturbances have dramatically increased the amount

of suitable habitat for this species, such that Melanoplus scudderi now also occupies

power/gas line right of ways, uncultivated fields, and other ruderal environments with a

mix of shrubs, grasses, and forbs such as Ambrosia, Helianthus, and Solidago.

Remarks

During the course of this revision, a long-winged (macropterous) form of M.

scudderi was discovered (Fig 4. A-B). The wings of the macropterous individuals usually reach the end of the abdomen in females, and are slightly shorter than the abdomen in males. However, the wings are still somewhat thickened and pad-like. Thus, these wings may not be functional for flight. Macropterous individuals are apparently rare. Of the 1,669 specimens examined in this study, only 22 are macropterous. The macropterous form appears to be more common in females as out of the specimens

examined, only three were male. Macropterous individuals were encountered in the field

at two locations (TN, Davidson Co., Couchville Glade, and MS Lowndes Co, Black

Prairie WMA). At Couchville Glade only one macropterous individual (a female) was

observed, whereas other specimens observed were the typical brachypterous form. At

Black Prairie WMA, 15 of the 38 specimens captured (39%) in one hour were

macropterous, of which only one was male. Though these individuals have long-wings, it

does not appear they are capable of flight. In the field, macropterous individuals jumped

to escape capture, similar to brachypterous individuals. Several macrocropterous individuals were captured and brought back to lab where several attempts were made to induce them to fly, none of which were successful, nor did these individuals ever spread

their wings while leaping similar to other macropterous Melanoplus species. Several

macropterous specimens in the UMMZ collection were labeled as being collected at

“light” and “under street light”. This may be an indication that some individuals may be

able to fly, and the lights drew them in. However, brachypterous individuals of other

Melanoplus species have been observed leaping to blacklights from adjacent vegetation.

Figure 1 Melanoplus scudderi.

A. dorsal view of terminalia, B. lateral view of terminalia, C. dorsal view of phallic complex, D. lateral view of phallic complex, E. dorsal view of aedeagus, F. lateral view of aedeagus, G. dorsal view of epiphallus, H. lateral view of epiphallus, I. caudal view of epiphallus, J. habitus. a = ancorae, as = aedeagal sheath, br = bridge, c= cercus, dv = dorsal valve of aedeagus, f= furcula, l = lophus, lp= lateral plate, sap= supra anal plate, sgp = subgenital plate, vv= ventral valve of the aedeagus.

Figure 2 Variant of the male genitalia of M. scudderi found east of the Appalachian Mountains.

A. dorsal, B. lateral.

Figure 3 Locality of specimens of Melanoplus scudderi that were examined in this study.

Figure 4 Long-winged forms of Melanoplus scudderi (A-B) and Melanoplus Mississippi (C).

Melanoplus folkertsi new species (Figs. 5A-J, 8)

Male Measurements (mm)

(n = 12) Body length 18 – 22.7 (mean = 20.7); pronotum length 3 – 4.8 (mean =

4.3); tegmen length 3.7 – 5.6 (mean = 4.6); hind femur length 10 – 11,8 (mean = 11.1);

cerci length 1.2 - 1.3 (mean = 1.2); basal width of cerci 0.6 - 0.8 (mean = 0.7); mid-cercal

width 0.4 - 0.5 (mean = 0.4); cerci apex width 0.2- 0.3 (mean = 0.2).

Female Measurements (mm)

(n = 12) Body length 19.2 – 25.6 (mean = 23.8); pronotum length 5.1 – 6.0 (mean

= 5.5); tegmen length 4.3 – 6.6 (mean = 5.7); hind femur length 11.2 – 15.3 (mean =

13.1).

Diagnosis

Differing from other species in the shape of the internal male genitalia. (Fig. 5 C-

F) In dorsal view, the dorsal valves are subapically curved laterally and apically recurved posteriorly. The ventral valves are constricted at their apices, and extend approximately to the laterally curved portion of the dorsal valves. Laterally, the valves project posteriorly at a 45º angle dorsally, with the subapical curved portion of the dorsal valves slightly curved ventrally (Fig. 5C-F). This species is very similar to M. mississippi, but is distinguished from that species by not having the dorsal valves as strongly curved subapically, having longer ventral valves, and their separate geographic distributions (Fig

6).

Holotype

♂; ALA[BAMA], Baldwin Co., Splinter Hill Bog, 31º01'30"N 87º41'07"W, 5

October 2013, J.G. Hill, open hillside adjacent to bog. Deposited in the USNM.

Paratype

Alabama: same data as holotype (1♀).

Other specimens examined

Alabama: Barbour Co., Barbour Co. Mgt. Area, August 1962, M.E. Dakin (2♂).

Butler Co., 1 min N. Perote, 4 October 1962, M.E. Dakin (2♂, 1♀); Greenville, 23

August 1962 (1♂). Clark Co., 0.5 mi N Tombigbee River on U.S. 43, 12 September

1958, I.J. Cantrall (1♂); 4.3 mi W Gosport on US 43, 14 September 1954, T.H. Hubbell

& I.J. Cantrall (3♂, 6♀); 4.4 mi N Grove Hill on U.S. 43, 12 September 1958, I.J.

Cantrall (8♂, 2♀); 6.7 mi Grove Hill on U.S. 43, 12 September 1958, I.J. Cantrall (5♂,

4♀); Grove Hill, 12 September 1958, I.J. Cantrall (1♂, 1♀). Conecuh Co., .8 mi NE

Evergreen on U.S. 31, 13 September 1958, I. J. Cantrall (1♂); 1.7 mi S jct. HWY 31 and

54, 13 September 1954,T.H. Hubbell & I. J. Cantrall (23♂, 22♀); 2.3 mi N Sepulge River on U.S. 31, 21 August 1951, I.J. Cantrall (2♂); 2 mi N Castleberry, 14 September 1954,

T.H. Hubbell & I. J. Cantrall (2♂, 1♀). Covington Co., 4.8 mi NNW Red Level,

31º27’44”N 86º38’52”W, 12 September 2013, J.G. Hill (1♂). Dallas Co., 0.5 mi NW

Cahaba, 32°16’44”N 87°07’33”W, 19 September 2006, J.G. Hill (1♂). Escambia Co.,

1.2 mi N. Brewton, 26 August 1963, M.E. Dakin (2♂, 1♀); 4.3 mi E Brewton, 5 October

1962, M.E. Dakin (1♀); Range, 31°14’49”N 87°11’41”W, 12 November 2007, J.G. Hill

& M.E. Dakin (1♀). Geneva Co., 3 mi E Hartford on HWY 52, 30 September 1959, M.

E. Dakin (3♂, 4♀). Monroe Co., Frisco City, 24 September 1959, M. E. Dakin (1♂, 1♀).

Florida: Gadsden Co., 2 mi S Chattahoochee, 30°40’16”N 84°50’04”W, 14 October

2010, J.G. Hill (2♂, 3♀). Georgia: Decatur Co., 3.34 mi SW Faceville, 30°44’09”N

84°42’20”W, 14 October 2010, J.G. Hill (1♂, 1♀); Silver Lake WMA, 30°49’29”N

84°45’19”W, 14 October 2010, J.G. Hill (1♂, 2♀); Dougherty Co., 11.6 mi W Albany, 5

October 1945, T. H. Hubbell (1♂). Early Co., Blakely, 5 October 1945, T. H. Hubbell

(1♂); Williams Bluff Pres. 31°12’11”N 85°04’49”W, 15 October 2010, J.G. Hill (1♂,

1♀), 31°22’07”N 85°04’44”W (1♂, 1♀), 31°11’58”N 85°04’43”W (1♀). Terrell Co., 1 mi N Sasser, 8 September 1946, T.H. Hubbell & I.J. Cantrall (2♂).

Etymology

Named in honor of the late Dr. George Folkerts, who possessed a remarkable

passion and knowledge of natural history of the southeastern United States, and who was

an advocate for the conservation of rare and imperiled communities of the southeastern

United States.

Distribution

Melanoplus folkertsi is found in the southern third of Alabama, east of the

Tombigbee River and Mobile Bay to southwestern Georgia and northern Florida (Fig. 8).

Habitat

Melanoplus folkertsi has been collected in a variety of habitats, but is most

commonly found in the understory of longleaf pine savannahs and sandhills. In Gadsden

County, Florida, M. folkertsi was collected along a powerline right of way in tangles of

Ambrosia and Rubrus.

Figure 5 Melanoplus folkertsi.

Melanoplus relictus new species (Figs. 6A-J, 8)

Male Measurements (mm)

(n = 7) Body length 19.5 – 22 (mean = 20.6); pronotum length 4.2 – 4.6 (mean =

4.5); tegmen length 2.5 – 4.7 (mean = 4.0); hind femur length 10.8 – 12.3 (mean = 11.2); cerci length 1.2 - 1.4 (mean = 1.3); basal width of cerci 0.7 (mean = 0.7); mid-cercal width 0.5 - 0.6 (mean = 0.5); cerci apex width 0.2- 0.3 (mean = 0.2).

Female Measurements (mm)

(n = 8) Body length 24 – 26 (mean = 24.7); pronotum length 5.4 – 6.2 (mean =

5.8); tegmen length 5.2 – 6.3 (mean = 5.6); hind femur length 12.3 – 14.5 (mean = 13.8).

Diagnosis

Distinguished from other species in the group by having slightly longer and more slender cerci and the shape the internal male genitalia (Fig. 6C-I), which have the dorsal and ventral valves strongly curved medially, such that they overlap (Fig. 6C-F). This species is most similar to the following undescribed species, but the valves of the aedeagus are more strongly curved medially in M. relictus.

Holotype

♂; FL[ORIDA], Liberty Co., Apalachicola B[luffs] & R[avines] Pres[erve],,

30°29’40”N 84°58’19”W, 14 September 2013, J. G. Hill. Deposited in the USNM.

Paratypes

Same data as holotype (1♂, 2♀).

Other specimens examined

Florida: Liberty Co., T. 2N, R. 7W S.217, “Old Camp Torryea” 17, October

1925, T.H. Hubbell (1♂, 1♀), 5 November 1938 (2♀); 4 September 1954, T.H. Hubbell

& I. J. Cantrall, (7♂, 5♀); Torreya Park 19 Aug 1938 (2♂, 2♀); Apalachicola Bluffs and

Ravines Preserve, 30°29’40”N 84°58’19”W, 13-14 September 2013, J. G. Hill (1♂, 2♀).

Georgia: Thomas Co., Wade Tract Preserve, 30°45’34”N 84°00’03”W, 14 September

2013, J. G. Hill (4♂, 8♀).

Etymology

The name of this species refers to the relictual environment of the of the

Apalachicola region where this grasshopper occurs.

Distribution

At present, this species is known only from sandhills just east of the Apalachicola

River and southern Thomas County, Georgia. (Fig. 8).

Habitat

At Apalachicola Bluffs and Ravines Preserve M. relictus was found to be uncommon in thick growth of Callicarpa americana, Rhus, Rubrus, and Smilax along the edge of a ravine. Cantrall (unpublished field notes) described collecting the species at Old

Camp Torreya in a similar situation. At the Wade Tract Preserve in Georgia, M. relictus was very abundant in clumps of low growing oaks and grasses in an old-growth longleaf savannah.

Figure 6 Melanoplus relictus.

Melanoplus coreyi new species (Fig., 7A-J, 8)

Male Measurements (mm)

(n = 6) Body length 19 – 22.2 (mean = 20.3); pronotum length 4.4 – 5.2 (mean =

4.7); tegmen length 4 – 4.9 (mean = 4.5); hind femur length 10.8 – 11.8 (mean = 11.2);

cerci length 1.3 - 1.4 (mean = 1.4); basal width of cerci 0.7 (mean = 0.7); mid-cercal

width 0.5 - 0.6 (mean = 0.5); cerci apex width 0.2- 0.3 (mean = 0.2).

Female Measurements (mm)

(n = 3) Body length 21.1 – 23.4 (mean = 22.2); pronotum length 5.5 – 6.1 (mean

= 5.8); tegmen length 5.5 – 6.2 (mean = 5.8); hind femur length 12.2 – 13 (mean = 12.5).

Diagnosis

Most similar to M. relictus, in having more slender cerci and medially curved

dorsal and lateral valves. However, the valves of M. coreyi (Fig. 7C-F) are not as strongly

curved as in M. relictus (Fig. 7C-F). The two may also be separated by the distribution, as

M. relictus, at present is known only from the Apalachicola region of the Florida and

south Georgia and M. coreyi, at present, is known only from the Fall Line sandhills of

North Carolina (Fig 8). Specimens from the type locality are slightly darker that those

from other locales, apparently blending in with the frequently burned substrate of the

area.

Holotype

♂; N[ORTH] C[AROLINA], Moore Co., Weymouth Woods N[atural].A[rea].,

35°08’49”N 79°22’09”W, 6 October 2012, J. G. Hill, collected in sandhill. Deposited in

the USNM.

Paratypes

North Carolina: Same data as type (1♂, 2♀), 13 August, 2009.

Other specimens examined

North Carolina: Cumberland Co., Fort Bragg, 35º11'28"N 79º05'04"W, 16

September 2013, J.G. Hill, J.E. Corey, and S. Hall (2♂, 8♀). Hoke Co., Stedman Fish

Hatchery, 12 September 1952, T.H. Hubbell et al (19♂, 18♀). Moore Co., Southern

Pines, 18 August 1923, T.H. Hubbell (1♂), 12 September 1952, T.H. Hubbell et al (1♂);

near Vass, 4 September 1945, Rehn (1♂). Richmond Co., McKinney Lake Fish

Hatchery, 35º00'59"N 79º37'58"W, 16 September 2013, J.G. Hill (1♂, 3♀); Sandhills

Game Lands, 35.02060 N 79.54823W, 6 October 2012, J.E. Corey and J.C. Beane (1♂,

1♀); 35º01'14"N 79º32'55"W, 16 September 2013, J.G. Hill & J.E. Corey (1♂). Scotland

Co., Sandhills Game Lands 34.98428 N 79.54176W, 6 October 2012, J.E. Corey and J.C.

Beane (1♂).

Etymology

This species is named in honor Ed Corey, North Carolina state Park biologist,

who aided in documenting the distribution of the species.

Distribution

At present, this species appears endemic to the Fall Line sandhills of North

Carolina (Fig. 8)

Habitat

Thus far, M. coreyi has been collected only in sandhills of North Carolina. The habitat at the type locality had a longleaf pine (Pinus palustris) overstory with a 46

wiregrass (Aristida spp.) understory, which had been burned earlier in the year. The specimens from Richland and Scotland counties were collected on a mesic hillside with wiregrass and scrub oaks. At Fort Bragg, M. coreyi was collected along the edge of

hillside seepage bog in sandhills.

Figure 7 Melanoplus coreyi.

Melanoplus muscogee new species (Fig. 8, 9A-J)

Male Measurements (mm)

(n = 14) Body length 15.5 – 20.3 (mean = 18.9); pronotum length 4.0 – 4.8 (mean

= 4.5); tegmen length 4.0 – 5.5 (mean = 4.7); hind femur length 10.4 – 12.0 (mean =

10.9); cerci length 1.1 - 1.3 (mean = 1.2); basal width of cerci 0.5 - 0.6 (mean = 0.6); mid-cercal width 0.4 - 0.5 (mean = 0.5); cerci apex width 0.2- 0.3 (mean = 0.2).

Female Measurements (mm)

(n = 14) Body length 19.6 – 25.0 (mean = 23.2); pronotum length 5.2 – 6.3 (mean

= 5.7); tegmen length 4.7 – 7.2 (mean = 5.85); hind femur length 12.0 – 14.3 (mean =

13.2).

Diagnosis

Distinguished from other species in the group by the shape the internal male genitalia (Fig. 9C-I). In dorsal view, the dorsal valves appear slightly dorso-laterally flattened, taper to a point, and in the apical two-thirds are curved dorsally and diverge laterally; the ventral valves are bent along the lateral edge to cover the ventral lateral margin of the dorsal valves, taper to point similar to the dorsal valves, but are slightly shorter than the dorsal valves (Fig. 9C-F).

Holotype

♂; GA. [Georgia], Talbot Co., Black Jack Crossing Pres[erve]., 32°35’06”N

84°30’23”W, 18 September 2013, J.G. Hill, Collected in Fall Line Sandhill. Deposited in the USNM.

Paratypes

Same data as holotype (5♂, 4♀).

Other specimens examined

Georgia: Houston Co., Oaky Woods WMA, 32°30’11”N 84°24’25”W, J.G. Hill

(4♂, 4♀). Macon Co., Montezuma, 19-20 September, 1923, H. Fox (8♂, 5♀). Marion

Co., 2 mi SE Juniper, 32°30’41”N 84°35’13”W, 10 October 2010, J.G. Hill, M.E. Dakin

(2♂, 2♀). Talbot Co., Black Jack Crossing Pres. 32°35’06”N 84°30’23”W, 25 August

2010, J.G. Hill (1♂). Taylor Co., Black Creek, N.A., 32°33’53”N 84°24’25”W, 11

October 2010, J.G. Hill (1♂); Fall Line Sandhills N.A., 32°34’37”N 84°17’25”W, 25

August 2010, J.G. Hill (1♂). Twiggs Co., 6.12 mi E Bonaire, 32°32’20”N 84°29’05”W,

J.G. Hill (1♂); 4.7 mi S Bullard on U.S. 129, 22 September 1946, T.H. Hubbell (1♂).

Etymology

This species is named in honor of the Muscogee people, a Native American tribe, who once inhabited the region where this grasshopper occurs.

Distribution

Melanoplus muscogee is found along the Fall Line in western Georgia (Fig 8).

Habitat

This species has been found inhabiting sandhills and the edge of pine woods. In the sandhill habitat, it often occurred in areas with denser growths of various grasses and forbs and tangles of Smilax and Rubus.

Figure 8 Distribution of M. folkertsi, M. muscogee, M. coreyi, M. relictus, M. quercicola, and M. davisi.

Figure 9 Melanoplus muscogee.

Melanoplus quercicola (Hebard) (Fig. 8, 10A-J)

Eotettix quercicola Hebard 1918: 156; Otte 1995: 361.

Melanoplus davisi quercicola, Blatchley 1920: 397.

Melanoplus quercicola, Otte 1995: 391; Hill 2009: 32 Genitalia illustration.

Male Measurements (mm)

(n = 15) Body length 18 – 22.5 (mean = 20.5); pronotum length 4.2 – 5 (mean =

4.7); tegmen length 3.3 – 4.8 (mean = 4.06); hind femur length 10.3 – 12.3 (mean =

11.5); cerci length 0.7 - 1.2 (mean = 1); basal width of cerci 0.5 - 0.6 (mean = 0.6); mid- cercal width 0.3; cerci apex width 0.1- 0.2 (mean = 0.1).

Female Measurements (mm)

(n = 10) Body length 24 – 28 (mean = 25.3); pronotum length 6 – 7.2 (mean =

6.6); tegmen length 5 – 6.7 (mean = 5.8); hind femur length 13 – 16.2 (mean = 14.5).

Diagnosis

Distinguished from other species in the group by the male cerci, which are triangular, and have the apical fourth curved medially (Fig. 10A-B), and in the shape of the internal male genitalia. In dorsal view, the valves are subequal in length, triangular in shape and are curved ventrally in the apical third (Fig. 10C-I). This species is most similar to Melanoplus davisi, but differs from that species in having narrower cerci that bow medially at the apical tip, and with much shorter valves, as well as a distinct geographic distribution. (Fig. 8).

Figure 10 Melanoplus quercicola.

Holotype

♂; Woodville, FLA [Florida], IX, 1, 1915 (R[ehn]. & Hebard].). Deposited in the

ANSP.

Specimens examined

Florida: Alachua Co., Archer, 5 October 1924, F.W. Walker, (7♂); Gainesville,

15-17 Nov. 1959 (1♂). Franklin Co., Carrabelle, 2-3 September 1915, Rehn and Hebard

(1♂), 2 October 1945, T.H. Hubbell (6♂, 1♀). Lake Co., Weirsdale, 17 August 1938 (2

♂). Leon Co. Apalachicola N.F., 30º 21' 31"N 84º 15' 35"W, 5 August 2011, J.G. Hill

(12♂, 3♀). Liberty Co., 20 August 1944 (7 ♂); Alum Bluff, 6 Nov. 1931, T.H. Hubbell,

(5♂, 3♀); Apalachicola Bluffs and Ravines Pres., 30º 29'33"N 84º 58'54"W, 3 August

2011, J.G. Hill (1♂, 1♀); NE of 30.455018º N 84. 970686ºW, 18 Nov. 2009, D.T.

Almquist (2♂); Camp Torreya, Twp 2N, R 7W, 17 September 1925, T.H. Hubbell (4♂,

3♀). Marion Co., Ocala, 19-20 September 1917, Rehn and Hebard (2♂, 1♂). Georgia:

Bryan Co., .4 mi E Canoochee Rvr, 32º08'53"N 81º46'13"W, 30 July 2006, J. G. Hill

(1♂). Emanuel Co., 14.5 mi S Swainsboro, 11 October 2010, J.G. Hill (1♂); Ohoopee

Dunes N.A., 32º34'31"N 82º26'30"W, 8 October 2007, J.G. Hill (2♂, 2♀); 32º34'32"N

82º26'33"W, 8-9 October 2007 (4♂, 2♀); 32º32'15"N 82º27'40"W, 24 July 2006 (1♀); 9

October 2007 (2♂, 1♀), 32º32'15"N 82º27'40"W 5-6 October 2007 (5♂, 9♀).

Distribution

Melanoplus quercicola is distributed across most of southern Georgia and large

portion of northern Florida to eastern Bay County (Fig. 8).

Habitat

Found in sandhills ranging from inland sand dunes to longleaf pine savannah. At many sites this species is most abundant in clumps of low growing oaks.

Remarks

See comments about intermediate individuals between M. davisi and M. quercicola under the remarks section of M. davisi.

Melanoplus davisi (Hebard) (Fig. 8, 11A-J)

Eotettix davisi Hebard 1918: 153; Otte 1995: 361. .

Melanoplus davisi, Blatchley 1920: 396;1995: 391.

Male Measurements (mm)

(n = 6) Body length 19.1 – 22.6 (mean = 20.6); pronotum length 4.5 – 4.8 (mean

= 4.6); tegmen length 3.7 – 4.8 (mean = 4.1); hind femur length 11.3 – 12.8 (mean =

11.8); cerci length 0.7 - 1.0 (mean = .9); basal width of cerci 0.5- 0.6 (mean = 0.6); mid- cercal width 0.3 - 0.3 (mean = 0.3); cerci apex width 0.1.

Female Measurements (mm)

(n = 6) Body length 24.6 – 26.7 (mean = 25.5); pronotum length 6.5 – 6.8 (mean

= 6.6); tegmen length 4.4 – 5.7(mean = 5.1); hind femur length 13 – 16 (mean = 14.7).

Diagnosis

Distinguished from other species in the group by the shape of the male cerci, which are more triangular (Fig 11A-B), and in the shape of the internal male genitalia. In dorsal view, the dorsal valves that are subequal in length, project dorsally and posteriorly

at a 45 degree angle, parallel each other, and are curved ventrally in their apical third.

Additionally the valves have tiny projections that often give them a serrate appearance

(Fig. 11C-F). This species is most similar to Melanoplus quercicola (Hebard), but differs

from that species in having broader cerci and longer valves, and a distinct geographic distribution (Fig 6).

Holotype

♂; DeFuniak Springs, Fl[ORID]a. VIII, 30 1915. (R[ehn] & H[ebard]) .

Deposited in the ANSP.

Other Specimens Examined

Alabama: Covington Co., 4.8 mi NNW Red Level, 31º 27’44”N 86º 38’52”W, 12

September 2013, J.G. Hill (1♂). Florida: Bay Co., 10 mi WNW Youngstown, 30º

25’40”N 85º 35’25”W, 13 September 2013, J.G. Hill (1♂, 1♀). Okaloosa Co., Delaco,

11-12 August 1935, T.H. Hubbell and G.G Hubbell (22 ♂, 9♀); Niceville, 6 Nov., 1939,

T.H. Hubbell (2♂, 2♀). Walton Co., DeFuniak Springs, 30 August 1915, Rehn and

Hebard (4♂, 2♀), 3 August, 1925, T.H. Hubbell (1♂); 11 Sept 1929, T.H. Hubbell (1♂,

1♀); .5 mi NW DeFuniak Springs, 14 Nov. 1938, I. J. Cantrall (3♂, 3♀), Nokuse

Plantation, 30º 32’36”N 85º 55’47”W, 13 September 2013, J.G. Hill (7♂, 10♀).

Figure 11 Melanoplus davisi.

Distribution

Melanoplus davisi is distributed in the panhandle of Florida and extreme southern

Alabama to western Bay County, FL (Fig. 8).

Habitat

Sandhills, particularly in areas with many scrubby oaks.

Remarks

Intermediate individuals between M. davisi and Melanoplus quercicola can be

found in Bay County, Florida where the two species apparently come into contact. These

individuals are externally similar to M. davisi, but have internal genitalia similar to M. quercicola.

Melanoplus mississippi new species (Figs. 12A-J, 13)

Measurements (mm)

(n = 12) Body length 19.5 – 23.5 (mean = 21.1); pronotum length 4.5 – 5.0 (mean

= 4.7); tegmen length (brachypterous) 4.7 – 6.4 (mean = 5.5) (macropyerous) 10.9; hind

femur length 11.0 – 12.0 (mean = 12.5); cerci length 1.3 - 1.4 (mean = 1.3); basal width of cerci 0.8 – 1.3 (mean = 1.0); mid-cercal width 0.5 - 0.6 (mean = 0.6); cerci apex width

0.2- 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 12) Body length 22.5 – 26.5 (mean = 24.6); pronotum length 5.0 – 6.0 (mean

= 5.7); tegmen length (brachypterous) 6.0 – 9.0 (mean = 7.2), (macropterous) 14 – 15.5

(mean = 14.5); hind femur length 12.2 – 14.5 (mean = 13.3).

Diagnosis

Differing from other species in the group primarily in the shape of the internal male genitalia (Fig. 12C-F); In dorsal view, the dorsal valves taper to a point, and are curved laterally subapically and apically recurved posteriorly and the ventral valves extend approximately two-thirds length of dorsal valves. In lateral view, the valves project posteriorly, with the subapical curved portion of the dorsal valves slightly curved ventrally (Fig. 12C-F). The genitalia are very similar to M. folkertsi, but are distinguished from that species by having the dorsal valves more strongly curved subapically, having shorter ventral valves, and their separate geographic distributions

(Fig. 13). A long winged form is known in this species.

Holotype

♂; AR[KANSAS], Cross Co., Village Creek St[ate].P[ark]., 35º09'20"N

90º43'09"W, 24 September 2013, J.G. Hill. Deposited in the USNM.

Paratypes

Same data as holotype (1♂, 4♀).

Other specimens examined

Illinois: Alexander Co., Cairo (Peddie Barge Landing Park), 15 September 1954,

T. H. Hubbell & I. J. Cantrall (16♂, 5♀); 2.8 mi N Gale, 30 August 1958, I. J. Cantrall

(6♂); 3.4 mi S Mill Creek on IL-127, 31 August 1958, I. J. Cantrall (1♂, 3♀); 0.2 mi S

Olive Branch on IL-3, 30 August 1958, I. J. Cantrall (14♂, 12♀); Cache, 14 October

1932, Frison & Ross (5♂). Kentucky: Fulton Co., N edge of Fulton, 15 September

1954, T. H. Hubbell & I. J. Cantrall (3♂, 23♀). Hickman Co., 5.2 mi NW Clinton on

KY-58, 15 September 1954, T. H. Hubbell & I. J. Cantrall (8♂, 7♀). Henderson Co., 14

September 1952, D. C. Eades (1♀). Madison Co., Berea, 20 September 1946, T. H.

Hubbell (1♀). Louisiana: LaSalle Par., 1 mi NW jct. US-4 & LA-8, 24 October 1957,

Cantrall, Cohn & Eades (1♂). Lincoln Par., 2.2 mi E center of Choudrant, 9 September

1958, I. J. Cantrall (19♂, 15♀); 0.6 mi E center of Choudrant, 9 September 1958, I. J.

Cantrall (3♀); 2.0 mi W Choudrant on US-80, 9 September 1958, I. J. Cantrall (5♂, 2♀);

3.5 mi E Choudrant, 9 September 1958, I. J. Cantrall (7♂, 5♀). Ouachita Par., 3.5 mi E

Calhoun on US-80, 8 September 1958, I. J. Cantrall (6M, 9♀); 1.4 mi W Calhoun on US-

80, 8 September 1958, I. J. Cantrall (7♂, 7♀). Mississippi: Tallahatchie Co., Tutwiler,

34º00'41"N 90º25'36"W, 19 September 2012, J.G. Hill (4♂, 3♀). Missouri: Butler Co.,

On US-67 3.6 mi SW jct. US-60 and 67 at Poplar Bluff, 26 August 1954, T. H. Hubbell

& I. J. Cantrall (30♂, 25♀). Cape Girardeau Co., 1.5 mi SW Cape Girardeau, 25 August

1954, T. H. Hubbell & I. J. Cantrall (28♂, 18♀). Crawford Co., Onandaga Cave, 7

September 1930, T. H. Hubbell (1♀). Mississippi Co., Bertrand, 23 October 1957,

Cantrall, Cohn & Eades (10♂, 16♀). St. Louis Co., Washington University Tyson

Research Tract, ~3 mi W Valley Park, 11 October 1969, I. J. Cantrall (2♂). Stoddard

Co., 0.8 mi W Grayridge on US-60, 26 August 1954 T. H. Hubbell & I. J. Cantrall (5M,

6♀); 3.5 mi W Dexter, 26 August 1954, T. H. Hubbell & I. J. Cantrall (11♂, 8♀).

Tennessee: Dyer Co., (1♂). Lauderdale Co., 3 mi S Ripley, 31 August 1924, T.H.

Hubbell (1♀). Shelby Co., Memphis, 2 August 1924, T.H. Hubbell (1♀).

Etymology

Named for the Mississippi River along, which the species is distributed.

Distribution

Melanoplus mississippi is distributed in the Mississippi River floodplain from

Illinois south to Louisiana and Mississippi. (Fig.13).

Habitat

Melanoplus mississippi has been collected in ruderal areas with some shrubs, mixed grasses and forbs. At Helena, Arkansas, the species was collected along a roadside in a stand of Sorghum halepense and Solidago spp.. At the type locality, M. mississippi

was collected along the edge of a hardwood forest and an open field among Ambrosia

and Solidago. Cantrall’s field notes describe the species being collected in pastures on bluffs above the Mississippi river with short grasses, various forbs, and scattered trees; as well as in open second growth hardwood forests with thin leaf litter and Solidago, grasses, Helanthus, Vitis, and Toxicodendron radicans growing in the understory.

Remarks

The series of 58 specimens from Cape Girardeau, MO contained four

macropterous female specimens of M. mississippi (Fig. 4C). See additional comments

about macropterous individuals under the M. scudderi remarks.

Figure 12 Melanoplus mississippi.

Figure 13 Distribution of Melanoplus mississippi.

Melanoplus arkansas new species (Fig. 14A-J, 17)

Male Measurements (mm)

(n = 13) Body length 20.0 – 23.5 (mean = 22.1); pronotum length 4.2 – 5.2 (mean

= 4.8); tegmen length 3.2 – 5.7 (mean = 4.7); hind femur length 10.3 – 12.5 (mean =

11.6); cerci length 1.0 - 1.3 (mean = 1.1); basal width of cerci 0.8 – 0.9 (mean = 0.8);

mid-cercal width 0.5 - 0.6 (mean = 0.6); cerci apex width 0.3.

Figure 14 Melanoplus arkansas.

Female Measurements (mm)

(n = 13) Body length 21.5 – 25.5 (mean = 23.3); pronotum length 5.3 – 6.0 (mean

= 5.8); tegmen length 4.8 – 7.1 (mean = 5.8); hind femur length 11.8 – 14.4 (mean =

13.4).

Diagnosis

Distinguished from other species in the group by the shape the internal male genitalia (Fig. 14C-I). In dorsal view, the dorsal valves taper to a point, are weakly bowed laterally (Fig 14C and E); the ventral valves parallel the dorsal valves along most of their length laterally. In a lateral view, the valves project posteriorly and are bowed ventrally; the ventral valves are widened dorso-ventrally, such that they largely obscure the view of the dorsal valves (Fig. 14C-F).

Holotype

♂; AR[KANSAS]., Logan Co., Mt. Magazine S[tate].P[ark]., 35º10'28"N

93º38'45"W, 16 September 2011, J.G. Hill, Collected in open glade like area

Paratypes

Same data as holotype (2♂, 2♀).

Other specimens examined

Arkansas: Conway Co., 2.3 mi S Arkansas River on Hwy 9., 8 September 1958,

I.J. Cantrall (3♂). Franklin Co, Cherokee Prairie Natural Area, 35º20'07"N 94º02'28"W,

27 September 2013, J.G. Hill (4♂, 5♀). Logan Co., 2.1 mi S Booneville on Ark Hwy 23,

29 August 1954, Hubbell & Cantrall (5♂, 1 ♀); 5 mi S Booneville, 35º08'28"N

93º00'10"W, 16 September 2011, J.G. Hill (2♂, 1♀); Mt. Magazine, 29 August 1905, 66

A.P. Morse (20♂, 7♀); 28 August 1954, Hubbell & Cantrall (1♂); Mt. Magazine State

Park, 35º10'28"N 93º38'45"W, 20 September 2012, J.G. Hill (3♂, 1♀), (2♀); Short

Mountain Creek, 1 mi S Paris, 25 August 1954, Hubbell & Cantrall (4♂). Yell Co., 0.5 mi E foot of Mt. Nebo on Ark. Hwy 155, 28 August 1954, Hubbell & Cantrall (1♂); 2 mi

SW Chickalah, 12 August 1939, Rehn and Rehn (2♂), Dardanelle, 31 August 1905, A.P.

Morse (2♂), Summit of Mt. Nebo, 28 August 1954, Hubbell & Cantrall (2♂); Mt. Nebo

State Park, 35º13'01"N 93º15'30"W, 20 September 2012, J.G. Hill (2♂, 1♀). Oklahoma:

Le Flore Co., 0.7 mi S Arkansas River bridge on US 59, 6 September 1958, I.J. Cantrall

(2♂, 3♀); 8 mi W Spiro, 35º14'10"N 94º32'14"W, 21 Sept, 2012, J.G. Hill (1♂); Poteau

35º06'25"N 94º40'14"W, 21 September 2012, J.G. Hill (1♂, 3♀). Muskogee Co., 3 mi S

Porum, 35º18'44"N 95º15'05"W, 15 September 2011, J.G. Hill (1♂).

Etymology

Named for the Arkansas River Valley, where the species is apparently endemic to

the, south side of the river.

Distribution

Melanoplus arkansas is distributed in Arkansas Valley physiographic region,

south of the Arkansas River, in western Arkansas and eastern Oklahoma (Fig. 17).

Melanoplus ouachita new species (15A-J, 17)

Male Measurements

(n = 12) Body length 19.0 – 24.0 (mean = 22.0); pronotum length 4.2 – 5.3 (mean

= 4.9); tegmen length 3.7 – 6.0 (mean = 4.7); hind femur length 10.7 – 12.3 (mean =

11.8); cerci length 0.9 - 1.1 (mean = 1.0); basal width of cerci 0.7 – 0.9 (mean = 0.8);

mid-cercal width 0.7 - 0.9 (mean = 0.8); cerci apex width 0.3 - 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 12) Body length 21.2 – 28.2 (mean = 25.2); pronotum length 5.3 – 6.6 (mean

= 6.0); tegmen length 4.5 – 7.8 (mean = 6.3); hind femur length 12.4 – 15.4 (mean =

14.2).

Diagnosis

Distinguished from other species in the group by having sub-quadrate shaped

cerci (Fig. 15A-B), and by the unique shape of the internal male genitalia (Fig. 15C-I). In

dorsal view, the dorsal valves are dorso-laterally flattened basally becoming cylindrical

and tapering to a point apically, are bowed and are apically curved dorsally. The ventral

valves are slightly broader than the dorsal valves, apically curved dorsally, taper to a

point, and are subequal in length to the dorsal valves. In lateral view the valves project

posteriorly. Specimens from Le Flore County, Oklahoma have the apical tips of the

dorsal valves recurved dorsally and laterally. This species is most similar to Melanoplus baronei and M. cohni, but differs from those species by having shorter dorsal and ventral valves, much thinner dorsal valves, and a distinct geographic distributions (Fig. 17).

Holotype

♂; AR[KANSAS]., Montgomery Co., Ouachita N[ational] F[orest], 34º36’09”N

93º34’05”W, 26 September 2013, J.G. Hill, Collected in roadside glade.

Paratypes

Same data as type (1♂, 3♀). 68

Other specimens examined

Arkansas: Conway Co., Petit Jean State Park, 35º06'55"N 92º56'16"W, 5-7

August 2008, J.G. Hill (5♂, 5♀); 35º06'47"N 92º56'50"W, 20 September 2013, J.G. Hill

(2♂, 3♀). Garland Co., Ouachita National Forest, 34º42’11”N 93º22”47”W, 26

September 2013, J.G. Hill (2 ♂, 3♀). Perry Co., 2.2 mi N Casa, 35º03'09"N 93º03'07"W,

20 September 2012, J.G. Hill (1 ♂, 5♀). Polk Co., 1 mi S Cove on Buffalo Creek, 29

August 1954, T.H. Hubbell & I.J. Cantrall (11♂, 11♀); Wolf Pinnacle summit, 29 August

1954, T.H. Hubbell & I.J. Cantrall (2♂, 2♀); 1.5 mi S Polk-Scott Co. line on US Hwy 71,

29 August 1954, T.H. Hubbell & I.J. Cantrall (10 ♂, 11 ♀). Pulaski Co., 18.6 mi E Perry

– Pulaski Co. Line on Hwy 10, 8 September 1958, I.J. Cantrall (5♂, 9♀); 4.2 mi N

Pulaski – Saline Co. Line on U.S. 167, 8 September 1958, I.J. Cantrall (4♂, 2♀). Scott

Co., 1 mi N Boles on Ark Hwy 71, 29 August 1954, T.H. Hubbell & I.J. Cantrall (10♂,

3♀); jct. AR Hwy 23 & U.S. Hwy 71, 29 August 1954, T.H. Hubbell & I.J. Cantrall

(2♀). Sevir Co., 2.8 mi N. DeQueen on U.S. Hwy 71, 29 August 1954, T.H. Hubbell &

I.J. Cantrall (3♂, 1♀); 6 mi W DeQueen on U.S. Hwy 70, 29 August 1954, T.H. Hubbell

& I.J. Cantrall (2♂, 1♀). Yell County, Ola, 35º02'26"N 93º12'29"W, 20 September 2012,

J.G. Hill (1 ♂). Oklahoma: Le Flore Co., Howe I.T., 5 August 1905, A.P. Morse (1 ♂);

Ouachita National Forest, 34º41'00"N 94º38'10"W, 27 September 2013, J.G. Hill (2♂,

1♀); Wister Lake, 34º56'00"N 94º42'46"W, 27 September 2013, J.G. Hill (1♂, 1♀);

Figure 15 Melanoplus ouachita.

Etymology

Named for the Ouachita Mountains, where this species is apparently endemic.

Distribution

Melanoplus ouachita is distributed over most of the Ouachita Mountains of west-

central Arkansas to southeastern Oklahoma (Fig. 17).

Habitat

At the type locality. M. ouachita was collected in a small glade surrounded by

Juniperus and scattered clumps of S. scoparium and an Echinacea sp. At other locales

the species was collected on roadsides among Lespedza and mixed grasses. Cantrall’s

field notes describe several of the localities where he collected M. ocuachita as being

grassy openings with Ambrosia, Andropogon, Smilax and at the summit of Wolf Pinnacle

in scrubby oak, dwarf hickory with large openings of Danthonia with patches of

Helianthus and Solidago. At Wister Lake the species was collected on an open hillside with short grasses and Rubus, and at the top of the hill along the edge of an open post oak forest.

Melanoplus cohni new species (Figures 16A-J, 17)

Male Measurements

(n = 12) Body length 19.8 – 24.2 (mean = 22.2); pronotum length 4.5 – 5.5 (mean

= 5.1); tegmen length 4.2 – 5.9 (mean = 5.0); hind femur length 10.3 – 13.2 (mean =

11.9); cerci length 1.0 - 1.3 (mean = 1.1); basal width of cerci 0.8 – 1.0 (mean = 0.9);

mid-cercal width 0.6 - 0.8 (mean = 0.7); cerci apex width 0.3 – 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 12) Body length 22.3 – 28.0 (mean = 25.4); pronotum length 5.3 – 6.5 (mean

= 6.0); tegmen length 4.9 – 7.2 (mean = 6.0); hind femur length 12.7 – 15.9 (mean =

14.0).

Diagnosis

Distinguished from other species in the group by having triangular shaped cerci

(Fig. 16A-B), and in the shape the internal male genitalia (Fig. 16C-I). In lateral view the valves project posteriorly and dorsally at approximately a 45-degree angle, and are strongly curved dorsally in the apical half. The dorsal valves taper to a point apically, and are slightly bowed, and in in some specimens are apically curved laterally. The ventral valves, are approximately half a long as the dorsal valves and parallel the dorsal valves (Fig.16C-I). Melanoplus cohni is most similar to Melanoplus ouachita and M. baronei, a new species described below, but has much longer and strongly apically curved dorsal and ventral valves than those species, and by their distinct geographic distribution (Fig. 17).

Holotype

♂; OK[LAHOMA]., 3.2 mi N Wilburton, 34º57'39"N 95º21'16"W, 21 September

2012, J.G. Hill, collected on edge of forest and prairie along rodside.

Paratypes

Same data as holotype (5 ♂, 2♀).

Other specimens examined

Oklahoma: Latimer Co., 7 mi E Hartshorne, 17 August 1951, T.J. Cohn (3 ♂,

4♀); Wilburton, 27 August 1905, A.P. Morse (1 ♂). Le Flore Co., Howe, 5 August 1905,

A.P. Morse (1 ♂). McCurtain Co., 1.6 mi W Idabel on US Hwy 70, 30 August 1954, T.H.

Hubbell & I.J. Cantrall (27 ♂, 25 ♀); 2.4 mi W Idabel in US Hwy 70, 30 August 1954,

T.H. Hubbell & I.J. Cantrall (3 ♂, 5♀). Pushmataha Co., 5 mi N Finley, 8 September

1950, M.J.D. White (1 ♂); Kiamichi River Bridge in US 271 N of Antlers, 30 August

1954, T.H. Hubbell & I.J. Cantrall (5 ♂, 3♀).

Etymology

Named in honor of the late Dr. Theodore “Ted” Cohn for his numerous efforts in furthering Orthopteran research.

Distribution

Melanoplus cohni appears to be endemic to the Ouachita Mountains of southeastern Oklahoma (Fig. 17).

Habitat

At the type locality, M. cohni was collected in a roadside prairie with

Schizachryium scoparium and Lespedeza along the edge of mixed hardwood/pines forest.

Cantrall’s field notes describe most of the localities where he collected specimens of this species in a similar manner, but mentions Ambrosia, Liatris, Rubus, and Solidago as characteristic forbs. At 2.4 miles west of Idabel, Cantrall collected this species in “very dry post oak woods”.

Figure 16 Melanoplus cohni.

Figure 17 Distribution of M. arkansas, M. ouachita, M. cohni, M. seltzerae, M. baronei, and M. decurvus.

Melanoplus seltzerae new species (Figs 17, 18A-J)

Male Measurements

(n = 8) Body length 19.1 - 23.2 (mean = 21.3); pronotum length 4.5 – 5.5 (mean =

5.1); tegmen length 4.3 – 6.0 (mean = 5.1); hind femur length 11.0 -12.2 (mean = 11.7);

cerci length 1.0 - 1.2 (mean = 1.131); basal width of cerci 0.6 – 1.0 (mean = 0.9); mid-

cercal width 0.7 – 0.9 (mean = 0.8); cerci apex width 0.3 – 0.5 (mean = 0.4).

Female Measurements (mm)

(n = 5) Body length 23.8 – 24.5 (mean = 24.1); pronotum length 5.5 – 6.3 (mean

= 6.0); tegmen length 5.5 – 6.2 (mean = 5.9); hind femur length 12.3 – 14.5 (mean =

13.9).

Diagnosis

Distinguished from other species in the group by the shape of the internal male

genitalia (Fig. 18C-I). In lateral view the valves project dorsally at a 45 degree angle

posteriorly; the dorsal valves taper to a point, and have the apical third curved ventrally;

the ventral valves extend approximately two-thirds length of dorsal valves and are

significantly less sclertotized than the dorsal valves (Fig. 18C-F). The genitalia are

similar to M. texarkana but are distinguished from that species by having the ventral

valves reduced in length and being much less sclerotized and, having thinner and less

strongly curved dorsal valves. In addition their geographic distributions differ (Fig 18 and

Fig.24).

Holotype

♂; ARK[ANSAS]., Saline Co., Middle Fork Barrens N[atural].A[rea].,

34º38’16”N 92º50’31”W, 26 September 2013, J.G. Hill, Collected in shale barren.

Deposited in the USNM.

Paratypes

Same data as type (19♂, 12♀). Garland Co., Hot Springs, 34º33'17"N

93º00'42"W, 22 September 2012, J.G. Hill, (1♂, 1♀).

Etymology

Named in honor of my wife, Jennifer Seltzer, for her encouragement and support

during this revision.

Distribution

At present, M. seltzerae is known only from areas near Hot Springs, Arkansas

(Fig. 17.)

Habitat

At the type locality, M. seltzerae was very abundant in a shale barren where

patches of Croton, Grindelia, Silphium, Smilax, and various grasses were scattered among the bare rock. At Hot Springs, this species was found along the edge of a

hardwood forest among resprouting trees and Rubus.

Figure 18 Melanoplus seltzerae.

Melanoplus baronei new species (17, 19A-J)

Male Measurements

(n = 8) Body length 21.0 – 23.0 (mean = 21.8); pronotum length 4.5 – 5.2 (mean

= 4.9); tegmen length 4.3 – 5.5 (mean = 4.8); hind femur length 11.5 – 12.3 (mean =

11.9); cerci length 1.0 - 1.1 (mean = 1.1); basal width of cerci 0.7 – 1.0 (mean = 0.9); mid-cercal width 0.7 - 0.8 (mean = 0.8); cerci apex width 0.3 (mean = 0.3).

Female Measurements (mm)

(n = 5) Body length 24.0 – 25.5 (mean = 24.8); pronotum length 5.8 – 6.2 (mean

= 6.0); tegmen length 5.5 – 6.3 (mean = 6.0); hind femur length 13.0 – 14.4 (mean =

13.7).

Diagnosis

Distinguished from other species in the group by having more quadrate shaped cerci (Fig. 19A-B) and by the shape the internal male genitalia (Fig. 19C-I). In dorsal view, the dorsal valves taper to a point apically, are somewhat flattened medially, and are bowed with the apices nearly attingent. The ventral valves are broader than the dorsal valves, taper to a point apically, and are subequal in length to the dorsal valves. In lateral view the valves are straightened and project posteriorly and dorsally at approximately a

45º angle. This species is most similar to Melanoplus ouachita, but differs from that species by having more elongate doral and ventral valves, and a majority of specimens with dark patches the on first few abdominal tergites.

Figure 19 Melanoplus baronei.

Holotype

♂; AR[KANSAS]., Perry Co., Ouachita N[ational].F[orest]., 34º51'25"N

92º46'02"W, 26 September 2013, J.G. Hill, Collected along roadside with ruderal vegetation. Deposited in the USNM.

Paratypes

Same data as holotype (6♂, 4♀).

Other specimens examined

Arkansas: Perry Co., 0.6 mi E jct. state Hwy 9 &10 on 10, 8 September 1958, I.J.

Cantrall (3♂, 1♀).

Etymology

Named for Dr. John Barone, for his work with grasslands in the southeastern

United States, and for his years of friendship and mentorship.

Distribution

At present, M. baronei is known only from eastern Perry County, Arkansas (Fig.

17).

Habitat

At the type locality M. baronei was collected in a large stand of Lespedeza between the road and a mixed pine – hardwood forest.

Melanoplus latus Morse New Status (Figs. 20A-J, 21A-D, 22J)

Melanoplus scudderi latus Morse 1906:122; Morse & Hebard 1915:103

(Lectotype designation). 81

Male Measurements (mm)

(n = 17) Body length 18.5 – 23.8 (mean = 21.2); pronotum length 4.5 – 5.2 (mean

= 4.8); tegmen length 3.6 – 5.2 (mean = 4.7); hind femur length 11.0 – 12.0 (mean =

11.6); cerci length 0.8 - 1.2 (mean = 1.0); basal width of cerci 0.6 - 0.9 (mean = 0.8); mid-cercal width 0.6 - 0.8 (mean = 0.7); cerci apex width 0.2- 0.3 (mean = 0.3).

Female Measurements (mm)

(n = 17) Body length 22.5 – 24.5 (mean = 23.3); pronotum length 5.1 – 6.3 (mean

= 5.6); tegmen length 4.7 – 6.5 (mean = 5.8); hind femur length 12.1 – 14.0 (mean =

13.1).

Female Measurements (mm)

(n = 17) Body length 22.5 – 24.5 (mean = 23.3); pronotum length 5.1 – 6.3 (mean

= 5.6); tegmen length 4.7 – 6.5 (mean = 5.8); hind femur length 12.1 – 14.0 (mean =

13.1).

Figure 20 Melanoplus latus.

Diagnosis

Distinguished from other species in the group by having more quadrate shaped cerci (Fig. 20A-B), and by the shape the internal male genitalia (Fig. 20C-I). In lateral view the valves project posteriorly and are slightly bowed dorsally. In dorsal view, the dorsal valves taper to a point apically, are bowed and apically are recurved medially, and slightly dorsally; the ventral valves are broader than the dorsal valves, are subequal in length to the dorsal valves, and are broadly rounded apically. Appearance of the aforementioned aedagal valves slightly varies throughout the range of the species (Fig.

21A-F). Specimens from central Kansas and Nebraska have valves that apically are more strongly bowed laterally curved ventrally (Fig. 21E-F). This species is most similar to

Melanoplus ouachita, but differs from that species by having longer and more robust dorsal and ventral valves.

Lectotype

Bonita, TX. 14 Aug 14, 1905. Deposited in the ANSP.

Figure 21 Variation in the genitalia of M. latus.

A-B. Dorsal and lateral view, Tarrant Co., Texas; C-D. dorsal and lateral view, Commanche Co., Oklahoma, and E-F. dorsal and lateral views, Otoe Co., Nebraska.

Other specimens examined

Kansas: Butler Co., 1910, R.H. Beamer. Riley Co., Manhatten, 17 March 1932,

H.C. Smith (1♀); 1 mi NE Ogden, 39º07'24"N 96º41'37"W, 14 September 2011, J.G. Hill

(1♂, 1♀). Nebraska: Otoe Co., Dunbar, 25 August 1958, I.J. Cantrall (13♂, 6 ♀).

Lancaster Co., Lincoln, 3 October 1929 (1♂); Roca, 31 October 1933 (1♂); Roca, 31

October 1933 (1♂). Oklahoma: Caddo Co., 3 mi E Hydro, 15 August 1939, Rehn &

Rehn (1 ♂). Carter Co., 5 mi E Ardmore, 30 August 1939, Rehn & Rehn (1♂). Choctaw

Co., between Hugo and Grant, 30 August 1939, Rehn & Rehn (1♂). Cleveland Co.,

Norman, 1927, M. J. Brown (1♂). Comanche Co., Ft. Sill Military Res., 27 October - 1

December 1918, T.H. Hubbell (4♂, 6♀). Grady Co., Chickasha, 13 August 1929 (1♂).

Love Co., .5 mi W Thackerville, 33º47'33"N 97º08'02"W, 23 85

September 2012, J.G. Hill (3♂, 7♀). McCurtain Co., 12 October 1933 (1♂); Beavers

Bend State Park, 29 August 1954, T.H. Hubbell & I.J. Cantrall (11♂, 3♀). Potawatomie

Co., Shawnee, 26 August 1915, A.P. Morse (1♀). Texas: Concho Co., 8 mi S Eden, 13

September 1951, Rehn (1♂). Cooke Co., Myra, (1♂). Coryell Co., 7 mi W Moody, 8

July 1955, T.J. Cohn (2♂). Gillespie Co., 14 mi E Fredricksburg, 6 Aug 1955, T.J. Cohn

& J.R. Hilliard (1♂). Jones Co., 3 mi E Hawley, 9 October 1958, J.R. Hilliard (2♂).

Montague Co., 4 mi NW St. Jo, 33º44'06"N 97º34'44"W, 23 September 2012, J.G. Hill

(2♂, 1♀). Parker Co., 8 mi NE Cresson, 19, August 1951, T.J. Cohn (3♂). Tarrant Co.,

19, July 1927 (1♂); Lakeside, 32º50'31"N 97º28'32"W, 23 September 2012, J.G. Hill

(1♂, 1♀); Sagamore Hill, 27 September 1912, Rehn & Hebard (4♂, 3♀). Taylor Co.,

Abilene, J.R. Hilliard (1♂).

Distribution

Melanoplus latus is distributed from central Texas northward to central Nebraska

(Fig. 34)

Habitat

Prairies, plains, and open forests with mixed grasses and forbs.

Figure 22 Distribution of Melanoplus latus.

Melanoplus decurvus new species (Figs. 17, 23A-J)

Male Measurements (mm)

(n = 9) Body length 20.1 – 23.4 (mean = 21.2); pronotum length 4.5 – 5.4 (mean

= 4.9); tegmen length 4.0 – 5.8 (mean = 4.9); hind femur length 11.2 – 12.4 (mean =

11.6); cerci length 0.9 - 1.0 (mean = 1.0); basal width of cerci 0.5 - 0.8 (mean = 0.7);

mid-cercal width 0.6 - 0.8 (mean = 0.7); cerci apex width 0.3 (mean = 0.3).

Female Measurements (mm)

(n = 9) Body length 23.0 – 25.8 (mean = 24.3); pronotum length 5.5 – 6.0 (mean

= 5.7); tegmen length 4.7 – 6.5 (mean = 5.3); hind femur length 13.0 – 13.5 (mean =

13.8).

Diagnosis

Distinguished from other species in the group by having more quadrate shaped

cerci (Fig. 23A-B) and in the shape of the internal male genitalia (Fig. 23C-I). In lateral

view the valves are strongly decurved. The dorsal valves taper to a point apically, are

slightly curved laterally and are apically recurved medially. The ventral valves are

truncated apically. Melanoplus decurvus is most similar to M. latus, but differs by

having the valves strongly decurved, unlike M. latus, where the valves project

posteriorly, and by their distinct geographic distributions (Fig. 17).

Holotype

♂; OKLAHOMA Atoka Co., 2 mi E Lane, 30 August 1954, T.H. Hubbell & I.J.

Cantrall (1♂). Deposited in the USNM.

Paratypes

Same data as holotype (5♂, 9♀).

Other specimens examined

Oklahoma: Atoka Co., 5 mi E. Farris, 6 August 1955, Rehn (1♂); 7.5 mi SE

Atoka, 6 August 1955, Rehn (1♂). Bryan Co., Caddo, 8 August 1905, A.P. Morse (4♂,

1♀); 1.8 mi N Colbert on U.S. 75, 31 August 1954, T.H. Hubbell & I.J. Cantrall (1♂).

Etymology

From the Latin “de” meaning down and “curvus” meaing curved in reference to the decurved nature of the male genitalia.

Distribution

At present, M. decurvus is known only from Atoka and Bryan Counties in the south central plains ecoregion (Fig. 17).

Habitat

At the holotype locality, field notes of Cantrall state that the specimens were collected in a valley of a small dry creek in small patches of Betula nigra that was approximately 1.5 m tall, and Cantrall describes the habitat at the 1.8 mi north of Colbert locality as “ a swale in open country with patches of Andropogon, Glyceria, and herbs beside a grove of Quercus spp. and M. pomifera.”

Figure 23 Melanoplus decurvus.

Melanoplus ozarkensis new species (Fig. 24A-J, 23)

Male Measurements

(n = 12) Body length 20.6 – 22.8 (mean = 21.6); pronotum length 4.2 – 5.1 (mean

= 4.7); tegmen length 4.0 – 5.5 (mean = 4.57; hind femur length 10.8 – 12.7 (mean =

11.5); cerci length 1.2 - 1.4 (mean = 1.3); basal width of cerci 0.6 – 0.9 (mean = 0.8); mid-cercal width 0.4 - 0.7 (mean = 0.5); cerci apex width 0.2- 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 12) Body length 22.0 – 25.0 (mean = 24.1); pronotum length 5.3 – 6.3 (mean

= 5.9); tegmen length 4.8 – 6.5 (mean = 5.7); hind femur length 12.5 – 15.0 (mean =

13.2).

Diagnosis

Distinguished from other species in the group by the shape the internal male genitalia (Fig. 24C-I). In dorsal view, the dorsal valves taper to a point and are bowed laterally (Fig. 24C and E), and the ventral valves parallel the dorsal valves, and extend approximately to the apically recurved portion of the dorsal valves (Fig. 24C-F). In lateral view, the valves project posteriorly and are curved ventrally and are apically recurved dorsally.

Holotype

♂; AR[KANSAS], Boone Co., Baker Prairie, 36º14’28”N 93º08’04”W, 25

September 2013, J.G. Hill, Collected in tall grass prairie. Deposited in the USNM.

Paratypes

Same data as holotype (2♂, 3♀). 91

Figure 24 Melanoplus ozarkensis.

Other specimens examined

Arkansas: Benton Co., 4 mi E, Gravette, 5 August 1958, T.J. Cohn (2♂); 4 mi S

Pea Ridge, 4 August 1958, T.J. Cohn (1♂). Boone Co., 2.6 mi N Burlington, 3

September 1958, I.J. Cantrall (4♂, 12♀); 2.7 mi S jct, St Hwy 68 & 16, 4 September

1958, I.J. Cantrall (2♂, 4♀); 3.5 mi S Harrison, 27 Aug 1954, T.H, Hubbell & I.J.

Cantrall (11♂, 10♀). Carroll Co., 2.9 mi W. Green Forest on US 62, 3 September 1958,

I.J. Cantrall (7 ♂, 1 ♀); 1.8 mi E. jct. US 62 & state Hwy 21 in Eureka Springs, 3

September 1958, I.J. Cantrall (7 ♂, 3♀). Crawford Co., 1.1 mi N. jct. 64 & 71 on 71 at

Alma, 6 September, 1958, I.J. Cantrall (8♂, 5♀); Winfrey, 2 September 1952, D. C.

Eades (2♂); East slope of Gaylord Mtn., 2 September 1952, D.C. Eades (1♂); Ozark

National Forest, 35º37'11"N 94º24'15"W, 21 September 2012, J.G. Hill (2♂, 1♀).

Franklin Co., 3 mi N. Cass on state Hwy 23, 6 September, I.J. Cantrall (9♂, 5♀).

Johnson Co., 2 mi W Clarksville, 6 September 1958, I.J. Cantrall (11♂, ♀3). 1.5 mi N

Ozone, 7 September, 1958, I.J. Cantrall (2♂); Ozark National Forest, 35º35'11"N

93º15'01"W, 16 September 2011, J.G. Hill (1♂); 35º10'28"N 93º38'45"W (1♂),

35º34'59"N 93º14'42"W (1♂). Madison Co., 3.4 mi N jct.St Hwy 23 & 16 on 23, 6

September 1958, I.J. Cantrall (2♂, 2♀); jct. St Hwy 12 & 45, 3 September 1958, I.J.

Cantrall (1♂). Newton Co., Mill Creek x Hwy 7, 36º06’36”N 93º08’02”W, 25

September 2013, J.G. Hill (9♂, 7♀); Ozark National Forest, 35º44’02”N 93º05’37”W, 25

September 2013, J.G. Hill (1♂, 1♀). Washington Co., Winslow 5 September 1905, A.P.

Morse (9♂, 2♀); Fayetteville, 5 September 1905, A.P. Morse (3♂, 1♀); N. edge of

Fayetteville on U.S. 71, 3 September, 1958, I.J. Cantrall (26 ♂, 20♀); Winslow, 2

September 1905, A.P Morse (6♂). Kansas: Brown Co., 0.2 mi S Hiawatha on U.S. 159,

26 September 1958, I. J. Cantrall (2♂). Douglas Co. Lawrence, 1937, H.C. Beamer

(1♂); 1.8 mi S Kansas River at jct U.S. 40 & Hwy 10. 25 August, 1958, I. J. Cantrall

(4♂, 3♀); 6 mi NW Kansas River at Lawrence, 26 September 1958, I. J. Cantrall (1♂,

3♀); Lawrence, 20 October 1910 (1♂); top of high ridge on E side of Kill, Creek, 20

September 1946, H. S. Wallace (4♂, 3♀). Jefferson Co., 1.9 mi N Oskaloosa, 26 August

1958, I. J. Cantrall (4♂, 3♀). Miami Co., 2.4 mi W Louisburg on Hwy 68, 27 August

1958, I. J. Cantrall (1♂, 1♀). Missouri: Cass Co., 2.1 mi E. Harrisonville, 27 August

1958, I.J. Cantrall, (2♂). Johnson Co., 4.1 mi W Leeton on St. Hwy 2, 27 August 1958,

I.J. Cantrall (3♂, 2♀). McDonald Co., 1.6 mi S Lanagan on US 71, 5 September 1958,

I.J. Cantrall (2♂); 2.1 mi S Goodman on US 71, 5 Sept 1958, I.J. Cantrall (2♂). Miller

Co., 2.5 mi E Kaiser on Hwy 42, 29 August 1958, I.J. Cantrall (2♂, 3♀). Newton Co., .3

mi S jct. US 71 & 60, 5 September 1958, I.J. Cantrall (2♂). Saline Co., 1 mi E Marshall,

28 August 1958, I. J. Cantrall (1♂, 3♀). Taney Co., 2.5 mi N Branson on US 65, 13

September 1958, I.J. Cantrall (2♂). Webster Co., .8 mi W Rogersville on US 66, 3

September 1958, I.J. Cantrall (2♂); 3 mi E Rogersville, 3 September 1958, I.J. Cantrall

(5♂, 3♀). Wright Co., 1 mi W Mountain Grove on U.S. 60, 2 September 1958, I.J.

Cantrall (8♂, 10♀). Oklahoma: Delaware Co., .8 mi E Leach, 4 September 1958, I.J.

Cantrall (5 ♂). Mayes Co., 200 yds. W Grand River, 4 September 1958, I.J. Cantrall

(6♂, 9♀); 1mi E Grand River, 4 September 1958, I.J. Cantrall (1♂); 3.1 mi N Pryor on

US 69, 4 September 1954, I.J. Cantrall (2♂). Ottawa Co., 3.3 mi E Fairland, 5 September

1958, I.J. Cantrall (9♂, 4♀). Sequoyah Co., 200 yds N bridge over Arkansas River on

US 59, 6 September 1958, I.J. Cantrall (3♂, 2♀). 2.8 mi S jct. US 64and 59 on 59, 6

September 1958, I.J. Cantrall (2♂), 3.3 mi E Arkansas River on US 60, 5 September

1958, I.J. Cantrall (2♂) 3.9 mi NW N Edge Arkansas River on US 64, 6 September 1958,

I.J. Cantrall (4♀), Sallisaw, 35º31'17"N 94º47'47"W, 21 September 2012, J.G. Hill (8♂,

3♀); 4.2 S Sallisaw, 35º23'23"N 94º48'55"W, 21 September 2012, J.G. Hill (4♂, 1♀); 14 mi NNE Sallisaw, 5 August 1958, T.J. Cohn (1 ♂). Wagoner Co., Cornell, 14 September

1935, T.H. & G.G. Hubbell (1♂).

Etymology

Named in reference to the Ozark Mountains, over which the species is widely

distributed.

Distribution

Melanoplus ozarkensis is endemic to the southern Ozark Mountains, north of the

Arkansas River over areas of Arkansas, Missouri Oklahoma; and north along the

Missouri river into eastern Kansas and South Dakota (Fig, 25).

Habitat

Inhabits prairies, glades, forest edges and other open areas with ruderal

vegetation.

Figure 25 Distribution of M. ozarkensis, and M. francoisensis.

Melanoplus francoisensis new species (Figs. 25, 26A-J)

Male Measurements (mm)

(n = 10) Body length 19.2 – 24.0 (mean = 21.2 ); pronotum length 4.0 – 5.2 (mean

= 4.6); tegmen length 3.6 – 6.0 (mean = 5.2); hind femur length 9.0 – 12.2 (mean = 11.0);

cerci length 1.0 – 1.3 (mean = 1.1); basal width of cerci 0.7 – 0.9 (mean = 0.8); mid-

cercal width 0.4 – 0.6 (mean = 0.5); cerci apex width 0.2 – 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 5) Body length 22.8 – 24.2 (mean = 23.5); pronotum length 5.5 – 7.0 (mean

= 6.1); tegmen length 5.8 – 7.5 (mean = 6.0); hind femur length 13.0 – 15.0 (mean =

13.9).

Diagnosis

Differing from other species primarily in the shape of the internal male genitalia

(Fig. 26C-I). In dorsal view the dorsal valves taper to a point and are curved laterally subapically, and the ventral valves extend to the curved portion of the dorsal valves and are truncated apically. In lateral view, the dorsal valves project posteriorly and slightly ventrally, are slightly bowed dorsally, and the apically curved portions are strongly curved ventrally; the ventral valves are slightly bowed, (Fig. 26C-F). The genitalia are very similar to M. scudderi and M. mississippi, but this new species is distinguished by having the genitalia with more bowed in lateral view and in dorsal view with more strongly bowed dorsal valves, strongly truncated ventral valves, along with their separate geographic distributions (Fig 25).

Holotype

♂; MISSOURI: Reynolds Co. Johnson Shut-in Sta. Park, 29 August 1958, I.J.

Cantrall, #35. Deposited in the USNM.

Paratypes

Same data as type (34♂, 4♀).

Other specimens examined

Missouri: Dent Co., Bunker, within 30 yds of Reynolds Co. line, 29 August 1958,

I. J. Cantrall (5♂, 4♀).

Etymology

Named for the Saint Francois Mountains, where the species is apparently

endemic.

Distribution

At present, M. francoisensis is known only from Reynolds and Dent Counties in southeastern Missouri (Fig. 25).

Habitat

Cantrall’s field notes indicate that the specimens from Johnson Shut-In State Park

were collected in a fallow field with “Rubrus, a very small amount of Helianthus, and

some goldenrod”, and those from Bunker from a “roadside at edge of forest in fairly well-

mown grass, ragweed, and scattered Rubrus and poison oak”.

Figure 26 Melanoplus francoisensis.

Melanoplus texarkana new species (Figs. 27A-J, 28)

Male Measurements

(n = 12) Body length 19.5 – 24.0 (mean = 22.0); pronotum length 4.0 – 5.5 (mean

= 4.8); tegmen length 3.8 – 5.5 (mean = 4.7); hind femur length 11.2 – 12.5 (mean =

11.8); cerci length 1.0 – 1.3 (mean = 1.2); basal width of cerci 0. – 1.0 (mean = 0.8); mid- cercal width 0.5 – 0.6 (mean = 0.5); cerci apex width 0.2- 0.3 (mean = 0.3).

Female Measurements

(n = 12) Body length 19.5 – 25.5(mean = 23.3); pronotum length 5.0 – 6.5 (mean

= 5.6); tegmen length 3.8 – 7.0 (mean = 5.3); hind femur length 12.0 – 15.2 (mean =

13.4).

Diagnosis

Distinguished from other species in the group by the shape the internal male genitalia. In lateral view, the valves project posteriorly and dorsally at approximately a

45º angle, and are strongly curved ventrally in their apical third; the dorsal valves taper to a point apically, the ventral valves parallel the dorsal valves along their length (Fig. 27C-

I). Melanoplus texarkana is most similar to the Melanoplus seltzerae, but has the ventral valves fully sclerotized unlike M. seltzerae, The two species have separate geographic distributions (Figs 17 & 28).

Holotype

♂; AR[KANSAS}, Miller Co., Miller County Sandhills Natural Area, 33º11'23"N

94º01'40"W, 28 September 2013, J.G. Hill, Collected in sandhill along powerline.

Deposited in the USNM.

100

Paratype

Same data as holotype (1♂, 3♀).

Other specimens examined

Arkansas: Calhoun Co., 2.3 mi. SW Thornton on US 79, 6 September 1958, I.J

Cantrall (12♂, 11♀); 0.9 mi SW Bearden on U.S. 79, 8. September 1958, I.J Cantrall

(7♂, 5♀). 3.5 mi NE Bearden on US 79, 8 Sept 1958, I.J Cantrall (13♂, 7♀); Thornton, 8

September 1958, I.J. Cantrall (3♂). Columbia Co., Magnolia, 31 August 1939, Rehn &

Rehn (6♂, 1♀). Hempstead Co., Bois De Arc WMA, 33º36'12"N 92º46'42"W, 22

September 2012, J.G. Hill (2♂, 3♀). Miller Co., 6 mi E Texarkana, 29 August 1955,

Rehn (3♂, 1♀); 7 mi E Texarkana, 31 August 1939, Rehn & Rehn (1 ♂). Ouachita Co.,

0.9 mi SW Bearden on US 79, 8 September 1958, I.J. Cantrall (2 ♂); 8.7 mi NE Camden

on US 79, 24 October 1957, Cantrall, Cohn, & Eades (2 ♂). Louisiana: Beauregard Par.,

3.1 mi N Merryville, 2 September 1954, T.H. Hubbell & I.J. Cantrall (2 ♂). Bienville

Par., Arcadia, 20 August 1915, Rehn & Rehn (1♂). Caddo Par., Shreveport, 19 August

1915, Rehn and Hebard (7♂); 22 Dec. 1931, H.H. Beamer (1♂). Jackson Par., 2.5 mi S.

Jonesboro, 24 October 1957, Cantrall, Cohn, & Eades (2♂). Lincoln Par., 0.6 mi E center Choudrant, 9 September 1958, I.J. Cantrall (2♂); 1 mi S Vienna on US 167, 24

Oct 1957, Cantrall, Cohn, and Eades (2♂); 1 mi N DuBach, 31 August 1939, Rehn &

Rehn (1♂); 11.8 mi W Choudrant on US 80, 9 September 1958. I.J. Cantrall (2♂); 2 mi

W Choudrant on US 80, 9 September 1958. I.J. Cantrall (2♂). Natchitoches Co.,

Natchitoches, 16 September, 1955, T.H. Hubbell and G.G. Hubbell (2 ♂); Kisatchie

National Forest, 32º06'45"N 93º01'42"W, 16 August 2006, J.G. Hill, J.A. MacGown

(2♂). Rapides Par. 1.6 mi N Red River, N of Alexandria on US 167, 9 September 1958. 101

I.J. Cantrall (2♂). Winn Par., 2 mi S. center of Winnfield on U.S. 167, 9 September 1958,

I.J. Cantrall (2 ♂). Texas: Palestine Co., 16 August 1915, Hebard (2♂). Camp Co.,

Pittsburg, 9 August 1905, F.C. Bishop (1♂). Cass Co., 3.6 mi N Atlanta, 26 August

1955, Rehn, (2♂); Jefferson, 32º56'30"N 94º22'18"W, 17 September 2011, J.G Hill (2♂).

Jasper Co., 9.2 mi W Jasper on US 190, 2 September 1954, Hubbell & Cantrall (1♂).

Gregg Co., Longview, 18 August 1915, Rehn & Hebard (1♂, 2♀). Hardin Co., Larson

Sandyland Preserve., 30º20'54"N 94º14'13"W, 17 September, 2011, J.G. Hill, (10♂, 7♀);

25 September, 2012, (2♂, 2♀). Henderson Co., 1.7 mi E Trinidad on Texas Hwy 31, 17

Aug 1961, I.J. Cantrall and T.J. Cohn (11♂, 4♀). Lamar Co., Paris, F.C. Bishop (1♂).

Polk Co., 1.9 mi N Trinity River on US 190, 2 Sept 1954, Hubbell & Cantrall (1♂); Big

Thicket Nat. Pres., 30º38'34"N 94º39'41"W, 25 September 2012, J.G. Hill (6♂, 3♀). Red

River Co., 2 mi W Detroit, 30 August 1939, Rehn & Rehn (1 ♂). Sabine Co., Sabine

N.F., 31º10’43”N 93º42’29”W, 21 June 2012, J.G. Hill, P. Loos, (1♂).

Etymology

Melanoplus texarkana, inhabits areas of Texas, Arkansas, and Louisiana. The

word “Texarkana” is a portmanteau using portions of the names of those three states like

the cities of Texarkana, AR and Texarkana, TX.

Distribution

Melanoplus texarkana is distributed in east Texas, southwestern Arkansas and

western Louisiana. (Fig 24).

102

Habitat

Open pine forests, longleaf savannahs, sandhills, and forest edges, areas with ruderal vegetation. At many of the locales where M. texarkana has been collected, it was found amongst stunted or resprouted oaks and a mixture of various grasses and forbs.

103

Figure 27 Melanoplus texarkana.

104

Figure 28 Distribution of Melanoplus texarkana.

Melanoplus taurus new species (Figs. 29A-J, 30)

Male measurements (mm)

(n = 12) Body length 21.2 – 24.0 (mean = 23.1); pronotum length 4.7 – 5.5 (mean

= 5.1); tegmen length 4.3 – 5.5 (mean = 5.0); hind femur length 12.0 – 13.5 (mean =

12.6); cerci length 1.1 – 1.2 (mean = 1.2); basal width of cerci 0.8 – 0.9 (mean = 0.8);

mid-cercal width 0.5 – 0.7 (mean = 0.6); cerci apex width 0.3.

105

Female measurements (mm)

(n = 12) Body length 23.8 – 26.5 (mean = 24.9); pronotum length 5.4 – 6.7 (mean

= 5.9); tegmen length 5.4 – 6.8 (mean = 6.2); hind femur length 13.6 – 15.5 (mean =

14.4).

Diagnosis

Distinguished from other species in the group primarily in the shape of the internal male genitalia (Fig. 29C-I). In dorsal view, the dorsal valves taper to a point and, are curved laterally subapically, and the ventral valves extend to the curved portion of the dorsal valves and are narrowed and slightly curved apically. In lateral view, the dorsal valves project posteriorly and are slightly bowed dorsally, and the ventral valves project dorsally, often arising above the dorsal valves (Fig. 29C-F). The genitalia are similar to

M. scudderi, but are distinguished from that species by having slightly different shaped cerci, distinctly shaped ventral valves that project over the dorsal valves, unlike in M. scudderi in which they lie below the dorsal valves. In addition, the two species have separate geographic distributions (Fig. 30).

Holotype

♂; TX [TEXAS], San Jacinto Co., 1 mi NW Point Blank, 30º45'39"N

95º12'01"W, 24 September 2012, J.G. Hill, Edge of forest and roadside in Schizachyrium,

Rubus, and hardwood resprouts. Deposited in the USNM.

106

Figure 29 Melanoplus tarus.

107

Paratypes

Same data as holotype (3♂, 3♀). Walker Co., Huntsville, 30º42'46"N

95º31'30"W, 30 September 2013, J.G. Hill (3♂, 8♀).

Other specimens examined

Texas: Harris Co., Channelview, 29 December 1969, R. Etheridge (1♂); 3.6 mi.

NW Huntsville on 190, 2 September 1954, Hubbell & Cantrall (1♂); Harrisburg, 12

August 1915, Rehn & Hebard (2♂). San Jacinto Co., 1 mi NW Point Blank, 30º45'39"N

95º12'01"W, 24 September 2012, J.G. Hill (3♂, 2♀). Walker Co., 3.8 mi NE Huntsville on U.S. 190, 2 September 1954, T.H. Hubbell & I.J. Cantrall (6 ♂, 14 ♀), 5 mi NE

Huntsville, 6 October 1968, J.R. Hilliard (9♂, 5♀).

Etymology

Taurus from the Latin for “bull”, which refers to the shape of the dorsal valves of male genitalia that resemble a bull’s horns.

Distribution

Thus far, Melanoplus taurus is known only from Harris, Henderson, San Jacinto,

and Walker Counties in southeastern Texas. (Fig. 30).

Habitat

Melanoplus taurus has been collected on forest edges in mixed grasses, shrubs

and various herbaceous plants.

108

Figure 30 Distribution of Melanoplus optimus, M. latus, M. texarkana, M. texensis, M. irwinorum, M. ottei, and M. tarus.

Melanoplus optimus new species (Fig. 30, 31A-J)

Male measurements

(n = 12) Body length 20.2 – 23.2 (mean = 21.5); pronotum length 4.1 – 4.7 (mean

= 4.5); tegmen length 3.6– 5.5 (mean = 4.5); hind femur length 10.5 – 12.0 (mean =

11.5); cerci length 1.0 – 1.1 (mean = 1.0); basal width of cerci 0.7 – 0.9 (mean = 0.8);

mid-cercal width 0.6 – 0.8 (mean = 0.7); cerci apex width 0.3 – 0.4 (mean = 0.3).

109

Female measurements (mm)

(n = 12) Body length 20.2 – 23.5 (mean = 21.3); pronotum length 4.2 – 5.5 (mean

= 4.8); tegmen length 3.2 – 4.8 (mean = 4.4); hind femur length 11.5 – 12.5 (mean =

12.0).

Diagnosis

Distinguished from other species in the group by having more quadrate cerci (Fig.

31A-B) and distinctive shape of the internal genitalia (Fig 31C-I). In lateral view, the

valves project posteriorly and are curved dorsally in their apical third. In some specimens

the valves curve more strongly such that their apices point back to the basal area of the

valves. The dorsal valves taper to a point apically, and are attingent throughout their

length. The ventral valves are subequal in length to the dorsal valves, are dorso-laterally

flattened basally, dorso-ventrally flattened in the apical third, tapering to a point apically,

and parallel the dorsal valves.

Holotype

♂; TX [TEXAS], Ellis Co., Ennis, 32º18'51"N 96º39'15"W, 28 September 2013,

J.G. Hill, Collected in disturbed prairie in ruderal vegetation. Deposited in USNM

Paratypes

Same data as holotype (9♂, 12♀).

Other specimens examined

Texas: Ellis Co., 0.5 mi N Ennis, 1 September 1954, T.H. Hubbell & I.J. Cantrall

(6♂, 2♀); Waxahachie, 11 November 1931, F. B. Isely (1 ♂).

110

Etymology

Optimus from the Latin for best, in references to the day the type series was collected, which was one of the best days I have had in the field.

Distribution

At present, M. optimus, is known only from Ellis Co. Texas (Fig. 30).

Habitat

At the type locality, M. optimus was collected along the edge of a disturbed

hilltop prairie remnant in a patch of shin-high mixed grasses, Ambrosia, Croton, and

Gaillardia along the edge of a Juniperus sp, Maclura pomifera, and Prosopis woodland.

Cantrall's field notes indicate that their specimens were collected in "waist high to chest

high coarse herbaceous growth, including Solidago, Helianthus, but mostly rosin weed".

111

Figure 31 Melanoplus optimus.

112

Melanoplus ottei new species (Fig. 30, 32A-J)

Male Measurements

(n = 12) Body length 19.5 – 24.0 (mean = 22.1); pronotum length 4.5 – 5.2 (mean

= 4.8); tegmen length 4.0 – 5.7 (mean = 5.9); hind femur length 11.0 – 13.0 (mean =

11.9); cerci length 0.8 - 1.2 (mean = 1.0); basal width of cerci 0.7 – 0.9 (mean = 0.8); mid-cercal width 0.5 - 0.6 (mean = 0.5); cerci apex width 0.2 – 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 8) Body length 23.0 – 27.2 (mean = 24.6); pronotum length 5.5 – 6.0 (mean

= 5.8); tegmen length 5.3 – 6.4 (mean = 5.8); hind femur length 13.8– 15.3 (mean =

14.5).

Diagnosis

Distinguished from other species in the group by having more quadrate shaped cerci (Fig. 32A-B) and by the shape the internal male genitalia (Fig. 32C-I). In lateral view, the valves are relatively short and project posteriorly and dorsally at approximately a 40 degree angle. The dorsal valves taper to a point apically, and are bowed laterally.

The ventral valves are slightly broader at the apices than at the base, cover the dorsal valves laterally, and are subequal in length to the dorsal valves.

Holotype

♂; TX [Texas], Burrnet Co., Longhorn Caverns S.P., 30º40'54"N 98º20'60"W, 29

September 2013, J.G. Hill, Collected in Juniper-oak grassland. Deposited in the USNM.

Paratypes

Same data as holotype (5♂, 5♀). 113

Other specimens examined

Texas: Atascosa Co., 20 mi. S San Antonio, 8 November, 1942, H.R. Roberts

(1♂). Bastrop Co., 6 mi NE Cedar Creek, 26 August 1955, Rehn (1♂); Bastrop State

Park, 30 August1952, T.J. Cohn (2♂), 29 September 1952, M.J.D White (1♂). Bexar Co,

San Antonio, 19 Nov. 1934, L. Seaton (1♂). Blanco Co., Pedernales Falls State Park,

30º18'16"N 98º15'35"W, 29 September 2013, J.G. Hill (1♂, 2♀). Burnet Co., Inks Lake,

10 mi W Burnet, 29 August 1951, T.J. Cohn (2♂); Shovel Mountain, F.G. Schaupp (1♂,

1♀). Coldwell Co., 3-5 mi E Luling, 15 September 1942, H.R. Roberts (1♂); Milano,

15-16 August, 1935, T.H. and G.G. Hubbell (2♂). F

ayette Co., Flatonia. 19-20 August 1920, Rehn and Hebard (9♂, 8♀). Gonzales Co.,

Palmetto State Park, 23 October 1949, H.T. Spielt, 20 August 1952, T.J. Cohn (1♂);

Rutledge Swamp, 2 October 1956, J.R. Hilliard (1♂). Llano Co., Buchanan, 17 mi E

Llano, 11 July 1959, T.J. Cohn (2♂). Travis Co., 16 mi E Austin, 29 August 1956, J.R

Hilliard (1♂); Austin, 20 November 1952, M.J.D. White (1♂); Zilker Park, 15 September

1955, J.R. Hilliard (1♂, 6♀). Williamson Co., 8 mi SE Florence, 26 July 1955, T.J. Cohn

(2♂).

Etymology

Named in honor of Dr. Daniel Otte, for his numerous contributions to Orthoptera

research and for graciously allowing me to revise the Scudderi group.

114

Figure 32 Melanoplus ottei.

115

Distribution

Melanoplus ottei is distributed in central Texas largely on the Edwards Plateau

(Fig 30).

Habitat

At the type locality and at Pedernales Falls State Park, M. ottei, was collected in openings of S. scoparium and Bouteloua curtipendula in a Quercus virginiana/Juniperus/Prosopis woodland.

Melanoplus irwinorum new species (Fig. 30, 33A-J)

Male Measurements (mm)

(n = 12) Body length 21.0 – 23.5 (mean = 21.7); pronotum length 4.5 – 5.1 (mean

= 4.8); tegmen length 3.9 – 5.3 (mean = 4.6); hind femur length 11.5 – 12.6 (mean =

12.0); cerci length 1.0 – 1.2 (mean = 1.1); basal width of cerci 0.7 – 0.8 (mean = 0.8); mid-cercal width 0.4 – 0.6 (mean = 0.5); cerci apex width 0.2 – 0.3(mean = 0.3);

Female Measurements (mm)

(n = 12) Body length 22.0 – 25.4 (mean = 23.9); pronotum length 5.2 – 6.0 (mean

= 5.4); tegmen length 4.8 – 7.0 (mean = 5.7); hind femur length 12.8 – 14.2 (mean =

14.2).

Diagnosis

Distinguished from other species in the group by the shape of the internal male genitalia (Fig. 33C-I). In lateral view, the valves project posteriorly and are curved dorsally subapically and recurved posteriorly apically. The dorsal valves appear cylindrical and taper to a point apically, and are parallel throughout their length. The 116

ventral valves are subequal in length to the dorsal valves, are dorso-laterally flattened, taper to a point apically, and are parallel to the dorsal valves.

Holotype

♂; TX [TEXAS], Jasper Co., Angelina N.F., 31º02’32”N 94º11’16”W, 30

September 2013, J.G. Hill, Collected in sandstone barren. Deposited in the USNM.

Paratypes

Same data as holotype (10♂, 12♀).

Etymology

Named in honor of the Irwin family of the Australia Zoo in Queensland,

Australia, for their global conservation efforts.

Distribution

At present, M. irwinorum is known only from the type locality (Fig. 30).

Habitat

Thus far, Melanoplus irwinorum has only been collected in a Catahoula sandstone

barren in the Angelina National Forest where it was relatively common in open expanses

of Schizachyrium scoparium that were bordered by Pinus palustis, Quercus stellata, and

Quercus marilandica.

117

Figure 33 Melanoplus iriwinorum.

118

Melanoplus texensis Hart New Status (Fig. 30, 34A-J)

Melanoplus scudderi texensis Hart 1906: 158; Frison 1927: 143 (Lectotype designation).

Male Measurements (mm)

(n = 12) Body length 20.2 – 22.8 (mean = 21.7); pronotum length 4.3 – 5.2 (mean

= 4.6); tegmen length 3.8 – 5.5 (mean = 4.7); hind femur length 11.0 – 12.3 (mean =

11.6); cerci length 0.9 – 1.1 (mean = 1.0); basal width of cerci 0.5 – 0.7 (mean = 0.7); mid-cercal width 0.3 – 0.4 (mean = 0.4); cerci apex width 0.2- 0.3 (mean = 0.3).

Female Measurements (mm)

(n = 12) Body length 20.7 – 25.0 (mean = 23.5); pronotum length 4.8 – 6.0 (mean

= 5.3); tegmen length 4.0 – 6.0 (mean = 5.2); hind femur length 12.2 – 14.0 (mean =

13.0).

Diagnosis

Distinguished from other species by the shape of the internal male genitalia (Fig.

34C-I). In this species, the valves are greatly shortened and in lateral view, project posteriorly and dorsally at a 45° angle. The dorsal valves appear cylindrical and taper to a point apically, and are apically bowed medially. The ventral valves, are slightly shorter than the dorsal valves, are dorso-laterally flattened and truncated at their apices (Fig.

34C-F). The male furculae of M. texensis are longer than those of M. scudderi and more triangular in shape (Fig. 34A-B).

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Holotype

College Sta. Tex. Dec. 20, [19]05. Deposited in the Illinois Natural History

Survey Collection.

Other specimens examined

Texas: Brazos Co., 4 mi NE Kurten on US Hwy 190, 1 September 1954, T.H.

Hubbell & I.J. Cantrall (3♂); 10 mi NW Navasota, 9 August 1955. Rehn (3♂, 3♀);

College Station, 24 Dec. 1905 (1♂); September-12 October 1926, V.A. Little (8♂, 12♀),

Lick Creek Park 30º34'00"N 96º12'41"W, 24 September 2012, J.G. Hill (12♂, 11♀).

Comanche Co., 4 August 1924, V.A Little (1♂). Dallas Co., Dallas, 14-16 August 1915,

Rehn (5♂). DeWitt Co., 1.3 mi SE Thomaston, 18 August 1965, T.J. Cohn (1♂).

Eastland Co., Cisco, 21-22 Sept 1912 (1♂). Houston Co., 4 mi N Grapeland, 27 August

1955, Rehn (3♂ 3♀). Madison Co., 5.5 mi NE Madisonville, 27 August, 1955, Rehn

(24♂, 9♀); 12.5 mi SW Madisonville on US Hwy 190, 2 September 1954, T.H. Hubbell

& I.J. Cantrall (2♂). Robertson Co., Hearne, 14-16 August 1915, Rehn (5♂, 7♀).

Washington Co., 2.5 W Washington, 9 August 1955, Rehn (1♂).

Distribution

Melanoplus texensis inhabits a narrow band of counties in east central Texas (Fig

30).

Habitat

At Lick Creek Park, M. texensis was very abundant in a post oak savannah in

dense growth of Schizachyrium scoparium. Cantrall's field notes state that the species

was very abundant in a "brushy pasture with scattered post oaks and hickories, with

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sparse growth of coarse grasses" and "yaupon shrubs and trumpet creeper along fence".

Cantrall states that during the night, M. texensis "was perched everywhere on yaupon shrubs along the fences, up to 5 feet [1.5m] from the ground – also on red elm shrubs, trumpet vines, grasses etc." The "next morning they were "common all through the

pasture".

Melanoplus dawsoni (Scudder) Incertae sedis

Melanoplus dawsoni was placed in the Scudderi group by Blatchley (1920).

However, upon review, M. dawsoni does not belong in the Scudderi group as it possesses

lanceolate tegmina, smaller cerci, large furculae, significantly different morphology of

the male genital structures unlike species in the Scudderi group. In addition, the

coloration of M. dawsoni is generally darker brown dorsally with a single vertical stripe

on most abdominal segments and a brighter yellow ventral surface than species in the

Scudderi group. At present, the group placement of M. dawsoni is uncertain.

121

Figure 34 Melanoplus texensis.

122

The Carnegiei group

Though similar in most external morphology, the internal genitalia of species in

the carnegiei group have a greatly reduced aedeagal sheath that is not approximate to the

valves. The Melanoplus carnegiei species group is established here to include M. carnegiei and M. acidocercus along with three previously undescribed species,

Melanoplus carolina, Melanoplus chattahoochee and Melanoplus savannah. Most

species in the carnegiei group are endemic to the Coastal Plain of southeastern United

States, with the exception M. carnegiei, which has also has been documented in the

southern Appalachians.

Diagnosis of species in the Melanoplus carnegiei Group

Species of medium to small size (♂ 16.0 – 21.5 mm, ♀ 20.1 – 27 mm). Head

slightly wider than pronotum; fastigum steeply declivent, Eyes somewhat prominent,

especially in males. Antennae usually 22-merous, but often 23 to 26-merous; nearly

cylindrical; equal in width throughout, except two basal segments. Head slightly wider

than pronotum. Pronotum broadly convex with distinct, but broadly rounded shoulders,

anterior margin sub-truncate, often somewhat emarginate; lateral margins of prozona

parallel, metazona margins diverging posteriorly; median carina distinct and equal in height throughout; anterior and median sulci present laterally, approaching the median carina, but not dissecting it; posterior sulci dissecting the median carina; prozona mostly smooth, with very little punctation on the lobes ventrally; metazona densely punctate; posterior margin broadly obtuse-angulate, prosternal spine elongate, blunt. Tegmina ovate; dorsal margins overlapping; length variable but, typically extending to the midpoint of the second abdominal tergite. Pro- and mesothoracic legs moderately stout; 123

metathoracic legs with the femora greatly swollen, extending just beyond the caudal end of abdomen. Metathoracic tibia with twelve to fifteen pairs of spines, but typically 12-

13. Furcula forming rounded protuberances, projecting slightly beyond the end of the segment from which they originate; bases minutely separated. Supra-anal plate (Fig.

35A) triangular, slightly longer than wide, with the median groove anteriorly distinct with

elevated sides, and diverging and becoming less distinct posteriorly. Cercus variable, but

generally triangulate to weakly falciform (Fig. 35B), being broader at base than apex,

Subgenital plate slightly conical.

Phallic Structures

Dorsal valves conical, tapering to a point. Ventral valves typically dorso-

ventrally flattened and variously shaped depending on the species. Aedeagal sheath

reduced, covering only the base of the valves. The epiphallus is of the typical

melanoploid shape, having lophi, ancorae, and an undivided bridge, but more precisely,

species in the Carnegiei group have a concave bridge, broadly rounded lophi, convexly curved lateral plates that are subdeltate in shape with a rounded anterior lobe and an acuminate caudal tip, and ancora that are triangular in shape, often tapering to a point.

Coloration

Males: Antenna ferruginous with a dark band on the ventral surface of the second

basal segment. Antennal crescent complete. A black spot present at the juncture of the

clypeus, gena, and frons. Head, thorax, and abdomen ferrugineou-fuscous, infuscated

dorsally, testaceous ventrally. A lateral, well-defined, piceous, post-ocular stripe extends

from the caudal margin of the eye to the second abdominal tergite, but often indistinct or

124

lacking on the metazona; lateral area of head, thorax, and abdomen below post-ocular stripe greyish-yellow. Tegmina ferrugineo-fuscous with slightly lighter venation. Hind femora ferrugineo-testaceous dorsally and laterally, bimaculate or often feebly trimaculate, with fuscous spots often extending halfway across medial area laterally; castaneous ventrally, knees darker. Hind tibia bright red, often dulled basally; with black or black tipped spines (Fig. 35 J).

Melanoplus carnegiei Morse (Figs. 35A-J, 38)

Melanoplus carnegiei Morse 1904: 10-11; Morse & Hebard 1915:103 (Lectotype

designation); Hill 2009: 32 (genitalia illustration).

Male Measurements (mm)

(n = 12) Body length 18.5 – 20.5 (mean = 19.6); pronotum length 4.0 – 5.0 (mean

= 4.6); tegmen length 3.5 – 4.5 (mean = 4.5); hind femur length 9.6 – 11.9 (mean = 10.7);

cerci length .6 - .8 (mean = 0.8); basal width of cerci 0.7 - .8 (mean = 0.8); mid-cercal

width 0.3 - 0.4 (mean = 0.4); cerci apex width 0.1- 0.2 (mean = 0.1).

125

Figure 35 Melanoplus carnegiei.

126

Female Measurements (mm)

(n = 12) Body length 20.5 – 27.0 (mean = 24.1); pronotum length 5.2 – 7.0 (mean

= 6.0); tegmen length 3.5 – 6.0 (mean = 5.1); hind femur length 12.0 – 14.5 (mean =

13.1).

Diagnosis

Distinguished from other species in the group by the shape of the male cerci (Fig.

35A-B), which are broadly triangular, acute apically, and with the length often equal to

basal width, and in the shape of the internal male genitalia. In lateral view, the valves

project posteriorly, are bowed dorsally and are dorsally recurved apically. In dorsal view,

the dorsal valves appear cylindrical in shape, taper to a point, and in the apical third are

bowed laterally, such that their apices are sub-attingent. In dorsal view the ventral valves

are dorso-laterally flattened, are slightly longer than the dorsal valves, and in the apical fourth abruptly taper and curve above the dorsal valves. The process of the ramus of the cingulum is well developed in this species, such that it has the appearance of a third pair of valves (Fig. 35F) below the ventral valves, and extends approximately one to two- thirds the length of the ventral valves.

Holotype

♂; Denmark, South Carolina, August 15, 1903, Morse. Deposited in the ANSP.

Other specimens examined

Alabama: Macon Co., 3 mi W jct. 60 and 29, 20 November 1954, M.E. Dakin

(1♂). Georgia: Bryan Co., 1 mi E Groveland, 32º08'14"N 81º45'16"W, 30 June 2006,

J.G. Hill (1♂); Groveland, 32º07'03"N 81º44'56"W, 13 October, 2010, J.G. Hill (2♂,

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3♀). Burke Co., Waynesboro, 29 August 1922, T.H. Hubbell (3♂); Thompson Bridge

Rd. 33º03'17"N 81º54'55"W, 18 September 2013, J.G. Hill (1♂, 1♀); Yuchi WMA,

33º05'21"N 81º47'04"W, 12 October, 2010, J.G. Hill, (5♂, 2♀), 33º05'43"N 81º39'40"W

(6♂, 2♀). Chatham Co., Near Burroughs, 22 September, 1945, T.H. Hubbell (2♂).

Clarke Co., Athens, 2 December 1922 (1♂). Emanuel Co., George L. Smith State Park,

32º32'44"N 82º07'21"W, 7 October, 2007, J.G. Hill (8♂, 3♀); Little Ohoopee Rvr X Hwy

50, 32º30'25"N 82º25'41"W, 6 October, 2007, J.G. Hill (1♀); Ohoopee Dunes N.A.,

32º32'15"N 82º27'40"W, 9 October, 2007, J.G. Hill (3♂, 1♀), 32º31'51"N 82º27'23"W, 6

October, 2007, J.G. Hill (3♂, 1♀), 25 Oct 2010 (3♂); 3.3 mi N Swainsboro, 32º40'37"N

82º23'08"W, 11 August, 2009, J.G. Hill (1♂); 3.8 mi N Swainsboro, 32º39'00"N

82º22'07"W, 11 August, 2009, J.G. Hill (2♂); Swainsboro, 29 September, 1945, T.H.

Hubbell, (4♂). Fulton Co., Hapeville, 7 September 1937 (11♂, 4♀). Jefferson Co., 4.8

mi N Wrens on Hwy 17, 9 September 1954, T.H. Hubbell & I.J. Cantrall (1♂, 4♀);

Wadley, 29 September, 1945, T.H. Hubbell (1♂); 1.65 mi N Wadley, 32º54'25"N

82º24'57"W, 11 August 2009, J.G. Hill (1♂). Laurens Co., 3.6 mi W Scott, 30 September

1945, T.H. Hubbell (1♂, 1♀). Richmond Co., Augusta, 5 September, 1930, T.H. Hubbell

(1 ♂); 1.3 mi W Hephzibah, 9 September 1954, T.H. Hubbell & I.J. Cantrall (1♂, 2♀);

Screven Co., 2.3 m W jct. US 301 & GA 24, 5 September 1954, T.H. Hubbell & I.J.

Cantrall (6♂, 4♀); 8.8 mi N Sylvania, 20 October, 1946, T.H. Hubbell (1♂). Tattnall

Co., 4 mi W Reidsville, 6 August 1947 (5♂). Toombs Co., 5.5 mi E Altamaha on U.S. 1,

29 September, 1945, T.H. Hubbell (1♂); 9.1 mi S Lyons. T.H. Hubbell (5♂, 1♀).

Treutlen Co., 5 mi N Gillis Springs, 33º04'56"N 81º47'04"W, 25 August 2010, J.G. Hill

(1♂). North Carolina: Buncombe Co., Swannanao, 11 August 1965, J.R. Bailey, (2♂).

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Jackson Co., 1.5 mi W., Dillsboro, 2 September 1951, I.J. Cantreall (3♂). Cherokee Co.,

2.2 mi W Murphy om U.S. 64, 1500 ft, 7 Sept, 1952, T.H. Hubbell & party (34♂, 13♀).

Clay Co., near Chatuge Lake dam, 1923 ft, 7, September, 1952, T.H. Hubbell & party

(10♂). South Carolina: Barnwell Co., 5.5 mi W Denmark on U.S. 78, 10 September,

1952 T.H. Hubbell (4♂, 4♀). Colleton Co., 2.2 mi N Salkehatchie on U.S. 21, 21 August,

1947, T.H. Hubbell (1♂, 1♀). Jasper Co., 1.5 mi S. Coosawhatchie on U.S. 17, 21 Aug,

1947, T.H. Hubbell (6♂); 2.5 mi N Tillman, 20 Oct 1946, T.H. Hubbell (2♂); 5.6 mi N

Ridgeland, 25 September-3 October 1930, T.H. Hubbell (4♂, 3♀). Orangeburg Co., 6.3

mi N Bamburg, 10 September 1952, T.H. Hubbell, (1♂).

Distribution

Melanoplus carnegiei is distributed from eastern Alabama across most of Georgia

and South Carolina, and extreme western North Carolina (Fig. 38).

Habitat

Across most of its distribution, M. carnegiei is often found in sandhills in

Georgia. Hill (2009) reports the species from aeolian sand dunes along the Little

Ohoopee River in Georgia. During the present study, M. carnegiei was found inhabiting

the edge of an open pine forest in dense growth of grasses and Rubus and Smilax.

Specimens from western North Carolina lack habitat information.

Melanoplus acidocercus Hebard (Figures 36A-J, 38)

Melanoplus carnegiei acidocercus, Blatchely 1920: 396.

Melanoplus acidocercus Hebard 1939: 290; Eades et al. 2014.

129

Male Measurements (mm)

(n = 12) Body length 18.8 – 21.5 (mean = 20.0); pronotum length 4.4 – 5.5 (mean

= 4.9); tegmen length 3.9 – 4.9 (mean = 4.3); hind femur length 10.9 – 12.2 (mean =

11.3); cerci length .8 – 1.1 (mean = 1.0); basal width of cerci 0.7 - .8 (mean = 0.8); mid- cercal width 0.3 - 0.4 (mean = 0.4); cerci apex width 0.1.

Female Measurements (mm)

(n = 10) Body length 20.1 – 26.9 (mean = 23.5); pronotum length 4.8 – 7.1 (mean

= 5.2); tegmen length 4.0 – 6.0 (mean = 5.2); hind femur length 12.2 – 15.5 (mean =

13.9).

Diagnosis

Distinguished from other species in the group by the shape of the male cerci (Fig

35A-B), which are triangular with the length longer than the basal width and acute apically, and in the shape of the internal male genitalia (Fig. 35C-I). In lateral view the valves project posteriorly, are slightly bowed dorsally and are apically recurved dorsally.

In dorsal view, the dorsal valves appear cylindrical in shape, taper to a point, and in the apical third are weakly bowed laterally, such that their apices are sub-attingent. In dorsal view the ventral valves are dorso-laterally flattened, are slightly longer than the dorsal valves, and in the apical fourth abruptly taper and curve above the dorsal valves. The enlarged process of the ramus of the cingulum present in M. carnegei is absent or greatly reduced in M. acidocercus.

Holotype

Bainbridge, Ga. IX 5-6 1915, (R. &H.). Deposited in the ANSP.

130

Other specimens examined

Florida: Columbia Co., Lake City, 21 August 1938 (2♂). Madison Co., Ellaville

(5♂). Georgia: Appling Co., 1.9 mi S Altamaha, 19 October 1946, T.H. Hubbell (4♂,

1♀); Bibb Co., Cross Keys, August 1964 (2♂, 1♀). Macon, 25 August 1938 (19♂, 4♀);

Mogul, 32º50’59”N 83º33’45”W, 10 August 2009, J.G. Hill (1♂). Coffee Co., 11.2 mi N

Broxton, 30 September, 1945 T.H. Hubbell (1♂, 1♀); Crawford Co., 1mi. S Gallard, 14

August 1947, T.H. Hubbell (2♂); 2.6 mi SE Knoxville, 32º41'58"N 83º57'45"W, 10

August 2009, J.G. Hill (1♂). Decatur Co., Bainbridge, 11 September, 1954, T.H.

Hubbell & I.J. Cantrall (4♂, 1♀); Houston Co., 5 mi N Perry, 10 September, 1924, T.H.

Hubbell (6♂, 4♀); Perry, 10 September, 1924, T.H. Hubbell (3♂); Laurens Co., Garetta,

30 September, 1945 T.H. Hubbell (2♂, 2♀).

Distribution

Melanoplus acidiocercus is distributed on the Coastal Plain of southern Georgia and in northern Florida (Fig. 38).

Habitat

Melanoplus acidocercus has been collected in open pine forest and sandhills.

131

Figure 36 Melanoplus acidocercus.

132

Melanoplus chattahoochee new species (Fig. 37A-J, 38)

Male Measurements (mm)

(n = 13) Body length 17.2 – 21.0 (mean = 18.8); pronotum length 4.2 – 5.4 (mean

= 4.6); tegmen length 3.2 – 4.7 (mean = 4.2); hind femur length 9.5 – 11.5 (mean = 10.9); cerci length 0.6 - 0.9 (mean = 0.8); basal width of cerci 0.7 - 0.9 (mean = 0.8); mid-cercal width 0.4 - 0.5 (mean = 0.4); cerci apex width 0.1- 0.2 (mean = 0.1).

Female Measurements (mm)

(n = 7) Body length 22.0 – 26.0 (mean = 23); pronotum length 5.7 – 6.5 (mean =

6.1); tegmen length 4.8 – 6.5 (mean = 5.4); hind femur length 13.5 – 14.5 (mean = 13.8).

Diagnosis

Distinguished from other species in the group by the shape of the male cerci (Fig

37A-B), which are broadly triangular, blunt apically, and with the length often subequal to the basal width and in the shape of the internal male genitalia (Fig. 37C-I). In lateral view the valves project posteriorly, are slightly apically curved ventrally. In dorsal view, the dorsal valves appear broadly cylindrical in shape being slightly dorso-ventrally flattened, taper to a point, and in the apical third are weakly bowed laterally, such that their apices are sub-attingent. In dorsal view the ventral valves are dorso-laterally flattened, are slightly longer than the dorsal valves, and in the apical fourth abruptly taper and curve slightly above the dorsal valves. In M. chattahoochee, the tips of the cingulum are bent medially, whereas in other species of the group they are straight. The enlarged process of the ramus of the cingulum present in M. carnegei is greatly reduced and

133

typically obscured from view by the aedeagal sheath in most specimens of M.

chattahoochee.

Holotype

♂; GA [GEORGIA], Taylor Co., Black Creek Nat[URAL]. Area, 32°34’23”N

84°24’11”W, 24 August 2010, J.G. Hill, Collected in Fall Line sandhill. Deposited in the

USNM.

Paratypes

Same data as holotype (1 ♂).

Other specimens examined

Georgia: Marion Co., 2 mi SE Juniper, 32°30’32”N 84°39’15”W, 25 August

2010, J.G. Hill (1 ♂); .25 mi E jct GA 103 & 41 at W limits Buena Vista, 10 September

1954, T.H. Hubbell and I.J. Cantrall (2 ♂). Talbot Co., Black Jack Crossing Pres.,

32°34’38”N 84°29’55”W, 11 October, 2010, J.G. Hill (2 ♂, 3 ♀); 32°35’06”N

84°30’23”W, 25 August 2010, J.G. Hill (2 ♂, 3 ♀); .5 mi N Junction City, October, 1964,

Sandford and Hayes (2 ♂). Taylor Co., Black Creek N.A., 32°34’11”N 84°24’27”W, 11

October 2010, J.G. Hill (3 ♂, 1 ♀); 32°33’53”N 84°24’25”W, 11 October, 2010, J.G. Hill

(4 ♂, 2 ♀).

134

Figure 37 Melanoplus chattahoochee.

135

Etymology

Named after the Chattahoochee River and in particular the so called

"Chattahoochee Fall Line", an area of sandhills to the east of the Chattahoochee River

along the Fall Line where this species is apparently endemic.

Distribution

At present, M. chattahoochee is known only from the Fall Line sandhills of

western Georgia (Fig. 38).

Figure 38 Distribution of Melanoplus acidocercus, M. carnegiei, M. carolina, M. chattahoochee, and M. savannah.

136

Melanoplus savannah new species (Figs. 38, 39A-J)

Male Measurements (mm)

(n = 14) Body length 16.0 – 19.7 (mean = 18.6); pronotum length 3.7 – 5.2 (mean

= 4.5); tegmen length 3.3 – 4.4 (mean = 3.8); hind femur length 9.8 – 12.2 (mean = 11.1); cerci length 1.1 - 1.4 (mean = 1.3); basal width of cerci 0.5 - 0.7 (mean = 0.6); mid-cercal width 0.5 - 0.7 (mean = 0.6); cerci apex width 0.3- 0.4 (mean = 0.3).

Female Measurements (mm)

(n = 5) Body length 20.5 – 22.5 (mean = 21.7); pronotum length 5.0 – 6.0 (mean

= 5.6); tegmen length 3.6 – 5.0 (mean = 4.6); hind femur length 12.3 – 14.0 (mean =

13.2).

Diagnosis

Distinguished from other species in the group by the shape of the male cerci (Fig

39A-B), which are sub-triangular, being slightly depressed dorsally and slightly rounded ventrally and broadly rounded apically. (Fig 39A-B), and in the shape of the internal male genitalia (Fig. 39C-I). In lateral view, the valves project dorsally, are subapically curved posteriorly and apically recurved laterally and dorsally. In dorsal view, the dorsal valves appear cylindrical, taper to a point, and in the subapical third are curved laterally, and then recurved dorsally. The ventral valves are dorso-laterally flattened, taper to a point, and extend apically to approximately the laterally curved portion of the dorsal valves.

The enlarged process of the ramus of the cingulum in M. carnegei is apparently reduced or absent in M. savannah.

137

Holotype

♂; S.C. [SOUTH CAROLINA], Barnwell Co. Ditch Pond Her[itage]. Pres[erve].,

33°24’57”N 81°28’13”W, 12 October 2010, J.G. Hill, sandhill near Carolina Bay.

Deposited in the USNM.

Paratypes

South Carolina: Same data as holotype (3♂, 2♀).

Other specimens examined

South Carolina: Aiken Co., 2.7 mi W Montmorenci, 33°31’59”N 81°35’10”W,

12 October 2010, J.G. Hill (6 ♂, 3 ♀); 5.2 mi ESE Beech Island, 33º24'42"N

81º48'12"W, 15 September 2013, J.G. Hill (1 ♀); ≈10 mi NW Aiken, 33°35’19”N

81°52’55”W, 23 July 2008, J.G. Hill (4♀); Savannah River Site, 33°21’51”N

81°35’12”W, 5 October 2012, J.G. Hill (1♀). Barnwell Co. Ditch Pond Heritage

Preserve, 33°24’57”N 81°28’13”W, 15 September 2013 (2 ♀).

Etymology

In reference to the Savannah River, which seems to be the western boundary of

the species.

Distribution

At present, M. savannah, is known only from Aiken and Barnwell Counties in

South Carolina (Fig. 38).

138

Figure 39 Melanoplus savannah.

139

Habitat

Melanoplus savannah has been collected in Fall Line sandhills on the east side of

the Savannah River in South Carolina. All the locales where this species has been collected are open areas with sandy soil, vegetated with wiregrass (Aristida stricta), and

prickly pear (Opuntia sp.). Some sites have an overstory of longleaf pine (Pinus palustris). At the Savannah River Site, this species was quite numerous among the smoldering understory of a longleaf pine forest that had been recently burned.

Melanoplus carolinensis new species (Fig. 38, 40A-J)

Measurements (mm)

(n = 1) Body length 19.1; pronotum length 4.1; tegmen length 4.0; hind femur

length 10; cerci length 0.8; basal width of cerci 0.7; mid-cercal width 0.8; cerci apex

width 0.3.

Diagnosis

Distinguished from other species in the group by the shape of the male cerci (Fig

40A-B), which are sub-falcate and truncated apically (Fig 40A-B), and in the shape of the

internal male genitalia (Fig. 40C-I). In lateral view the valves project dorsally and

posteriorly at a 45º angle and are subapically curved ventrally. In dorsal view, the dorsal

valves appear cylindrical, taper to a point, and are subapically curved ventrally. The

ventral valves are dorso-laterally flattened, and are broadly rounded apically, and are

subequal in length to the dorsal valves. The enlarged process of the ramus of the

cingulum in M. carnegei is apparently reduced or absent in M. carolinensis.

140

Holotype

♂; SC.[SOUTH CAROLINA], Chesterfield Co., Carolina Sandhills NWR,

34º31’19”N 80º13’27”W, 17 September 2013, J. G. Hill..

Etymology

Named in reference to the Carolina sandhills where the species was discovered.

Distribution

At present, M. carolinesis is known only from the type locality (Fig. 38).

Habitat

At the type locality, M. carolinesis, was collected in a clump of low-growing

Vaccinium in an open sandhill with abundant bare sand and several small Quercus laevis and an overstory of Pinus palustris.

141

Figure 40 Melanoplus carolinensis.

142

Molecular Analysis

Utilizing the barcode data, Neighbor-joining, Maximum Likelihood, and

Maximum Parsimony trees were constructed using Mega6 software. All the trees produced similar overall topology and support for major clades. The relationships based on COI sequences appear well supported for some species such as M. savannah (.77

bootstrap support) whereas for other species such as M. acidocercus, M. carnegiei, and

M. querciola, the relationships are not consistent with the morphological species concepts

presented above, thus they appear as polyphyletic in Figure 41. These species level

results are consistent with other DNA barcode studies in Orthoptera as well as

Calliphorid flies whereby barcoding does not often yield consistent identifications (Song

et al. 2007, Walker and Funk 2014, Whitworth et al. 2007). The three phylogenetic

techniques consistently divided the samples into three main clades (values presented are

for Maximum Likelihood analysis): a western clade of specimens from west of the

Mississippi River (76% bootstrap support), a central clade consisting of Scudderi group specimens from west of the Chattahoochee and Apalachicola Rivers and east of the

Mississippi River (97% bootstrap support), and an divergent eastern clade comprised of

Scudderi and Carnegiei group species from east of the Apalachicola - Chattahoochee

Rivers (46% bootstrap support, but sub-clades with greater support). (Fig. 41 and 42)

The cases where the barcoding apparently accurately delineated species are likely just an

artifact of this geographic structure as the species delineated all have relatively small

geographic ranges, whereby species with broader distributions were often polyphyletic,

which is consistent with other studies of Melanoplus phylogeography (See Knowles

2000)

143

Pairwise genetic distances were compared between members of all species and M.

scudderi in each clade (intraclade) and similarly between individuals from different

clades (interclade) in MEGA6. For the COI gene, all interclade distances are greater than

all intraclade distances (Table 1.), which suggests in concert with the phylogenies, that

these mtDNA clades represent independent evolutionary lineages, and that members of

each clade are more closely related to each other than they are members of other clades

(See also Avise 1994 and Burbrink et al. 2000).

Table 2 Estimates of Inter and Intraclade Evolutionary Divergence over sequence pairs.

Clade Eastern Central Western Eastern All 8.74 (1.4) M. scudderi 15.0 (3.46) Central All 14.78 (1.80) 2.43 (.57) M. scudderi 19.0 (4.05) * Western All 15.73 (3.15) 5.89 (2.9) 1.0 (.69) M. scudderi 20 (2.60) 7.2 (4.3) 0 (0) The number of amino acid differences per sequence from averaging over all sequence pairs for all species and Melanoplus scudderi only between clades are shown. Standard error estimates are shown in parentheses and were obtained by a bootstrap procedure (100) replicates. * Melanoplus scudderi is the only species present in this clade.

144

Figure 41 Molecular phylogentetic analysis of species in the Melanoplus scudderi and carnegiei groups by maximum likelihood using Jukes-Cantor model.

The tree with the highest log likelihood is shown (-2146.6867). Bootstrap values above 45 are shown to allow for better observation of the split between the clades. Numbers for the eastern sub-clades correspond to those mentioned in the text.

145

Figure 42 Distribution of samples from each clade showing the Apalachicola- Chattahoochee River /Mississippi River discontinuity.

Discussion

Given the amount of diversity discovered in recent revisions of Melanoplinae

(Otte 2012, 2014), it was not unexpected that the Scudderi group would be comprised of

additional cryptic species; however, the amount of diversity discovered was surprising.

The Scudderi group as defined above is made up of 23 sibling species that are allopatric

or parapatric with clearly defined distributions that often correlate with physiographic or

biotic regions suggesting that the species are geographically and possibly ecologically

isolated. 146

Biology

Like many short winged grasshoppers, members of the Scudderi and Carnegiei

groups prefer brushy conditions. More precisely, members of the group are often found

in areas with young trees, shrubs, and tangles of Rubus and Smilax or in dense growths of

various grasses and members of the Asteraceae such as Ambrosia, Helianthus and

Solidago. In most cases of field collections, one species appears to be replaced by another

species within a few miles under seemingly identical ecological conditions.

Before large-scale human disturbances, these grasshoppers were likely denizens of the ruderal areas of relatively open habitats, including prairie, savannahs, barrens,

glades, or forest openings made by wind or fire. Human disturbances, namely the

clearing of forests, have increased the amount of suitable habitat for these species such

that they now inhabit fence rows, power/gas line right of ways, uncultivated fields,

unmown margins of roadsides, and other ruderal environments. One might surmise that

the pervasive amount of anthropogenic disturbance present across eastern North America

that has greatly increased the amount of habitat for these species would have made a

hodge-podge of the distribution of these species by creating artificial dispersal routes.

Unhindered by traditional barriers such as forests and swamps, these species would be

free to move out in all directions, coming into contact with other species of the group and

potentially interbreed with them. Based on this study, the mixing of morphologically

defined species has not happened, at least not extensively. Several demes have been

discovered that are apparently hybrids or intermediates where two species come into

contact with each other (a phenomenon documented in several other Melanoploid groups

(See Cantrall and Cohn 1972 and Dakin 1985). However, this is the exception and not

147

the rule. At most it appears that anthropogenic disturbances have possibly increased number of demes, particularly with the widespread M. scudderi. The habitats preferred

by most of the members of the Scudderi group are ruderal in nature and can also be

brought about by numerous natural disturbances (fire, wind, drought) and edaphic

conditions making it likely that their distributions have been in balance for some time.

The present distribution of some species of the Scudderi group in the southeastern

United States (such as M. muscogee and M. folkertsi) can be adequately explained on the basis of natural barriers such as major rivers (such as the Chattahoochee, Flint, and

Tombigbee among others). The species of the Scudderi group are closely related and

have relatively discrete distributions across the region. These facts point to a fairly recent

origin, probably no earlier than the late Pliocene. However, why widespread species,

such as Melanoplus scudderi, have not spread across the entire range is not clear. Perhaps

some other mechanism, such as competitive exclusion, which is typical of sibling species

is retarding dispersal.

The basic ecology and life history of species in the Scudderi and Carnegiei

groups need further study. For instance, though most species of Melanoplus are thought

to be generalist herbivores, the diet of species in the Scudderi and Carnegiei groups is

unknown. Additionally, more in depth natural history and ecological studies, such as

ecological niche modeling, may help elucidate the factors that result in the allopatric

distribution of species. These studies are particularly important for species with small

distributions that are habitat limited, as the resulting information may aid in any

necessary conservation or management actions.

148

Molecular Discussion

Based on the molecular data available at present, the Melanoplus scudderi group

is composed of three mtDNA clades: 1) an eastern clade, also containing the Carnegiei

group, distributed east of the Chattahoochee and Apalachicola River; 2) a central clade,

distributed between the Chattahoochee and Apalachicola Rivers to the east and the

Mississippi River to the west; and 3) a western clade, distributed west of the Mississippi

River (Figs 41-42). This pattern has been documented for several organisms including the

American rat snake, Elaphe obsoleta (Say); eastern fence lizard, Sceloporus undulatus

(Bosc and Daudin); and the northern short-tailed shrew, Blarina brevicauda (Say) (Brant

and Orti 2003, Buribrink et al. 2000, Leache and Reeder 2002, also see review in Soltis et

al 2007). In these examples, each clade is thought to have arisen as a result of isolation

during Pleistocene glaciation and subsequent isolation by major river systems. The

effects of Pleistocene glaciation have not been well documented for grasshoppers in the

southeastern United States, but studies from the western U.S. have documented that

glaciation did indeed play an important role in promoting genetic drift and speciation in

Melanoplus by displacing populations to multiple glacial refugia (Knowles 2000, 2001,

and Knowles and Otte 2000, Knowles and Richards 2005).

The eastern and central clades are separated by the Chattahoochee and

Apalachicola Rivers. This watershed is a known site of morphological and molecular

discontinuity as well as being a center of endemism. (See reviews in Avise 1996, Hubbell

et. Al, 1956, Soltis et. Al 2006). At the maximum rise in sea level during the Pliocene

and Pleistocene, the Apalachicola River was embayed as a large saltwater channel well

into Georgia and Alabama, possibly up to the Fall Line in central Georgia (Cooke, 1945,

149

Neill 1957). For some taxa, it is at this point where the Appalachian Mountains, just to the north, further serve as a barrier for eastern and central clades (See Soltis et al. 2006 for review). This may be the case for the Scudderi group as well, which would be

supported by the morphological variant of M. scudderi that occurs east of the

Appalachians (Fig. 2). The central clade is separated from the western clade by the

Mississippi River, which again, has been documented for several groups of organisms

including fish, reptiles, and amphibians, and has been attributed to Pleistocene glaciation

(Burbrink et. al 2000).

Many of the species of the Scudderi and Carnegiei groups evidently developed in

classical areas of endemism, which is typical of the trend of many insects with low

vagility such as flightless grasshoppers (Hubbell 1932, Noonan 1988). Thirteen species in

the Scudderi group and four species of the Carnegiei group are southeastern Coastal

Plain endemics, and seven species of the Scudderi group are endemic to the Interior

Highlands.

The Coastal Plain extends from Massachusetts south to the Florida Keys, west to eastern Texas, and southward to Tamaulipas, Mexico with a northward projection, termed the Mississippi Embayment, extending to Southern Illinois (Fenneman 1938, Sorrie and

Weakley 2001). The Coastal Plain is the exposed portion of the continental shelf, and is composed of sedimentary deposits of Cretaceous Age or younger (Fennenman 1938

Sorrie and Weakley 2001). In the southeastern United States, the Coastal Plain covers large portions of Alabama, Arkansas, Florida, Georgia, Louisiana, Mississippi,

Oklahoma, North Carolina, South Carolina, and Texas, and spans the southern area of three mtDNA clades.

150

Eastern Clade

The largest representation of species available for the molecular analysis comes

from the eastern clade. This is due, in part, to the lack of species diversity with the groups

in the area covered by the central clade and poor sampling from the area covered by the

western clade. The eastern clade in comprised of samples from specimens collected on

the Coastal Plain in Georgia and Florida, with the exception of one specimen of M.

folkertsi from south Alabama. As seen from the phylogeny, this clade has many

divergent lineages between and within species. This structure suggests a complex pattern

of multiple refugia within the region: sub-clade 1 is comprised of specimens of two

species (with the exception of one specimen of M. folkertsi from Alabama) from southwest Georgia and nearby northern Florida; sub-clade 2 is comprised of specimens

M. savannah from the Fall Line Sandhills of western South Carolina, sub-clade 3 is

comprised of specimens from three species from southeastern GA, sub-clade 4 is

comprised of a single specimen of M. carnegiei from the eastern Fall Line Sandhills of

Georgia, and sub-clade 5 is comprised of three species from western Fall Line sandhills

of Georgia.

Sorrie and Weakely (2001) studied the phytogeographic patterns of Coastal Plain

endemic plants and documented 27 patterns of species distributions, many of which

demarcated classical areas of endemism. A comparison of the distribution of the species

in the Scudderi and Carnegiei groups to Sorrie and Weakely’s patterns revealed that nine

of the 13 Coastal Plain endemics could be assigned to one of their phytogeographic

patterns. Five species are endemic to sandhills along the Fall Line in Georgia, South

Carolina and North Carolina (Sorrie and Weakly pattern #5). Of these five species, two

151

are largely sympatric (though from different species groups) and the rest are allopatric with their ranges being segregated by large river systems of the region. This may be a possible explanation for structure seen in the sub-clades of the larger eastern clade as three of the clades (2, 4, and 5) are comprised of specimens from the Fall Line and may be isolated by larger rivers such as the Savannah and Oconnee or Ocmulgee Rivers.

Additional species distributions corresponding to Sorrie and Weakly’s

phytogeographic patterns include: M. acidocercus and M. quercicola roughly

corresponding to pattern #6 (southern South Carolina to peninsular Florida), M. relictus

to pattern #11 (central panhandle of Florida), and M. davisi and M. folkertsi to pattern

#12 (western panhandle of Florida and adjacent Alabama).

Central Clade

Only three species, M. scudderi, M. folkertsi, and M. mississippi occupy the area

between the Apalachicola/Chattahochee and Mississippi Rivers. None of these species

are endemic to the area covered by the central clade. Only two of the species, M.

scudderi and M. folkertsi were available for molecular analysis. The specimen of M.

folkertsi grouped with other folkertsi in the eastern clade, and was the only specimen to

cross barriers in the study (Figs. 41-42). The lowest amount of genetic divergence (2.43

total changes) is found in the area occupied by the central clade. This may indicate that

M. scudderi has colonized this area relatively recently and has not had time to diverge.

This may explain why grassland habitats typically rich in endemic species in this area

such as the cedar glades of the Central Basin of Tennessee and the Bibb County glades in

Alabama do not contain endemic Scudderi group species.

152

Western Clade

Specimens from the western clade were not well represented in the molecular

study, however, given the diversity and amount of major rivers and mountains, a

divergent pattern similar to that of the eastern clade is expected. West of the Mississippi

River on the Coastal Plain, M. irwinorum, M. taurus, and M. texarkana conform to Sorrie and Weakley’s phytogeographic pattern # 14 West Gulf Coastal Plain. Four other species, M. decurvus, M. mississippi, M. optimus, M. texensus, do not fit one of their

patterns of endemism, but all except, M. optimus, correspond to physiographic or

geographic features.

Endemism in the Interior Highlands

Seven species of the Scudderi group are endemic to the Interior Highlands. The

Interior Highlands are the only appreciable montane region between the Appalachian and

Rocky Mountains, and cover considerable parts of Arkansas, Missouri, Oklahoma, and

smaller portions of Kansas and Illinois. The highlands are comprised of three distinct regions: the Ozark Plateau, Arkansas River Valley, and the Ouachita Mountains. The

Ozark Plateau is a deeply dissected broad dome that is centered over the Saint Francois mountains of southeastern Missouri that has been slowly uplifted and eroded, which has resulted in high levels of topologic (elevations up to 793 m), edaphic, and hydrologic

diversity (Fenneman 1938, Zollner et al 2005). The Arkansas River valley, or the Arkoma

Basin, is a broad alluvial plain with several relatively high isolated areas, such as Mount

Magazine the highest point in Arkansas (839 m) that lies between the Ozark Plateau and

the Ouachita Mountains (Croneis 1930, Zollner et al. 2005). The Ouachita Mountains

(elevations up to 817 m) are a series of east-west fold-belt mountain ranges that in the 153

north consist of long ridges of sandstone, separated by broad valleys, and in the south are

sharp novaculite ridges separated by narrow, stony valleys (Croneis 1930, Zollner et al.

2005).

Much like on the Coastal Plain, the seven endemic species of the Scudderi group

are associated with a distinct portion of the region. Two species, M. ozarkensis and M. francoisi, are restricted to the Ozark Plateau, with M. ozarkensis occurring across much

of the Plateau and M. francoisi being restricted to the Saint Francois Mountains

subdivision of the Plateau. One species, M. arkansas, is endemic to the Arkansas River

Valley, south of the Arkansas River. Four species, M. baronei, M. cohni, M. ouachita,

and M. seltzerae, are endemic to the Ouachita Mountains.

The interior highlands are a well-known area of endemism with 36 plant species and 68 insect species considered endemic to the region (Allen 1990, Robinson and Allen

1995, Zollner 2005). Of the plants, the highest level of endemism in the Interior

Highlands is found in the Ouachita Mountains with 14 species (39%) being found only in that region (Zollner 2005). This is mirrored in the Scudderi group as four out of the seven

(57%) Interior Highland endemic species are endemic to the Ouachita range.

Events that may have led to speciation of the Scudderi and Carnegiei groups

Difficulty in determining the polarity of transformations of the male genitalic

structures and a lack of external variation prohibits the construction of a phylogeny with

morphological characters (Knowles and Otte 2000). The molecular data presented here, is

of little use due to the problems involved the short sequences used in DNA barcoding as mentioned above. In order to construct a sound phylogenetic framework, a more rigorous

molecular study involving more complete taxon sampling, larger sequences, and more 154

genes must be used, and indeed a study of this type is underway. However, until that study is complete, the hypothetical scenario below is presented for evolution of the group in the southeastern United States.

As previously mentioned, species in the of the Scudderi and Carnegiei groups are

inhabitants of open or grassland habitats, and as such are likely derived from an ancestor

that inhabited the less forested, western portion of their range. This ancestral species

could have subsequently spread east during periods of drier climatic conditions that

favored the spread of savannah and scrub habitat. On the Coastal Plain, this most likely occurred along the Gulf Coastal Corridor, whereby the Madro-Tertiary Geoflora

(semiarid live oak-conifer woodlands, arid subtropic scrub, grassland, subdesert to desert vegetation) and related fauna developed in Mexico and southwestern North America during the Miocene spread into the Great Plains and eastward along the Gulf Coast during the late Miocene and persisted into the early Pleistocene (Albright 1998, Axelrod

1958, Noss 2013, Webb 1977). During the last Glacial Maximum (26,500-19,000 years ago), sea levels dropped as a greater proportion of water was incorporated into glacial ice,

exposing more of the continental shelves thus making the Coastal Plain broader than it is

today. During this period, a warm thermal enclave that extended from Florida north to

Cape Hatteras, NC (35ºN) produced a climate that was likely warmer and drier,

supporting prairie and savannah adapted biota as evident from fossil and pollen data.

(Russell et. al 2009). Subsequent climate changes and rising sea levels after the retreat of

continental ice resulted in disappearance of the enclave. As a result, the climate of the

region became wetter, resulting in grassland habitats being supplanted by mixed forests.

These mixed forests resulted in reduced sediment load in rivers, which along with

155

increased moisture, led to the shallow shallow braided rivers of the region forming meandering channels and floodplains (Leigh 2006), which would have further isolated areas of grassland. The distribution of M. mississippi, which is endemic to the Mississippi

River Alluvial valley on both sides of the River may also be explained by events during the late Pleistocene as during this period the River was series of braided stream terraces

(Rittenour 2003, Saucier 1974) that would have allowed the flightless grasshoppers to migrate across the river as stream channels changed or dried.

In the Interior Highlands, edaphically controlled glades, prairies, and relatively open woodlands are common today. Again, during the last Glacial Maximum, drier climatic conditions would have likely allowed these open habitats to spread and into the area and then be isolated as the climate became increasingly wet resulting in widening rivers expanding forests. Indeed, more than half (58%) of the plant endemism in the

Interior Highlands is associated with glades, and an additional 22% is associated with dry

pine-oak and oak woodlands (Zollner 2005), both of which are suitable habitat for

members of the Scudderi group.

Many studies suggest that Pleistocene glaciation as a causal factor that led to

speciation in various groups in North America; however, as Allen (1990) indicates, some

areas, such as the Interior Highlands have been a positive feature on the North American

continent since the Pennsylvanian (320 mya). Nonetheless, for the Scudderi and

Carnegiei groups the timing of Pleistocene glaciation is likely appropriate as previous

studies have shown that many species of Melanoplus originated during the Pleistocene

and were strongly affected by the movement of glaciers and other climatic changes

(Knowles 2000, 2001, and Knowles and Otte 2000). Thus, it is feasible that the ancestral

156

form of the Scudderi and Carnegiei groups could have spread out of Texas eastward

along the Gulf Coastal Corridor and into the Interior Highlands and diverged as they

became subsequently isolated as a result of concomitant habitat changes due to climate

change. Given the amount of morphological differentiation present, it is likely the

ancestral species of the Carnegiei group as well as that of M. davisi and M. quericicola of

the Scudderi group were in the region for more time and may have been isolated during a

period of glaciation. During a later interstadial the ancestor of other members of the

group such as (M. relictus, M. coreyi, and M. folkertsi) could have colonized the area,

later to be isolated during interglacial periods.

The broad distribution of Melanoplus scudderi

As far is known Melanoplus scudderi is a non-migratory animal with very low

vagility. However, even with the reduction of range resulting from this revision, M.

scudderi still has the largest distribution of any brachypterous grasshopper in the eastern

United States. The wings are so short in most specimens that, as far as is known, they are functionless. Macropterous forms of the species, along with M. mississippi, have been

documented, but at present it does not appear that long-winged individuals are capable of

flight, making them no more vagile than the brachypterous individuals.

One would expect dispersal in the group to be slow resulting from peripheral expansion of the range; however, the opposite may be true. Given its broad distribution and relative abundance, despite not being capable of flight, M. scudderi is apparently a

competent disperser. The presence of M. scudderi in all three mtDNA clades suggests

that it emerged from at least several refugia, which may have aided its spread, to colonize

its present wide-ranging distribution. Melanoplus scudderi was often the first 157

grasshopper to colonize small (10 x 20 m) forest openings as part of Black Belt prairie restoration project (JGH Pers. Obs.). These plots are otherwise surrounded by dense

stands of eastern red cedar, and M. scudderi was not detected in the plots before they

were cleared. At present it is not known how new populations of Melanoplus scudderi or any brachypterous species colonize islands of suitable discontinuous habitat.

Conservation

Given the apparent ruderal nature of the habitat favored by these species, one might expect them to be omnipresent in modern landscapes. In fact, the opposite is sometimes the case. Anthropogenic disturbances while often creating favorable conditions for some Scudderi group species has also likely added to the isolation of many

of the populations. Over the western portion of the group’s range, namely in Texas north

through the Dakotas, large amounts of area was once covered with native grasslands that

consisted of a diverse mix on grasses and forbs in concert with the actions of drought, fire

and free-ranging mega-herbivore grazing and wallowing provided habitat for these

western species. Today, this area is largely covered in row crops or pastures planted in

exotic grasses such as Bermuda grass [Cynodon dactylon (L.) Pers.], Johnson grass

[Sorghum halepense(L.) Pers.], and Old world bluestems (Bothriochloa spp.). The

grazing action of cattle (Bos primigenius Bojanus) is said to be analogous to that of the

once dominant American bison (Bison bison L.), however fenced in cattle often graze the vegetation close to the ground such that there is no cover. Additionally, the wallowing behavior possessed by bison, but not cattle, has been found to greatly influence the local plant community, especially by promoting the abundance of Ambrosia (McMillian et al.

2011, Gill 2013), a plant which species in the Scudderi group are commonly associated 158

with as well as other grasshoppers such as Campylacantha olivacea Scudder. As a result

of these anthropogenic actions, it was often difficult to find suitable habitat to sample

species such as M. latus and M. optimus during this revision..

Many of the species in the eastern portion of the range are associated with

longleaf pine (Pinus palustris) savannahs. Today, it is estimated that 3% of the longleaf

ecosystem remains (Frost 1993, Platt 1999). Much of this land has been cleared for

agriculture or is in timber production of loblolly (Pinus taeda) and slash pine (Pinus

elliottii) and is under management practices that do not favor the growth of understory

vegetation. Many species like M. davisi, M. quercicola, M. relictus are frequently found in low-growing oaks that have been stunted by fire. Fire suppression, on feral lands, allows these oaks to grow tall and shade out the understory and eliminating the habitat for the savannah biota. Thus, throughout the range of the group numerous anthropogenic actions have resulted in fragmenting populations of these species.

List of Groups and Species

A key to the species groups and the species is available in Appendix A.

Scudderi group

Melanoplus scudderi (Uhler) — Figures 1A-J, 2A-B, 3, 4A-B

Melanoplus folkertsi NEW SPECIES — Figures 5A-J, 6

Melanoplus relictus NEW SPECIES — Figures 6, 7A-J

Melanoplus coreyi NEW SPECIES — Figures 6, 8A-J

Melanoplus muscogee NEW SPECIES — Figures 6, 9A-J

Melanoplus quercicola Hebard — Figures 6, 10A-J

159

Melanoplus davisi (Hebard) — Figures 6, 11A-J

Melanoplus mississippi NEW SPECIES — Figures 12A-J, 13

Melanoplus arkansas NEW SPECIES — Figures 14A-J, 17

Melanoplus ouachita NEW SPECIES — Figures 15A-J, 17

Melanoplus cohni NEW SPECIES — Figures 16A-J, 17

Melanoplus seltzerae NEW SPECIES — Figures 17, 18A-J

Melanoplus baronei NEW SPECIES — Figures 17, 19A-J

Melanoplus latus Morse, NEW STATUS — Figures 20A-J, 21A-D, 22

Melanoplus decurvus NEW SPECIES —Figures 17, 23A-J

Melanoplus ozarkensis NEW SPECIES — Figures 24A-J, 25

Melanoplus francoisensis NEW SPECIES — Figures 25, 26A-J

Melanoplus texarkana NEW SPECIES — Figures 27A-J, 28

Melanoplus taurus NEW SPECIES — Figures 29A-J, 30

Melanoplus optimus NEW SPECIES — Figures 30, 31A-J

Melanoplus ottei NEW SPECIES — Figures 30, 32A-J

Melanoplus irwinorum NEW SPECIES — Figures 30, 33A-J

Melanoplus texensis Hart, NEW STATUS — Figures 30, 34A-J

Carnegiei group

Melanoplus carnegiei Morse — Figures 35A-J, 38

Melanoplus acidocercus Hebard — Figures 36A-J, 38

Melanoplus chattahoochee NEW SPECIES — Figures 37A-J, 38

Melanoplus savannah NEW SPECIES — Figures 38, 39A-J

Melanoplus carolinensis NEW SPECIES — Figures 38, 40A-J 160

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Frison, T.H. 1927. A list of the insect types in the collection of the Illinois State Natural History Survey and the Univ. of Illinois. Bulletin of the Illinois State Laboratory of Natural History. 16(4): 137-309.

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CHAPTER III

THE GRASSHOPPER FAUNA OF SOUTHEASTERN GRASSLANDS: A

PRELIMINARY INVESTIGATION

“We came to a large prairie, that was about six miles across it, and in this I saw

the trail which I knowed was made by bear, deer, and turkeys. We went on

through it till we came to a large creek, and the low grounds were all set over

with wild rye, looking as green as a wheat field.” Quote from David Crocket

describing a grassland landscape near the headwaters of the Choctawhatchee

River in southern Alabama in 1815. Originally quoted in Crocket (1834) and

annotated by (Shackford and Folmsbee (1972).

Introduction

When most people think of the historic landscape of the South, they likely think

of vast forests and swamps stretching from the mountains to the coast. However, large

areas of the Southeast were once covered by natural grasslands of various types such as

prairies, glades, savannahs, marshes, sandhills, rock outcrops, and mountain top balds

(Fig.1). For example, longleaf pine savannahs covered approximately 89,600,000 acres

(36,259,833 ha) from Virginia to east Texas (The Longleaf Alliance 2013), and prairies were known to cover at a minimum 4,323,591 acres (1,749,695 ha) in Alabama,

Arkansas, Florida, Louisiana, and Mississippi (Barone 2005a,b, Barone in press, Barone

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unpublished data, Stephenson 2011). These grasslands contribute significantly to the biota of the region; for example, 927 plant taxa are endemic to the longleaf pine ecosystem alone (Sorrie 2011), and Black Belt prairies support more than 1,160 species of moths (Brown 2003 and unpublished data.). However, natural grasslands in the

Southeast have decreased greatly since European settlement, and many are now considered highly imperiled as a result of anthropogenic disturbances, existing only as small remnants of what were once dominant landscape features (Anderson et al. 1999,

Noss 2013). Given their rich biodiversity and conservation importance, grasslands have been an area of emphasis of an ongoing study of the grasshopper fauna of the southeastern United States, which is defined here to include the states of Alabama,

Arkansas, Florida, Georgia, Louisiana, Mississippi, North Carolina, South Carolina, and

Tennessee. This definition was used largely for practical purposes, but also because the

defined area has an identifiable fauna that can be distinguished from other regions.

In my opinion (and I admit that I am a bit biased), a book detailing the natural

history of grasslands would be quite deficient if some mention of grasshoppers was

lacking. I have been studying the grasshopper fauna of grasslands and other naturally

open habitats in the Southeast for the last 13 years and have found that grasshoppers are

often one of the most conspicuous members of the invertebrate fauna in these

communities. Grasshoppers are relatively large (for an insect), are often very abundant in

grasslands, and dazzle the eyes and ears with their colorful and crackling flight displays

adding to the ambiance of the habitat. Additionally, grasshoppers are important

components of temperate grasslands as they are often functionally dominant herbivores in

grassland ecosystems (Joern 1979, Otte 1981, Branson et al. 2006), affect nutrient cycling

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(Belovsky and Slade 2000), are important food items for vertebrate and invertebrate predators (Hamilton and Pollack 1961, Cherrill and Begon 1989, Fowler et al 1991), and are vectors of some vertebrate diseases and parasites (Hill and Goddard 2012). In addition, some species are restricted to remnants of habitats that were once expansive,

such as natural grasslands, and the reduction of these habitats raises concern about the

continued existence of these species (Panzer et al. 2009). In writing on the importance of

species and habitat conservation, Aldo Leopold (1953) stated that “To keep every cog

and wheel is the first precaution on intelligent tinkering.” Given their varied and

important roles, it could be argued that grasshoppers are a vital cog in the mechanism of

grassland function. For this reason I feel there is a strong basis for the inclusion of

grasshoppers in biological inventories and studies of grassland ecosystem function.

Studies of the southeastern grasshopper fauna started in earnest with Samuel

Scudder's (1878) paper on the Orthoptera collected in Florida by J. H. Comstock.

Thereafter, an impressive list of orthopterists have contributed to knowledge of the

regional fauna including John Capinera (and students), Matt Dakin, J. J. Friauf, Morgan

Hebard, Theodore Hubbell, J. McNeil A. P. Morse, Dan Otte, and James Rehn. Though

most of their work is descriptive in nature, enough conclusions can be drawn from their

work and some of my own to 1) explore the uniqueness of the Southeastern grasshopper

fauna in relation to the North American fauna, 2) examine whether the grasshopper fauna

of different types of grasslands in the region varies, and 3) explore and speculate on any

biogeographic affinities of the fauna. However, the information presented here should be

viewed as preliminary in nature. A more thorough treatment awaits a firmer taxonomic

framework (primarily of the Melanoplinae) and more detailed studies of the distribution,

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ecology, and phylogenetics of many of southeastern species. Thus, again I stress that this chapter should be considered only a starting place, not the final word on this topic.

Figure 43 Landscape photographs of various Southeastern grasslands.

A) Black Belt Prairie in Chickasaw Co., MS; B) cedar glade in Davidson Co., TN; C) longleaf pine savannah in Thomas Co., GA; D) sedge dominated wetland, Bibb Co., AL; E) riverine sand dune in Emmanuel Co., GA; F) rock outcrop in Stone Co., AR.

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Here I use the definition of a grassland as employed by Noss (2013) in his treatment of southeastern grasslands to include any natural community or ecosystem in which the herbaceous layer is dominated by grasses, other graminoid plants such as

sedges, and associated forbs. This does not exclude the presence of trees, but trees should

not shade out the herbaceous layer. Grasslands in the Southeast encompass many habitat

types such as prairies, glades, rock outcrops, sandhills, savannas, scrub, and several types

of wetlands. The term “grasshopper” is commonly used to refer to various members of the order Orthoptera. Here I use the term grasshopper to refer to species of “short-horned

grasshoppers” and “lubber grasshoppers” belonging to the families Acrididae and

Romaelidae respectively. The Tettigoniidae, which are often called katydids, meadow

grasshoppers, or long-horned grasshoppers, are likely an important group regionally, but

are not considered here.

The Southeastern Fauna

To answer the first question of the uniqueness of the grasshopper fauna of

southeastern grasslands, one must examine the regional fauna as a whole and then

examine which species inhabit grasslands. Data on grasshoppers associated with

southeastern grasslands come largely from species accounts in numerous taxonomic

works from the region, several faunistic studies, and my own personal collections. To be

considered a grassland inhabitant, a species must primarily be associated with at least one

of habitats described above. Several taxa considered here as grassland inhabitants, such

as Syrbula admirabilis (Uhler), are often associated with grasslands of anthropogenic

origin, such as old fields or roadsides; however, these species are also found in naturally

occurring grasslands and have been able to colonize and adapt to these new habitats. 170

Thus, these species are still considered grassland inhabitants. Intuitively, it makes sense that grasshoppers would be associated with grasslands. However, several species groups, such as the Melanoplus tribulus and M. nigrescens groups, are near endemics or have their center of diversity in the Southeast, but are denizens of forests, woodlands, and forest/grassland interfaces. Further, some species, such as Romalea microptera are

habitat generalists, meaning they occupy in a wide variety of habitats including forests,

swamps, and various types of grasslands. Species that fit this category were not included

in the grassland inhabitant list.

Grasshopper taxonomy is currently undergoing rapid changes as new

morphological and molecular approaches are being applied in revisions, but based on

current taxonomy approximately 751 grasshopper taxa (Acrididae and Romaleidae) are documented from North America north of Mexico (Eades 2013). Of these, 211 taxa

(28% of the North American fauna) occur in the southeastern United States.

Furthermore, more than half of the regional fauna, 130 taxa (62% of the Southeastern

fauna and 17% of the North American fauna) are endemic or are near endemics to the

Southeast, and a significant proportion (82%) of the endemic Southeastern fauna is

associated with grasslands of various types (Tables 2 and 3). Thus, it appears that the

natural grasslands of the southeastern United States are responsible for the majority of the

of the grasshopper diversity in the region. Notably, 71 species (34% of the regional

fauna and 56% of the regional endemics) are associated with longleaf pine

savannahs/sandhills including the five southeastern endemic genera, Aptenopedes,

Floridacris, Floritettix, Eotettix, and Gymnosciretes (Table 3, Fig. 4 A, B, C, and E).

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Most of the endemism is found in the subfamily Melanoplinae, which includes

many brachypterous (short-winged) or apterous (no wings) species that are flightless and

are restricted to specific habitats, which often results in many of the regional endemics

having small geographic distributions. For example, Melanoplus dakini Hilliard (Fig. 2A)

is known only from three parishes in Louisiana where it inhabits open pine forests with a

grassy understory; Melanoplus ingrami Hill (Fig.1B, Fig. 2B) is endemic to the cedar glades in the Central Basin of Tennessee; Melanoplus stegocercus Rehn and Hebard (Fig.

2D) is known only from dune or sandhill sites in east-central Georgia, and Melanoplus

pygmaeus Davis (Fig. 2C) is found only in sandhill/dune habitats in southern Alabama

and northwest Florida.

Table 3 Proportion of North American and southeastern grasshoppers that occur in southeastern grasslands, are endemic to the Southeast, are endemic to natural grasslands in the Southeast, and are longleaf endemics.

Number of % of North % Southeastern species American Fauna Fauna North American Fauna 751 100 Occur in the Southeast 211 28 100 Occur in open habitats in the 173 23 82 Southeast (natural and disturbed) Endemic to the Southeast 130 17 62 Endemic to natural grasslands of the 111 15 53 Southeast Longleaf pine system endemics 74 10 34 Only Acrididiae and Romalidae are considered.

Two well-studied species groups containing short-winged, grassland-inhabiting

species in the region are the Melanoplus puer species group and the Melanoplus scudderi

species group (sensu lato). The Melanoplus puer species group consists of smaller

flightless species that inhabit the sandhill/scrub habitats of sand ridges in peninsular

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Florida and longleaf pine ecosystems (Hubbell 1932, Folkerts and Deyrup 1993, Squitier et all 1998, Otte 2012). The sand ridges inhabited by these flightless grasshoppers are

relatively elevated compared to most of Florida and were likely islands during periods of

higher sea levels in the past. Although sea levels are much lower now, these

grasshoppers are still isolated from each other by areas of habitats that are unsuitable to

them, such as oak hammocks, swamps, and agricultural areas. As a result of their

interesting biogeographic relationships, this group of grasshoppers has been used as a

model of evolutionary divergence, serving as one of the strongest pieces of evidence of

the relative isolation of various sand ridges systems in peninsular Florida (Deyrup 1996).

The Melanoplus scudderi species group (sensu lato) contains several species with

interesting distributions in the region, including five species that are endemic to sandhills

along the Fall Line in Georgia and the Carolinas (Fig. 3). Of these five species, two are

largely sympatric and the rest are allopatric with their ranges being segregated by large

river systems of the region (Hill in prep). Additionally, several other species in the

scudderi group are associated with longleaf pine habitats on the Coastal Plain.

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Figure 44 Restricted distributions of some flightless grasshoppers in the Southeast.

A) Melanoplus dakini, B) M. ingrami, C) M. stegocercus, D.) M. pygmaeus. Produced in Google Earth Pro®.

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Figure 45 Members of the Melanoplus scudderi group (s.l.) endemic to sandhills along the Fall Line.

White circles = M. muscogee, blue circles = M. chattahoochee, red circles = yellow circle = M. carolinensis, and green circles = M. coreyi. Produced in Google Earth Pro®.

Affinities With Western Grasslands

Different grassland types within the southeastern region support distinct biotas

that have biogeographic affinities with the grasslands of the west. Whereas the biota of

the prairies of the Southeast are associated more with the tall grass prairie, the biota of

the sandhill habitats are associated more with the arid habitats of the southwestern United

States. The sandhills of the Southeast often contain one or two species of a diverse

western group of congeners, which suggests that the eastern species are derived from the

western group (Folkerts et al 1993). This pattern has been documented for numerous

other taxa, both extant and extinct (see Noss 2013 for a summary). This relationship has

been explained by a possible corridor of savanna habitat that developed during the 175

Miocene and persisted into the early Pleistocene along the coast of the Gulf of Mexico, with this corridor allowing an exchange of xeric-adapted biota between the semiarid west

and the Southeast (Webb 1977, Emslie and Czaplewski 1999, Noss 2013).

Grasshoppers are no exception to this western species in the east phenomenon.

Several southeastern endemic grasshopper taxa are sole members of “western” or

Neotropical genera or families including Achurum carinatum (Walker) (Fig 4C),

Amblytropidia mysteca (Saussure), and Romalea microptera (Beauvois). Additionally, at

least 10 widely distributed western species have disjunct populations associated with

grasslands in the Southeast. The primarily western genus Hesperotettix contains three

sandhill inhabiting species (H. floriidensis Morse, H. gemmicula Hebard, and H. osceola

Hebard) that are endemic to the Southeast. Hespertotettix viridis Thomas, a grassland

species that is widely distributed in the west, has two subspecies Hesperotettix viridis

brevipennis (Thomas) and H. v. pratensis Scudder that inhabit grasslands in the east. The

genus Eritettix is most diverse in the western U.S, but two species, Eritettix simplex

(Scudder) and Eritettix obscurus (Scudder) (Fig. 6F) inhabit grasslands of the Southeast.

Additionally, Melanoplus angustipennis (Dodge) and Melanoplus bispinosus Scudder are

species that are typically associated with sandy or arid habitats in the west and occur in

sandhill habitats in the Southeast. Morse (1904) suggested that the genera Arphia,

Trimeroptropis, Spharagemon (Fig. 7) were likely of western origins. Given the habitat

preferences of most of the species in these genera, I would agree with his assertion. See

Table 3 for a summary of taxa with disjunct distributions and their habitat associations in

the Southeast.

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Figure 46 Some spur-throat grasshopper species that inhabit southeastern grasslands.

A) Floritettix borealis, B) Gymnoscirtetes pusillus, C), Aptenopedes sphenariodes apalachee D) Hesperotettix viridis brevipennis, E) Campylacantha olivacea. F) Eotettix pusillus.

The grasslands of the Southeast are well known for supporting a distinct flora and

for harboring rare or endangered vertebrates such as the indigo snake (Drymarchon

couperi), gopher tortoise (Gopherus polyphemus), and Bachman’s sparrow (Peucaea

aestivalis), but as indicated above, they also support unique invertebrate communities.

One of the most common misconceptions about grasslands in the Southeast is that they

are of anthropogenic origin, having arisen though burning and clearing by Native

Americans. Other authors have pointed out that this is not the case given the high level of

endemism and biodiversity in these systems and the amount of time it would take for

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these systems to evolve (Noss 2013). The results from the analysis here underscore this conclusion.

Figure 47 Some spur-throat grasshopper species that inhabit southeastern grasslands.

A) Schistocerca rubiginosa (Harris), B) Melanoplus ingrami, C), Melanoplus pygmaeus, D) Melanoplus stegocercus E) Melanoplus savannah, F) Paroxya clavuliger.

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Figure 48 Some slant-faced and tooth-legged grasshopper species that inhabit southeastern grasslands.

A) Dichromorpha elegans, B) Mermiria picta, C) Achurum carinatum, D) Pseudopomala brachyptera, E) Metaleptea brevicornis F) Eritettix obscurus.

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Figure 49 Some band-winged grasshopper species that inhabit southeastern grasslands.

A) Psinidia fenestralis, B) Trachyrhachys kiowa, C), Trimerotropis saxatalis, D) Spharagemon cristatum, D) Mermiria picta, E) Arphia xanthoptera. F) Pardalophora phoenicoptera.

Grasshopper Composition of Grasslands

The grasslands of the Southeast occur in a variety of environmental conditions

(edaphic, hydrologic, elevation, etc.) and geographic regions, which could likely

influence their faunistic compositions. For example, Barone (2005a) found that different

prairie types in the eastern U.S. tended to have similar floristic composition to their

nearest neighbors. A database of grasshoppers was constructed with 117 species from 16

distinct grassland types from around the region to investigate their faunistic relationships

from the various grasslands in the Southeast. For comparison the database included the

faunal list from the Konza Prairie in Kansas as an example of a western grassland and 180

from cedar-hardwood forests of the Central Basin of Tennessee as an open forest type.

The database was constructed from using lists gathered from the literature (referenced in legend) as well as unpublished data of JGH.

A cluster analysis was performed on the database using PC-ORD software

McCune and Mefford (2011) with the Flexible Beta (β= -0.25) as the linkage method and

the Sorensen (Bray-Curtis) as the distance measure as recommended by McCune and

Grace (2002).

All the grasslands with the exception of the xeric limestone prairies of the

Western Highland Rim of Tennessee and northwestern Alabama branched independently,

suggesting that each habitat type possessed a relatively distinct fauna for the most part

(Figure 8 .). Geographic region appears to be important in structuring the dendrogram.

The top cluster is made of grasslands (prairies and glades) from Alabama, Mississippi, and Tennessee, the second cluster consists of the Arkansas glades/prairies and Konza

Prairie, and the third major cluster is comprised of sandhill habitats from Florida and the southern half of Georgia. However, habitat type also appears to have an influence, as the sites represented in the first cluster are found on limestone or chalky clay soils, and the third major cluster of habitats all have well drained sandy soil. Balds, cedar-hard wood forests and marshes are found on long branches of the dendrogram, which is an indicator of their very distinctive faunas. The fauna of ruderal grasslands of the Welka region

Florida includes habitat generalists from the region and appears to be a subset of the

sandhill/scrub fauna. The cedar hardwood forests and southern Appalachian balds are

distantly clustered together, which is likely a result of both habitats having depauperate

faunas partly composed of species that inhabit open forests or forest edges.

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The cluster analysis illuminates some interesting patterns in the species composition of grasslands of the region with sandhills, prairie, balds and forests all distinguishing themselves. A more detailed look into the faunas of these grasslands is presented below.

Figure 50 Cluster analysis results of grasshopper species composition from 15 different grassland types across the southeastern United States and Konza Prairie in Kansas.

Data are for 117 species. Abbreviations: BBP = Black Belt prairie (Hill 2007), BCG = Bibb County Glades (Hill 2007 and recent collections), XPL=xeric limestone prairies of the Central Basin of Tennessee (Hill 2012), TCG = Tennessee Cedar Glades in the Central Basin (Hill 2012), TNWR = xeric limestone prairies of the western highland rim of Tennessee (Hill in prep), ALMV= xeric limestone prairies in the Moulton Valley of NW Alabama (in prep), AROG = Glades in the Ouachita Mountains of Arkansas (Hill unpublished data Morse 1907), KPR = Konza Prairie (Wright et al. 2005), OD= Ohoopee Dunes of Georgia (Hill 2009), GAFL = Fall Line sandhills in Georgia (Hill in prep), FLSH = sandhills in Florida (Friauf 1953, Squitier and Capinera 2002), FLSC = Florida scrub Friauf 1953, Squitier and Capinera 2002), FLRG = ruderal grasslands in Florida (Friauf 1953, Squitier and Capinera 2002), CHF = cedar hardwood forests of the Central Basin of Tennessee, SMKAB = balds from Great Smoky Mountain National Park in Tennessee and North Carolina (Hill unpublished data), FLM = marshes in Florida along the St Johns River (Friauf 1953).

Calcareous Grasslands and Glades

The first large cluster of the dendrogram (Figure 8) is composed of calcareous

grasslands, including blackland prairies (MS: Black Belt and Jackson Prairie); xeric

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limestone prairies (AL: Moulton Valley, TN: Central Basin and Western Highland Rim);

glade habitats (AL: Bibb County, TN: Central Basin), Arkansas glades/prairies, and the

Konza Prairie. There is some obvious geographic structuring of the cluster, with the

Arkansas glades, and Konza prairies lying at the eastern most limits for some “western” taxa, and the eastern grasslands containing widespread eastern species. The grasslands in this cluster share many grasshopper species, but one interesting aspect is the occurrence of native grassland specialists that are predominantly distributed in the Great Plains, namely Paratylotropidia brunneri Scudder, Pseudopomala brachyptera (Scudder) (Fig

6D), and Trachyrhachys kiowa (Thomas) (Fig. 7B). (See Table 3 for a summary.)

Brown (2003) examined the distributions of moths and butterflies (Lepidoptera) and a few other insects from Black Belt prairies in Mississippi and observed numerous examples of species primarily distributed in the Great Plains. This observation led Brown to hypothesize that a grassland corridor existed between the Black Belt and the Great

Plains prior to and following the Wisconsin Glaciation that provided dispersal routes for

grassland species, and that the Black Belt may have served as refugium for grassland

species during this glacial episode. Further, his hypothesis proposed that the fauna of the

tall grass prairie in the post-glacial Midwest was more likely derived from the refugium

in the Black Belt than from the southwestern United States. The grasshopper data

presented here and below add support Brown’s hypothesis. Additionally, the flora of

these southeastern prairies is similar to that of other eastern tall grass prairies with the

dominant grasses being little bluestem (Schizachyrium scoparium), big bluestem

(Andropogon gerardii), switch grass (Panicum virgatum), Indian grass (Sorghastrum

nutans) and drop seed (Sporobolus spp) and the common forbs being rosinweeds

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(Silphium spp.), prairie coneflower (Ratibida pinnata), prairie-clovers (Dalea spp.), and blazing stars (Liatris spp.) (See Barone and Hill 2007, DeSelm and Murdock 1993, and

Lawless et al. 2006, and Webb et al. 1997). Given the combination of these biotic associations, the prairies of the Southeast should be included as part of the Eastern Tall

Grass Prairie biome.

Longeleaf Pine Ecosystem (Fall Line Sandhills, Ohoopee Dunes, Florida Sandhill/Scrub)

The second major cluster of the dendrogram is composed of sandhill/scrub habitats. Data for these habitats were taken from several studies, and it should be noted that these represent subsets of the longleaf pine ecosystem. The clustering of these habitats is likely due to the presence of widespread species that are restricted to xeric sandy soil, such as Melanoplus angustipennis (Dodge), Melanoplus bispinosus Scudder,

Psinidia fenestralis (Serville) (Fig7A), Schistocerca rubiginosa (Fig. 5A), Spharagemon

cristatum (Scudder) (Fig. 7D), Spharagemon marmorata picta (Scudder), and

Trimerotropis maritima (Harris) and a high number of endemic species associated with

sandhills on the Coastal Plain. Structure in this cluster is organized by geography with

the Georgia and Florida sites separated from each other. This is likely due to the high

level of endemism associated with sandy habitats in Florida, such as seen in the

Melanoplus puer group (mentioned above).

Species shared between Prairies and Sandhills

The distant linkage of prairies and sandhills is likely a result of the sharing of

many widespread grassland generalists, such as Arphia xanthoptera (Burmeister) (Fig.

7E), Dissosteira carolina (Linnaeus) Melanoplus sanguinipes vulturnus Gurney and 184

Brooks, and Pardalophora phoenicoptera (Burmeister) (Fig 7F) among others.

Additionally, Eritettix simplex (Scudder), Melanoplus impudicus Scudder, Mermiria

bivittata (Serville) (Fig. 6B), and Mermiria picta (Walker) are patchily distributed,

grassland specialists shared between these broad habitat categories

Cedar Hardwood Forests

The third branch is the cedar hardwood forests of the Central Basin of Tennessee.

This was the only forest type included in the analysis, and was done so because this

habitat is relatively open and has a well-developed herbaceous layer. However, the

grasshopper fauna of this habitat is composed of species associated with edges or closed

canopies, such as Melanoplus gracilis (Bruner), Melanoplus walshii Scudder, and

members of the Melanoplus tribulus/obovatipennis species group. This association is

typical of most open forest/edge communities throughout the region.

Southern Appalachian Balds

Clustering somewhat with the cedar hardwood forests are the southern

Appalachian balds. The data used in this survey were collected in Great Smoky

Mountains National Park. These balds tend to be smaller and more disturbed than balds

to the north (Noss 2013), thus these results may not be applicable to all balds in eastern

U.S.. Balds in the Great Smoky Mountains National Park do not appear to have any

grassland specialists, and the fauna is mostly made of forest/edge species, such as

Melanoplus walshii and members of the Melanoplus viridpes species group. Perhaps the

cooler temperatures of these high elevations allow these sylvan species to escape the

shade of the forest canopy and exploit the lush herbaceous vegetation of the balds. The

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grasshopper fauna of larger and less disturbed balds should be studied to see if this is the typical fauna or if there are species that may be bald specialists.

Marshes

The final branch of the dendrogram represents Spartina/saw grass Marshes along

the St. Johns River in Florida. Though these data are from Florida, the species represented here are typical of many open wetlands across the Southeast, with species in the genus Paroxya (Fig. 5F), Dichromorpha (Fig. 6A), and Mermiria intertexta Scudder

being represented (JGH pers obs.). Further investigations of the grasshopper fauna of

marshes and wetlands throughout the region will likely illuminate a greater diversity of

faunal associations as there are more species that inhabit wet grasslands throughout the

region that are not present in these Florida marshes, including Leptysma marginicollis

(Serville), Metaleptea brevicornis (Johannson) (Fig.6E), Stenacris vitreipennis

(Marschall), and Stethophyma celata Otte.

Conservation/Management

Given the unique and diverse grasshopper fauna in the southeast, it is apparent

that the greatest threat to grasshopper biodiversity in the region is the loss and

fragmentation of grassland systems. Indeed, many of the southeastern grasslands are

considered endangered or imperiled as a result human activities. Surveys conducted by

the General Land Office in the 1830’s suggest that prairies once covered at least 144,000

hectares in the Black Belt of Mississippi and Alabama (Barone 2005). Since that time,

more than 99% of these prairies have been lost to agriculture and urban development

(Noss et al. 1995). Remaining remnants of prairie are threatened by further development,

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erosion, and the encroachment of eastern red cedar, Juniperus virginiana L.,

(Cupressaceae), the latter probably as a result of fire suppression. Likewise, the cedar

glades and xeric limestone prairies of the Central Basin of Tennessee are considered

highly imperiled due to anthropogenic disturbances including fire suppression,

recreational driving, and the suburban sprawl of the cities of Nashville and Lebanon

(Noss et al.1995, Nielmiller et al. 2011). Even the once vast longleaf pine ecosystem that

covered an area from southeastern Virginia to eastern Texas has been reduced to only 3%

of its pre-settlement extent as a result of tar and pitch production, logging, clearing for

agriculture, and modern forestry practices, namely site prep practices and stocking

densities (Frost 1993, Platt 1999).

As mentioned above, many of the flightless grassland specialists have small

geographic distributions, and given their narrow habitat breadth (such as sandhills), their

populations are often isolated by inhospitable natural features such as forests, rivers, and

swamps. Thus, these species often occupy relatively small areas, making them more

vulnerable to extinction.

However, their reliance upon a single habitat and small geographic area may also

have positive conservation implications. As grasshoppers are primary consumers, they

are often abundant in many grassland systems, and a small remnant can often be

inhabited by a significant number of individuals, such that even establishing a small

preserve, sometimes less than several hectares (or acres), could potentially protect an

entire population. Moreover, a series of small preserves could conceivably provide

adequate protection the species, as well as saving examples of their habitat and associated

biota.

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The effects of habitat management techniques on grassland species in the

Southeast has not been well-documented. Management practices aimed at maintaining grassland vegetation such as burning and grazing likely have a significant impact on the grasshopper fauna. For example, a species such Pseudopomala brachyptera (a grassland

endemic) that overwinters in the nymphal stage may be sensitive to early season burns

that are currently favored by many land managers. There is little doubt that fire is

important in maintaining the habitat for grassland species; however, for grasshoppers, the

level, timing or frequency of burning relative to being beneficial or disruptive is

unknown, and the answer is likely dependent on the species and grassland system in

question. That being said, many grasslands are fire-maintained systems, and as members

of that system, grasshoppers have several adaptations to cope with fire. For example,

species that are capable of flight often disperse away from the fire. Large numbers of

grasshoppers have been observed flying away from a prescribed fire in a longleaf pine

savannah (Matt Elliott pers. com.). In addition, short-winged, flightless species may

climb trees to avoid flames, particularly in longleaf savannahs. During a visit to a sandhill

at the Savannah River Site in the fall of 2012, numerous individuals of Melanoplus

savannah were observed perched about 4 meters above the ground on longleaf pine

trunks immediately after a burn (J.G. Hill pers obs.). Finally, though not a means to

escape fire, grasshoppers may be adapting to frequently burned areas by developing

darker coloration, which makes them more cryptic against the charred surface of their

habitat. On several occasions, dark morphs of Orphulella pelidna, Melanoplus coreyi, and Melanoplus strumosus have been observed in areas that experience frequent burning.

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Conclusion

Grasshoppers are ideal study organisms in temperate grasslands considering their biological importance, interesting distributions, relative abundance, and ease of sampling.

The grasslands of the Southeast offer an astounding array of interesting questions for researchers from a variety of fields. Biogeographers and systematists could answer questions about origins of the fauna of these grasslands and the dispersal routes taken by disjunct species, and further research could investigate the faunal relationships between various grassland types. Ecologists could investigate habitat requirements and life histories of many grassland endemic species, particularly those that may be of conservation concern. Conservationists and management oriented researchers could study the effects that various management issues, particularly those involving the effects of habitat fragmentation, importance of preserve size and connectivity, timing of prescribed burning, effects of disturbance and, in longleaf pines systems site prep and stocking rate of new plantations.

This introductory synthesis of the grasshopper fauna of grasslands in the southeastern United States further demonstrates that these varied and distinct grasslands are rich in endemic species that contribute significantly to the biodiversity of the region

and that of North America as well as providing further corroborating evidence that these

systems are of natural origin. Thus the continued conservation of these systems along

with the study of their biota are important for as they represent an important part of our

regional natural heritage. It is my hope that the information presented here inspires

further interest in research and conservation of southeastern grasslands.

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Table 4 List of grasshopper species that occur in the southeastern United States.

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic leidae Romalea microptera (Beauvois) X Melanoplinae Aptenopedes apalachee Hebard X X X X Aptenopedes chefixico Otte X X X X Aptenopedes chiaha Otte X X X X Aptenopedes clara Rehn X X X Aptenopedes menawai Otte X X X X Aptenopedes rufovittata Scudder X X X X Aptenopedes sphenarioides Scudder X X X X Aptenopedes sphenax Otte X X X X Aptenopedes yoholoi Otte X X X X Booneacris variegata (Scudder) X Campylacantha olivacea (Scudder) X Dendrotettix australis (Morse) X Dendrotettix quercus Riley Dendrotettix zimmermanni X (Saussure) Eotettix hebardi Rehn X X X X Eotettix palustris Morse X X X X Eotettix pusillus Morse X X X X Eotettix signatus Scudder X X X X Floridacris etami Otte X X X X Floridacris mariona Otte X X X X Floridacris orangina Otte X X X X Floridacris robusta (Hebard) X X X Floritettix aptera (Scudder) X X X X Floritettix borealis (Hebard) X X X X Floritettix calusa Otte X X X X

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Table 4 (continued)

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic Floritettix coquinae (Hebard) X X X Floritettix flodidana Otte X X X X Floritettix hadjoi Otte X X X X Floritettix holatamico Otte X X X X Floritettix hubbelli Ottei X X X X Floritettix ocilla Otte X X X X Floritettix osceola Otte X X X X Floritettix nigropicta (Hebard) X X X X Floritettix saturiba (Hebard) X X X X Floritettix simplex (Hebard) X X X X Gymnoscirtetes morsei Hebard X X X X Gymnoscirtetes pusillus Scudder X X X X Hesperotettix floriidensis Morse X X X X Hesperotettix gemmicula Hebard X X X X Hesperotettix osceola Hebard X X X X Hesperotettix viridis brevipennis X (Thomas) Hesperotettix viridis pratensis X Scudder Melanoplus acidocercus Hebard X X X X Melanoplus acrophilus Hebard X X X Melanoplus alabamae Hebard X X X Melanoplus adelogyrus Hubbell X X X Melanoplus angustipennis (Dodge) X Melanoplus apalachicolae Hubbell X X X Melanoplus attenuatus Scudder X X X Melanoplus arkansas Hill X X X Melanoplus australis Morse X X X X Melanoplus baronei Hill X X X Melanoplus bispinosus Scudder X Melanoplus bivitttatus (Say) X Melanoplus cantralli Dakin X X X X Melanoplus carnegiei Morse X X X Melanoplus carabellae Otte X X X Melanoplus carolinensis Hill X X X X Melanoplus celatus Morse X X X Melanoplus chattahoochee Hill X X X X

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Table 4 (continued)

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic Melanoplus cherokee Hebard X Melanoplus childsi Otte X X X X Melanoplus clypeatus (Scudder) X X X Melanoplus confusus Scudder X X X Melanoplus coreyi Hill X X X X Melanoplus dakini Hilliard X X X Melanoplus davisi Hebard X X X X Melanoplus deceptus Morse X X Melanoplus decoratus Morse X X Melanoplus decorus Rehn and Hebard X X X Melanoplus delaware Hebard Melanoplus devius Morse X Melanoplus differentialis differentialis

(Thomas) Melanoplus differentialis nigricans

Cockerell Melanoplus divergens Morse X X X Melanoplus eurycercus Hebard Melanoplus fasciatus (Walker) X Melanoplus folkertsi Hill X X X Melanoplus femurrubrum (DeGeer) X Melanoplus foedus Scudder X Melanoplus forcipatus Hubbell X X X X Melanoplus foxi Hebard X X X X Melanoplus furcatus Scudder X X X Melanoplus gracilis (Bruner) Melanoplus gurneyi Strohecker X X X X Melanoplus harrisi Hill X Melanoplus hubbelli Hebard Melanoplus impudicus Scudder X Melanoplus inconspicuus Caudell X Melanoplus ingrami Hill X X X Melanoplus keeleri keeleri (Thomas) X Melanoplus keeleri luridus (Dodge) X Melanoplus kissimmee Otte X X X X Melanoplus longicornis Saussure X X X Melanoplus mirus Rehn and Hebard X X X

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Table 4 (continued)

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic Melanoplus mississippi Hill X Melanoplus morsei Blatchley Melanoplus muscogee Hill X X X X Melanoplus nanciae Deyrup X X X X Melanoplus nossi Hill X Melanoplus nubilus Rehn and Hebard X X X X Melanoplus nigrescens (Scudder) X Melanoplus ordwayae Deyrup X X X Melanoplus ouachita Hill X X X Melanoplus ozarkensis Hill X Melanoplus pachycercus Hebard X X X Melanoplus packardii Scudder X Melanoplus peatus Otte X X X Melanoplus pegasus Hebard X X X Melanoplus peninsularis Hubbell X X X Melanoplus ponderosus (Scudder) X X X Melanoplus viola Thomas Melanoplus primestivus Dakin X Melanoplus propinquus Scudder X X X Melanoplus puer (Scudder) X X X X Melanoplus punctulatus arboreus

Scudder Melanoplus punctulatus punctulatus

(Scudder) Melanoplus pygmaeus Davis X X X X Melanoplus quercicola Hebard X X X X Melanoplus querneus Rehn and Hebard X Melanoplus relictus Hill X X X X Melanoplus rotundipennis (Scudder) X X X X Melanoplus sanguinipes vulturnus X Gurney and Brooks Melanoplus savannah Hill X X X X Melanoplus scapularis Rehn and X X X X Hebard Melanoplus scudderi (Uhler) X Melanoplus sebringi Otte X X X X Melanoplus seltzeri Hill X X X

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Table 4 (continued)

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic Melanoplus seminole Hubbell X X X X Melanoplus serrulatus Hebard X X X Melanoplus similis Morse X Melanoplus stegocercus Rehn and X X X X Hebard Melanoplus strumosus Morse X X X X Melanoplus sylvaticus McNeil X Melanoplus sylvestris Morse X Melanoplus symmetricus Morse X X X Melanoplus tepidus Morse X Melanoplus tequestae Hubbell X X X X Melanoplus texarkana Hill X X X X Melanoplus tumidicercus Hubbell X X X X Melanoplus tunicae Hebard X Melanoplus tribuloides Morse X Melanoplus tribulus Morse Melanopluss viola Thomas X Melanoplus vulnus Eades X X X Melanoplus walshii Scudder Melanoplus withlacoocheensis Squitier X X X X and Deyrup Melanoplus zento Otte X X X Paratylotropidia beutenmuelleri Morse X Paratylotropidia brunneri Scudder X Paratylotropidia morsei Rehn and Rehn X Paroxya atlantica Scudder X Paroxya clavuliger (Serville) X Paroxya hooseri (Blatchley) X Paroxya paroxyoides (Scudder) X X X Paroxya recta Scudder X X X Leptysma marginicollis (Serville) X Stenacris vitreipennis (Marschall) X Cyrtacanthacridnae Schistocerca alutacea (Harris) X Schistocerca americana (Drury) X

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Table 4 (continued)

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic Schistocera ceratiola Hubbell and X X X X Walker Schistocerca damnifica (Saussure) X Schistocerca lineata Scudder X Schistocerca obscura (Fabricius) X Schistocerca rubiginosa (Harris) X X X Gomphocerinae Achurum carinatum (Walker) X X X Ageneotettix deorum Scudder X Amblytropidia mysteca (Saussure) X Boopedon auriventris McNeil X Chloealtis conspersa Harris Chorthippus curtipennis (Harris) X Dichromorpha elegans (Morse) X X X Dichromorpha viridis (Scudder X Eritettix obscurus (Scudder) X X X X Eritettix simplex (Scudder) X Mermiria bivittata (Serville) X Mermiria intertexta Scudder X X X Mermiria picta (Walker) X Orphulella pelidna (Burmeister) X Orphulella speciosa (Scudder) X

Pseudopomala brachyptera (Scudder) X

Stethophyma celata Otte X X Syrbula admirabilis (Uhler) X Acridinae Metaleptea brevicornis (Johannson) X Oedipodinae Arphia granulata (Sassure) X X X X Arphia simplex Scudder X Arphia sulphurea (Fabricius) X Arphia xanthoptera (Burmeister) X Chortophaga austrailor Rehn & X X X X Hebard Chortophaga viridifasciata (DeGeer) X Dissosteira carolina (Linnaeus) X

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Table 4 (continued)

Endemic and SE Grassland Grassland Longleaf Species Endemic Specialist Specialist Endemic Encoptolophus soridus (Burmeister) X Hippiscus ocelote (Saussure) X Pardalophora apiculatus (Harris) X Pardalophora phoenicoptera X (Burmeister) Psinidia fenestralis (Serville) X Spharagemon bolli Scudder Spharagemon crepitans (Saussure) X X X Spharagemon cristatum (Scudder) X Spharagemon equale (Say) X Spharagemon marmorata picta X (Scudder) Trachyrhachys kiowa (Thomas) X Trimerotropis maritima (Harris) X Trimerotropis saxatalis McNeil X Totals: 211 species. 130 173 111 71

Compiled from Hill and Dakin 2011, Otte 2012, Hill 2014, and Otte 2014. X’s indicate if a species is endemic to the region, specializes in grasslands or open habitats, and is endemic to either grassland or longleaf pine savannah ecosystems. Taxonomy follows Eades et al. (2013).

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Table 5 Grassland-specific grasshopper taxa with notes on their distribution pattern and habitat preferences.

Taxa Pattern Grassland type Achurum carinatum (Walker) Sole member of a western Various natural and genus in the SE anthropogenic grasslands on the Coastal Plain. Amblytropidia mysteca Sole member of a western Various natural and (Saussure) genus in the SE anthropogenic grasslands. Aptenopedes Genus endemic to Open pine forests, AL, FL, and GA that is Prairies in Florida comprised of 9 species Arphia Genus likely of western Various natural and origin represented by 4 anthropogenic grasslands. species in the SE Campylacantha olivacea Sole member of a western Prairies – associated with (Scudder) (Fig. 4E) genus in the SE. Species Ambrosia psilostachya widespread in west, and restricted to natural grasslands in the Southeast. Eotettix Genus that is endemic to the Longleaf pine forests and SE and is comprised of 3 sandhills species Eritettix simplex (Scudder) Widely distributed in West, Prairies and sandhills. disjunct populations associated with grasslands in the SE Eritettix obscurus (Scudder) Member of a genus that is Scrub and sandhill in widely distributed in the peninsular Florida. West. Endemic to Florida Floridacris Genus of 4 species that Sandhill/scrub endemic to Florida Floritettix Genus endemic to Georgia Longleaf pine savannahs, and Florida that is comprised Sandhill/scrub of 13 species Gymnoscirtetes Genus comprised of two Longleaf pine savannahs species that are endemic (often mesic areas) and sandhills. Hesperotettix Genus likely of western Prairies, sandhills, and origin represented by three Florida glades and species endemic to the SE, prairies. and one species that is widely distributed in West Melanoplus angustipennis Widely distributed in West, Sandhills. (Dodge) disjunct populations in SE

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Table 5 (continued)

Taxa Pattern Grassland type Melanoplus bispinosus Scudder Widely distributed in West, Sandhills disjunct populations in SE Melanoplus impudicus Eastern U.S. Various naturally open habitats Melanoplus puer species group Group that is endemic to the Sandhills and scrub. Coastal Plain Melanoplus scudderi species Group that has apparently Various natural and group speciated in different anthropogenic grasslands. grassland types in the SE Mermiria bivittata (Serville) Widely distributed and Prairies and sandhills. common in West, disjunct populations associated with grasslands in the SE Mermiria picta (Walker) Widely distributed in West, Prairies and sandhills. disjunct populations associated with grasslands in the SE Paratylotropidia brunneri Widely distributed in West, Prairies. Scudder disjunct populations associated with prairies in TN. Pseudopomala brachyptera Widely distributed in West, Prairies. (Scudder) disjunct populations associated with grasslands in the SE Romalea microptera (Beauvois) Sole member of a western or Various habitat types, family in the SE including some forested. Spharagemon Genus likely of western Sandhills and open origin represented by 5 forests. species in the SE Trachyrhachys kiowa (Thomas) Widely distributed in West, Prairies, glades, and rock disjunct populations outcrops. associated with grasslands in the SE Trimeroptropis (Thomas) Genus likely of western Sandhills and rock origin represented by 2 outcrops. species in the SE

198

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APPENDIX A

KEY TO THE SPECIES OF THE MELANOPLUS SCUDDERI AND CARNEGIEI

SPEICE GROUPS

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Key to the Scudderi and Carnegiei species groups

1. Male genitalia with sheath reduced, covering only the base of the valves; apodemes of

cingulum with apicies bent medially (See Fig. 35C-F); cerci generally triangular,

except in M. savannah and M. carolinensis; southeastern United States Carnegiei

group

— Male genitalia with sheath produced; apodemes of cingulum straight (Fig 1C-F); cerci

generally falciform, except in M. davisi and M. quercicola; widespread across

eastern United States ...... Scudderi group

Key to species of the Melanoplus scudderi group based on male genitalia and distribution.

1. Occurring east of the Mississippi River ...... 2

— Occurring west of the Mississippi River ...... 9

2. Cerci falcate with rounded tips (Fig 1A-B) ...... 3

— Cerci triangular (Figs 9A-B, 10A-B) ...... 8

3. Genitalia with dorsal valves curved laterally (Figs 1C-F, 5C-F, 12C-F) ...... 4

— Genitalia with dorsal valves curved medially or dorsally (Figs 6C-F, 7C-F, 9C-F) 6

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4. Curved portion of dorsal valves short and not apically decurved (Figs 1C-I, 2A-B);

occurring across eastern United States, excluding the Mississippi River alluvial

valley and areas of southern Alabama and Georgia, northern Florida ...... scudderi

— Curved portion of dorsal valves long and apically decurved ventrally (Figs 1C-I,

2A-B); occurring in Mississippi alluvial valley and in southern Alabama and

Georgia and northern Florida ...... 5

5. Occurring in southern Alabama to southwest Georgia and northwest Florida;

genitalia as in

Fig 5C-I ...... folkertsi

— Occurring in Mississippi River alluvial valley; genitalia as in Fig. 12C-I mississippi

6. Occurring along Fall Line of western Georgia; genitalia with dorsal valves curved

dorsally (Fig 9C-F); ......

...... muscogee

— Not occurring near Fall Line of western Georgia; genitalia with dorsal valves curved

medially (Figs 6C-F, 7C-F) ...... 7

7. Northern Florida and southwest Georgia; genitalia as in Fig. 6C-I ...... relitcus

— Fall Line of North Carolina; genitalia as in Fig 7C-F ...... coreyi

8. Cerci with apex acutely triangular and straight (Fig 10A-B); extreme southern

Alabama and the panhandle of Florida; genitalia as in Fig. 10C-I ...... davisi

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— Cerci with apex rounded and often curved medially (Fig 9A-B); southern Georgia

and northern Florida; genitalia as in Fig 9C-I ...... quercicola

9. Specimen from Arkansas ...... 10

— Specimen from Arkansas ...... 17

10. Male genitalia with dorsal valves curved laterally (Figs 1C-F, 12C-F) ...... 11

— Male genitalia with dorsal valves not curving laterally (Figs. 14-16C-I, 18-19C-I,

24C-I) ...... 12

11. Curved portion of dorsal valves short and straight (Figs 1C-F); widespread species,

occurring mostly through the central (north to south) portion of Arkansas

...... scudderi

— Curved portion dorsal valves long and apically decurved ventrally; occurring in

Mississippi alluvial plain ...... mississippi

12. Dorsal valves bowed laterally and ventrally (Figs. 13C-F, 24C-F) ...... 13

— Dorsal valves not bowed laterally and ventrally (Figs. 15-6C-F, 18-19C-F)/...... 14

13. Dorsal valves long, strongly bowed laterally and ventrally with apices recurved

dorsally; ventral valves not dorso-ventrally expanded; occurring north of the

Arkansas River in western Arkansas; genitalia as in Fig. 24C-I / ...... ozarkensis

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— Dorsal valves short, slightly bowed laterally and ventrally with apices not curved

dorsally; ventral valves dorso-ventrally expanded; occurring in the Arkansas

River Valley, south of the Arkansas River; genitalia as in Fig 14C-I .... arkansas

14. Dorsal valves strongly decurved apically (Fig. 18C-F, Fig 27C-F) ...... 15

— Dorsal valves bowed dorsally (Fig. 15C-F, 19C-F) ...... 16

15. Genitalia with dorsal valves thin, ventral valves lightly sclerotized and extending

approximately half the length of the dorsal valves (Fig. 18C-I); occurring in

southeastern Ouachita Mountains ...... seltzerae

— Genitalia with dorsal valves thick; ventral valves fully sclerotized and subequal to

length of dorsal valves (Fig. 27C-I0; occurring in southwestern Arkansas in the

West Gulf Coastal Plain region ...... texarkana

16. Genitalia with dorsal valves short and thin (Fig. 15C-I ); occurring in Ouachita

Mountains; genitalia as in Fig. 15C-I ......

...... ouachita

— Dorsal valves longer and thicker, occurring in eastern Ouachita Mountains;

genitalia as in Fig. 19C-I ...... baronei

17. Specimen from Louisiana ...... 18

— Specimen from Louisiana ...... 20

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18. Male genitalia with dorsal valves curved laterally (Figs 1C-F, 12C-F); occurring in

… ...... 19

— Male genitalia with dorsal valves strongly decurved apically, occurring in western

Louisiana; genitalia as in Fig. 27C-I...... texarkana

19. Curved portion of dorsal valves short (Figs 1C-F); widespread species, occurring

mostly through the central (north to south) and southern portion of the state

...... scudderi

— Curved portion dorsal valves longer and apically decurved ventrally; occurring in

Mississippi alluvial plain ...... mississippi

20. Specimen from Texas ...... 21

— Not occurring in Texas...... 27

21. Male genitalia with shortened valves (Fig. 33C-F, 34C-F); longer furcula (Fig. 32

A-B, Fig 34 A-B) ...... 22

— Male genitalia with longer valves (Fig. 20C-F, 27C-F, 29C-F, 31C-F, 33-34C-F);

furcula shorter (Fig. 20A-B, 27A-B, 29A-B, 31A-B) ...... 23

22. Genitalia as in Fig. 34C-F, with dorsal valves curved laterally with apices

projecting posteriorly ...... texensis

— Genitalia as in Fig. 33C-F, with dorsal valves bowed medially with apices curved

dorsally ...... ottei

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23. Genitalia with apices of dorsal valves curved laterally (Figs 29C- F,); ventral valves

often projecting over dorsal valves; occurring in southeast Texas, genitalia as in

Fig. 29C-I ...... tarus

— Genitalia with apices of dorsal valves not curved laterally (Fig. 20C-F, 27C-F,

31C-F, 33C-F) ...... 24

24. Genitalia with apices of dorsal valves curved medially; genitalia as in Fig 20C-I or

21A-F) ......

...... latus

— Genitalia with apices with dorsal valves not curved medially; either curved dorsally

or ventrally (Figs. 27C-F, 31C-F, 33C-F) ...... 25

25. Genitalia with dorsal valves strongly decurved apically, genitalia as in Fig. 27C-I.

...... texarkana

— Genitalia with dorsal valves curved dorsally (Fig. 31C-F, 33C-F)...... 26

26. Genitalia as in Fig. 31C-I/ ...... optimus

— Genitalia as in Fig. 33C-I ...... irwinorum

27. Specimen from Oklahoma/ ...... 28

— Specimen from Arkansas or Oklahoma ...... 33

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28. Dorsal valves bowed laterally and ventrally (Figs. 13C-F, 23C-F, 24C-F) ...... 29

— Dorsal valves not bowed laterally and ventrally (Figs. 15C-F, 18-19C-F) ...... 31

29. Dorsal valves strongly bowed laterally and ventrally; occurring in southeastern

plains of Oklahoma; genitalia as in Fig. 23C-I...... decurvus

— Dorsal valves not a strongly bowed laterally and ventrally (Figs, 23C-F and 24C-F);

occurring in east-central to northeast Oklahoma; genitalia not as in Fig. 23C-F/30

30. Dorsal valves longer, bowed laterally and ventrally with apices recurved dorsally;

ventral valves not dorso-ventrally expanded; occurring north of the Arkansas

River in northeastern Oklahoma; genitalia as in Fig. 24C-I ...... ozarkensis

— Dorsal valves shorter and only slightly bowed laterally and ventrally; ventral

valves dorso-ventrally expanded; occurring in the Arkansas River Valley, south

of the Arkansas River in eastern Oklahoma; genitalia as in Fig 14C-I .. arkansas

31. Genitalia with dorsal valves strongly curved apically and dorsally; occurring in

extreme western Ouachita Mountains; genitalia as in Fig 16C-I ...... cohni

— Genitalia with dorsal valves a most weakly curved apically and dorsally (Fig 15C-

F, 20C-F, 21A-F); occurring in central to western Oklahoma or in the Ouachita

Mountains closer to the Arkansas boarder; genitalia not as in Fig. 16C-F ...... 32

32. Genitalia with dorsal valves, thinner and less robust, occurring in Ouachita

Mountains, close to the Arkansas border; genitalia as in Fig. 15C-I ...... ouachita

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— Genitalia with dorsal valves, thicker and more robust; occurring in central to

western Oklahoma; genitalia as in Figs 20C-I or 21A-F ...... latus

33. Male genitalia with apices of dorsal valves curved laterally (Figs 1C-F, 12C-F) .. 34

— Male genitalia without dorsal valves that curve laterally; genitalia strongly

recurved so that apices of dorsal valves project anteriorly (Figs. 14-16C-I, 26C-

F) ...... 36

34. Dorsal valves not strongly recurved, so that the project anteriorly, but instead

project laterally; genitalia not as in Fig. 1C-F, 12C-F ...... 35

— Dorsal valves so strongly recurved that apicies of dorsal valves project anteriorly;

occurring in the St. Francois Mountains in Missouri; genitalia as in Fig. 26A-F ..

...... francoisensis

35. Curved portion of dorsal valves short (Figs 1C-F); widespread species; genitalia as

in Fig. 1C-F ...... scudderi

— Curved portion dorsal valves longer and apically decurved ventrally; occurring in

Mississippi alluvial plain; genitalia as in Fig. 12C-I/ ...... mississippi

36. Dorsal valves bowed dorsally with apicies recurved dorsally, occurring mostly on

the Ozark Plateau, genitalia as in Fig. 24C-I ...... ozarkensis

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— Doral valves not bowed dorsally, or is so, only slightly and not recurved dorsally;

widespread over the western range of the group; genitalia as in Fig, 20A-F AND

21A-F ...... latus

Key to species of the Melanoplus carnegiei group

1. Cerci triangular ...... 2

— Cerci sub-falcate ...... 4

2. Cerci broadly triangular (Figs 35A-B, 37A-B), tapering evenly from base to apex 3

— Cerci elongate-triangular, tapering from base to middle, the apical half very narrow,

but nearly equal in width throughout (Fig 35A-B); genitalia as in Fig 35C-I

...... acidiocercus

3. Cerci acutely pointed apically, male genitalia with the process of the ramus of the

cingulum well developed giving the appearance of a third pair of valves,

widespread in the extreme southeastern United State; genitalia as in Fig. 35C-F

...... carnegiei

— Cerci bluntly pointed apically, male genital without a well-developed ramus of the

cingulum; genitalia as in Fig. 37C-F ...... chattahoochee

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4. Cerci apicies broadly rounded (Fig 39A-B), genitalia with dorsal valves recurved

such that they point apically; Genitalia as in Fig. 39C-I ...... savannah

— Cerci apicies truncated (Fig 40A-B); genitalia with dorsal valves curved ventrally,

ventral valves subequal in length to dorsal valves and bluntly rounded; genitalia as

in Fig 40C-I ...... carolinensis

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