• TABLE OF CONTENTS
HIDE
 Front Cover
 Title Page
 Division of plant industry technical...
 Table of Contents
 List of Figures
 Foreword
 Acknowledgement
 Introduction
 General account
 Systematic account
 Plates
 References
 Index
 Back Cover








The sand flies (Culicoides) of Florida
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Permanent Link: http://ufdc.ufl.edu/UF00000090/00001
 Material Information
Title: The sand flies (Culicoides) of Florida (Diptera : Ceratopogonidae)
Series Title: Contribution - Bureau of Entomology no. 424
Physical Description: xv, 204 p. : ill. ; 27 cm.
Language: English
Creator: Blanton, Franklin S
Wirth, Willis Wagner ( joint author )
Publisher: Florida Dept. of Agriculture and Consumer Services, Division of Plant Industry
Place of Publication: Gainesville, Fla.
Publication Date: 1979
 Subjects
Subjects / Keywords: Ceratopogonidae -- Classification   ( lcsh )
Insects -- Classification   ( lcsh )
Insects -- Classification -- Florida   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: p. 188-200.
General Note: Includes index.
Statement of Responsibility: by Franklin S. Blanton and Willis W. Wirth.
 Record Information
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: ltqf - AAA0026
notis - ACU1603
alephbibnum - 000520116
oclc - 05303425
lccn - 79624786
Classification:
System ID: UF00000090:00001

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Table of Contents
    Front Cover
        Front Cover 1
        Front Cover 2
    Title Page
        Page i
        Page ii
    Division of plant industry technical council and administrative staff
        Page iii
    Table of Contents
        Page iv
        Page v
    List of Figures
        Page vi
        Page vii
        Page viii
        Page ix
        Page x
    Foreword
        Page xi
        Page xii
        Page xiii
        Page xiv
    Acknowledgement
        Page xv
        Page xvi
    Introduction
        Page 1
        Page 2
        Page 3
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
    General account
        Page 10
        Page 11
        Page 12
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    Systematic account
        Page 38
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    References
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    Index
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        Page 203
        Page 204
    Back Cover
        Page 205
Full Text


ARTHROPODS OF FLORIDA
And Neighboring Land Areas
Volume 10

THE SAND FLIES

(CULICOIDES) OF FLORIDA

(DIPTERA: CERATOPOGONIDAE)
by
Franklin S. Blanton
and
Willis W. Wirth


FLORIDA DEPARTMENT OF AGRICULTURE AND CONSUMER SERVICES
Doyle Conner, Commissioner
___- ~x-~ __^, _^ j_ __ _






























HUME LIBRARY
INSTITUTE OF FOOD AND
AGRICULTURAL SCIENCES

UNIVERSITY OF FLORIDA
Gainesville

W^^^S






ARTHROPODS OF FLORIDA
And Neighboring Land Areas


Volume 10



THE SAND FLIES

(CULICOIDES) OF FLORIDA

(DIPTERA: CERATOPOGONIDAE)



by
Franklin S. Blanton
Emeritus Professor, Department of Entomology and Nematology
University of Florida, Gainesville, Florida 32611

and

Willis W. Wirth
Systematic Entomology Laboratory, IIBIII, USDA,
c/o United States National Museum of Natural History
Washington, D. C. 20560



FLORIDA DEPARTMENT OF AGRICULTURE AND CONSUMER SERVICES
Doyle Conner, Commissioner
DIVISION OF PLANT INDUSTRY
Halwin L. Jones, Director
Florida Department of Agriculture and Consumer Services
Division of Plant Industry
Post Office Box 1269
Gainesville, Florida 32602


Contribution No. 424. bureau of Entomology.



---c l-~2-.-.-


Release Date: June 20,1979


, A...._ .&-... & _-_-L0_0____ ,







FLORIDA DEPARTMENT OF AGRICULTURE
AND CONSUMER SERVICES

DIVISION OF PLANT INDUSTRY



Plant Industry Technical Council
Vernon Conner, Chairman (Citrus) ................ ...................... Mount Dora
Roy Vandegrift, Jr., Vice Chairman (Vegetable) ...................... ...... Canal Point
Lawrence Cutts (Apiary) .......................... ................... C.... Chipley
Colin English, Sr. (Citizen-at-Large) ................ ................... .. Tallahassee
John W. Hornbuckle (Citrus) ............... ............................ Dade City
John Morroni (Commercial Flower) ................ ........................ Ft. Myers
Lewis E. Wadsworth, Jr. (Forestry) . ................... .................... Bunnell
Joseph Welker (Ornamental Horticulture) ................................. Jacksonville
Stanley F. Cruse (Turfgrass) ............... ............................. Palmetto
Halwin L. Jones, Secretary ............... ............................ Gainesville



Administrative Staff
Halwin L. Jones, Director ................ .............................. Gainesville
S. A. Alfieri, Jr., Assistant Director ................ ...................... Gainesville
C. Youtsey, Chief of Budwood Registration ................................ Winter Haven
R. E. Brown, Chief of Methods Development ................ ................ Gainesville
J. K. Condo, Chief of Plant Inspection ................ ................... Gainesville
H. A. Denmark, Chief of Entomology ................ ................... Gainesville
J. Herndon, Chief of Apiary Inspection ................ ................... Gainesville
C. Poucher, Chief of Special Programs ................................... Winter Haven
C. P. Seymour, Chief of Plant Pathology ................ .................. Gainesville
D. E. Stokes, Chief of Nematology ............... ....................... Gainesville




This public document was promulgated at a cost of
$12,243.78 or $8.16 per copy. It makes available to all in-
terested persons the results of arthropod faunal studies
emphasizing Florida and the Circum-Caribbean Region.
PI78S-2







TABLE OF CONTENTS


Title page ......................................................... ...... i

Division of Plant Industry Technical Council & Administrative Staff ................... iii

Table of Contents ............... .......................................... iv

List of Figures ............................................................. vi

Foreword ............ ....... .............................................. xi

Acknowledgements ............... ........................................ xv

Introduction .............................................................. 01

H historical ............ ......................................... ........ 01

Economic Importance .................................................... 02

Disease Transmission .................................................... 03

Control Measures ..................................... ........ ....... 08

General Account .......... ... .................................... ......... 10

Biology .......... ........ ............................ ... ........ 10

Biting Habits .......... .................................... ......... 10

Mating Habits .......... ................................... ......... 11

Adult Food Habits and Autogeny ........................................ 11

Oviposition .......... ...................................... ......... 12

Larval Habitats .......... .................................. ......... 12

Saltmarsh Habitats .................................................. 13

Seasonal Distribution ...................................... ....... 15

Natural Enemies .......... ....-:................. .......... ....... 16

Colonization ................. .......................................... 17

Methods of Collection and Study ................. .......................... 18

Geography and Life Zones of Florida ................. ....................... 21

M orphology ........... ..... ................................... ....... 29

Adult............. ....................................... ....... 29

Immature Stages ..................................................... 34







System atic A account . . . ......................... .................... 38

Classification ........................... ............................... 38

Keys to Species ........................... .......................... 43

Females ........................... .... ....................... 43

Male Genitalia ...................... .................. 46

Pupae .................. ............................ ........... 48

Larvae ..... .......................................... .......... 51

Diagnostic Tables for Quick Identification ...................... ........... 52

Species D descriptions .. ...................... .......................... 54

Plates ................ ........... .................................. 172

References ............ .......... ....................... .......... 188

Index.................................................... ........... 201







LIST OF FIGURES
Fig. Pages
1 Culicoides furens (Poey), the little gray saltmarsh sand fly, adult female. ........... 04
2 Culicoides furens (Poey). a, eggs; b, larva; c, head of larva; d, male genitalia; e, pupa ... 05
3 Shorthorn heifer with acute symptoms of Bluetongue-swollen muzzle and lips, thick
saliva, and swollen, protruding tongue....................................... 06
4 Sheep with Bluetongue. a, typical stance when coronitis is severe; b, severely inflamed
coronary band; c, casting of fleece at 50 days after infection; d, break line where wool
has grown a quarter inch and is less dense. ..................................... 07
5 Breeding site of Culicoides variipennis in Kerr County, Texas, a Bluetongue outbreak
area.................................................................... 08
6 Tide gate at Vero Beach, Florida, effective in limiting Culicoides furens breeding by
changing a tidal saltwater swamp into a freshwater swamp. ...................... 09
7 Mating position of Culicoides melleus. ......................................... 11
8 Mating of Culicoides melleus; separation with spermatophore adhering to female ..... 11
9 Larval habitat of Culicoides loughnani in Opuntia cactus on Sanibel Island, Florida. . 12
10 Larval habitat of Culicoides melleus on sandy shore of tidal water of Indian River at
Fort Pierce, Florida ....................................................... 13
11 Grassy saltmarsh at St. Joseph State Park, Gulf County, Florida, with Juncus
roemerianus; larval habitat of C. mississippiensis and C. bermudensis ............. 13
12 Schematic drawing showing how, in Florida, the same tide intervals can flood the high
marsh continuously at one time of the year and not at all at another. MSHW, mean
spring high water; MHW, mean high water; MNHW, mean neap high water; MSL,
mean sea level ........................................................... 14
13 Intertidal profiles in Florida, with plants characterizing low and high marsh. MSHW,
mean spring high water; MHW, mean high water; MNHW, mean neap high water;
MLW, mean low water; MSLW, mean spring low water. ............. ......... 14
14 Low marsh of Spartina alterniflora at Morehead City, North Carolina, typical larval
habitat of Culicoides hollensis. Cone-shaped emergence trap between D. L. Kline (left)
and J. C. Dukes. ....................................................... 15
15 Red mangrove (Rhizophora mangle) swamp on Sanibel Island, Florida; larval habitat
of Culicoides furens and C. barbosai......................................... 15
16 Pneumatophores in wet soil under black mangrove (Avicennia nitida) in tidal marsh at
Vero Beach, Florida, typical larval habitat of Culicoides furens. ................... 15
17 Culicoides stellifer (Coquillett) male with a parasitic mermithid worm in abdomen. ... 17
18 Bennett trap in operation in Newfoundland: a, initial uncovering of bail fowl; b,
hoisting of uncovered bait to tree canopy; c, final covering of exposed bait preparatory
to collection of biting midges through sleeve cage; d, close-up of bait fowl in restrain-
ing cage on platform board and of sleeve cage. ................................ 19
19 New Jersey type suction light trap effective in surveys of adult Culicoides .......... 20
20 Malaise tent trap designed by H. Townes, useful in collecting adult Culicoides........ 20
21 Outline of the Floridian Plateau and Topographic Divisions of Florida ............. 22
22 Wicomico and pre-Wicomico shore lines in the Southeastern States. ............... 24
23 Penholoway shore line in the Southeastern States...... ...................... 24
24 Talbot and post-Talbot shore lines in the Southeastern States..................... 25
25 Pamlico and pre-Pamlico shore lines in the Southeastern States. .................. 25
26 Small stream in hardwood hammock in Highlands Hammock State Park, Highlands
County, Florida; larval habitat of Culicoides niger, C. scanloni, C. spinosus, and C.
stellifer. ...................................... .................. ...... 26
27 Small branch of the Myakka River in Sarasota Co., Florida, choked with water
hyacinth; larval habitat of Culicoides insignis, C. edeni, C. knowltoni, and C. stellifer. 26
28 Muddy cow pasture at the Ona Range Experiment Station in Hardee County, Florida;
larval habitat of Culcoides insignis, C. knowltoni, and C. pusillus. .................. 26
29 Everglades with sawgrass and cabbage palms in eastern Collier County, Florida; lar-
val habitat of Culcoides edeni, C. floridensis, C. insignis, C. knowltoni and C. stellifer.. 26
30 Early Pleistocene shore lines in the Southeastern States. ........................ 27
31 Adult female of Culcoidds furens, lateral view with left wing and right legs removed,
with parts labeled (cx, coxa; em, epimeron; ep, episternum; m, meron; pn, pronotum;
pp, propleuron; px, precoxale; tr, trochanter). ................ ................. 30







32 a, head and mouthparts of Culicoides female, schematic, parts labeled; b, male
genitalia of Culicoides, schematic, parts labeled. ............... . ...... .. . 31
33 Scanning electronmicrographs of Culicoides: a, b, female maxillary palpus of C. gut-
tipennis showing detail of sensilla in pit; c, same of C. crepuscularis; d, sensillum
coeloconicum on antenna of Culicoides sp. .................................. 32
34 Scanning electronmicrograph of male genitalia of Culicoides guttipennis ............ 33
35 Culicoides pupa: C. arboricola; C. melleus............................... ....... 35
36 Culicoides larva and egg: a-d, g, C. melleus larva; a, head, ventral view; b, head, dorsal
view; c, head, lateral view; d, hypopharynx, ventral view; g, larva, lateral view; e,
diagrammatic reconstruction of skeletal structures of Culicoides head, capsule not
shown except for strip joining dorsal ends of subgenal band; f, lateral view of extruded
anal papillae of C. nubeculosus; h, Culicoides egg. ........................... 37
37 Culicoides alachua: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed ...... 54
38 Florida distribution of Culicoides alachua. ................. ................... 55
39 Culicoides arboricola: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed .............................................................. 56
40 Florida distribution of Culicoides arboricola. ............................. . . 58
41 Culicoides barbosai: a, antenna; b, wing; c, palpus; d, eye separation; e, mesonotum; f,
spermathecae; g, hind femur and tibia; h, male parameres; i, male genitalia, parameres
rem oved................... ............. ............................... 59
42 Florida distribution of Culicoides barbosai ................................... 60
43 Culicoides baueri: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 62
44 Florida distribution of Culicoides baueri. ................................. 63
45 Culicoides beckae: a, antenna; b, palpus; c, wing; d, spermathecae; e, eye separation; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 64
46 Florida distribution of Culicoides beckae....................................... 64
47 Culicoides bermudensis: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed. .. .............................. ..... ...................... 65
48 Florida distribution of Culicoides bermudensis. ........................... .. 67
49 Culicoides bickleyi: a, antenna; b, palpus; c, wing; d, spermathecae; e, male parameres;
f, eye separation; g, hind femur and tibia; h, male genitalia, parameres removed. ..... 68
50 Florida distribution of Culicoides bickleyi. ..................................... 69
51 Culicoides biguttatus: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed. .. ................................. .. ...................... 70
52 Florida distribution of Culicoides biguttatus. ................................... 72
53 Culicoides chiopterus: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
rem oved. .................................. .... .................... 73
54 Florida distribution of Culicoides chiopterus. ................................... 74
55 Culicoides crepuscularis: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed. ...................................... ..................... 76
56 Florida distribution of Culicoides crepuscularis. ............................... 78
57 Culicoides debilipalpis: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed....................... ................. ...................... 79
58 Florida distribution of Culicoides debilipalpis. .................................. 80
59 Culicoides edeni: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 81
60 Florida distribution of Culicoides edeni. ..................................... 82
61 Culicoides floridensis: a, antenna; b, wing; c, palpus; d, eye separation; e, sper-
mathecae; f, hind femur antl tibia; g, male parameres; h, male genitalia, parameres
rem oved.. .......... ............................. ..................... 83
62 Florida distribution of Culicoides floridensis. ............................... 84

vii







63 Culicoides footei: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 85
64 Florida distribution of Culicoides footei...................................... 86
65 Culicoides furens: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 88
66 Florida distribution of Culicoides furens. ................................... 91
67 Culicoides guttipennis: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed. ............................................................... 93
68 Florida distribution of Culicoides guttipennis. .................................. 95
69 Culicoides haematopotus: a, antenna; b, palpus; c, wing; d, eye separation; e, male
parameres; f, spermathecae; g, hind femur and tibia; h, male genitalia, parameres
removed. ............................................................... 96
70 Florida distribution of Culicoides haematopotus................................ 98
71 Culicoides hinmani: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae;
f, hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. .... 99
72 Florida distribution of Culicoides hinmani........ ............................ 101
73 Culicoides hollensis: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae;
f, hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. .... 102
74 Florida distribution of Culicoides hollensis. ................................ 103
75 Culicoides husseyi: a, antenna; b, palpus; c, wing; d, tibial comb; e, eye separation; f,
spermathecae; g, male parameres; h, legs; i, male genitalia, parameres removed....... 105
76 Florida distribution of Culicoides husseyi ..................................... 106
77 Culicoides insignis: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 107
78 Florida distribution of Culicoides insignis. ............. .............. . . 108
79 Culicoides juddi: a, antenna; b, palpus; c, wing; d, spermathecae; e, hind femur and
tibia; f, eye separation ................. .... .............................. 109
80 Florida distribution of Culicoides juddi. ....................................... 110
81 Culicoides knowltoni: a, antenna; b, palpus; c, wing; d, eye separation; e, spermatheca;
f, hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. .... 111
82 Florida distribution of Culicoides knowltoni. .................................. 112
83 Culicoides loisae: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 113
84 Florida distribution of Culicoides loisae. ................ ................... 114
85 Culicoides loughnani: a, antenna; b, wing; c, palpus; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed. ...................................... ...................... 115
86 Florida distribution of Culicoides loughnani. ................................... 116
87 Culicoides melleus: a, antenna; b, wing; c, palpus; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 117
88 Florida distribution of Culicoides melleus. ..................................... 119
89 Culicoides mississippiensis: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
matheca; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
rem oved .............................................. ....... .......... 120
90 Florida distribution of Culicoides mississippiensis. ....... ............... . 122
91 Culicoides mulrennani: a, antenna; b, palpus; c, wing; d, eye separation; e, male
parameres; f, spermathecae; g, hind femur and tibia; h, male genitalia, parameres
removed. .................................. ............................. 123
92 Florida distribution of Culicoides mulrennani. ................................. 124
93 Culicoides nanus: a, antenna; b, palpus; c, wing; d, eye separation; e, male parameres; f,
spermathecae; g, hind femur and tibia; h, male genitalia, parameres removed......... 125
94 Florida distribution of Culicoides nanus. ..................................... 126
95 Culicoides niger: a, antenna; b, palpus; c, wing; d, eye separation; e, male parameres; f,
spermathecae; g, hind femur and tibia; h, male genitalia, parameres removed......... 127
96 Florida distribution of Culicoides niger. ..................................... 129
97 Culicoides ousairani: a, antenna; b, palpus; c, wing; d, eye separation; e, male
parameres; f, spermathecae; g, hind femur and tibia; h, male genitalia, parameres
removed .................................. .................. .......... 130
98 Florida distribution of Culicoides ousairani. ............................... 131







99 Culicoides paraensis: a, antenna; b, wing; c, mesonotum; d, tibial comb; e, sper-
mathecae; f, palpus; g, male parameres; h, male genitalia, parameres removed. ....... 132
100 Florida distribution of Culicoides paraensis................................ 134
101 Culicoides parapiliferus: a, antenna; b, palpus; c, wing; d, eye separation; e, male
parameres; f, spermathecae; g, hind femur and tibia; h, male genitalia, parameres
removed. ................................... ............................ 135
102 Florida distribution of Culicoides parapiliferus. ................................ 136
103 Culicoides pechumani: a, antenna; b, palpus; c, wing; d, spermathecae; e, eye separa-
tion; f, hind femur and tibia................ .... ......................... 137
104 Florida distribution of Culicoides pechumani. ................................ 137
105 Culicoides piliferus: a, antenna; b, palpus; c, wing; d, eye separation; e, male
parameres; f, spermathecae; g, hind femur and tibia; h, male genitalia, parameres
removed. ............................................................... 138
106 Florida distribution of Culicoides piliferus. .................................... 140
107 Culicoides pusillus: a, antenna; b, wing; c, mesonotum; d, tibial comb; e, spermathecae;
f, palpus; g, male parameres; h, male genitalia, parameres removed. ................. 141
108 Florida distribution of Culicoides pusillus ..................................... 142
109 Culicoides scanloni: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae;
f, male parameres; g, hind femur and tibia; h, male genitalia, parameres removed. .... 143
110 Florida distribution of Culicoides scanloni ................. ................. 144
111 Culicoides snowi: a, antenna; b, palpus; c, wing; d, eye separation; e, male parameres; f,
spermathecae; g, hind femur and tibia; h, male genitalia, parameres removed......... 145
112 Florida distribution of Culicoides snowi ....................................... 146
113 Culicoides spinosus: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae;
f, hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. .... 147
114 Florida distribution of Culicoides spinosus ............................. ...... 149
115 Culicoides stellifer: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 150
116 Florida distribution of Culicoides stellifer. ..................................... 152
117 Culicoides testudinalis: a, antenna; b, palpus; c, wing; d, spermathecae; e, eye separa-
tion; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. 153
118 Florida distribution of Culicoides testudinalis ................................. .154
119 Culicoides tissoti: a, antenna; b, palpus; c, wing; d, eye separation; e, male parameres;
f, spermathecae; g, hind femur and tibia; h, male genitalia, parameres removed. ...... 155
120 Florida distribution of Culicoides tissoti ...................................... 156
121 Culicoides torreyae: a, antenna; b, palpus; c, wing; d, tibial comb; e, eye separation; f,
spermathecae; g, male parameres; h, legs; i, male genitalia, parameres removed ....... 157
122 Florida distribution of Culicoides torreyae.. ................................ 158
123 Culicoides travisi: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
hind femur and tibia; g, male parameres; h, male genitalia, parameres removed. ...... 160
124 Florida distribution of Culicoides travisi. ..................................... 161
125 Culicoides variipennis: a, antenna; b, palpus; c, palpus of ssp. australis; d, wing; e, eye
separation; f, spermatheca; g, male parameres; h, hind femur and tibia; i, male
genitalia, parameres removed.......................... .................. 162
126 Florida distribution of Culicoides variipennis. ................................. 165
127 Culicoides venustus: a, antenna; b, palpus; c, pupal respiratory horn; d, wing; e, eye
separation; f, sperniathecae; g, hind femur and tibia; h, male genitalia; i, male
parameres................... ....... .......... ..... ............... 167
128 Florida distribution of Culicoides venustus. ................................... 168
129 Culicoides villosipennis: a, antenna; b, palpus; c, wing; d, eye separation; e, sper-
mathecae; f, hind femur and tibia; g, male parameres; h, male genitalia, parameres
removed ............................................................... 169
130 Florida distribution of Culicoides villosipennis...................... ........... 171







EXPLANATIONS OF PLATES I XII


Plate I, fig. 131-143, respiratory horns of Culicoides pupae: 131, arboricola; 132, barbosai; 133,
bergi; 134, bermudensis; 135, bickleyi; 136, biguttatus; 137, chiopterus; 138, crepuscularis;
139, floridensis; 140, footei; 141, furens; 142, guttipennis; 143, haematopotus.

Plate II, fig. 144-158, respiratory horns of Culicoides pupae: 144, hollensis; 145, insignis; 146,
loisae; 147, nanus; 148, niger; 149, parapiliferus; 150, piliferus; 151, scanloni; 152, spinosus;
153, stellifer; 154, testudinalis; 155, travisi; 156, variipennis; 157, venustus; 158, villosipen-
nis.

Plate III, fig. 159-170, opercula of Culicoides pupae: 159, arboricola; 160, barbosai; 161, bergi;
162, bermudensis; 163, bickleyi; 164, biguttatus; 165, chiopterus; 166, crepuscularis; 167,
floridensis; 168, footei; 169, furens; 170, guttipennis.

Plate IV, fig. 171-182, opercula of Culiciodes pupae: 171, haematopotus; 172, hollensis; 173,
insignis; 174, loisae; 175, nanus; 176, niger; 177, parapiliferus; 178, piliferus; 179, scanloni;
180, spinosus; 181, stellifer; 182, testudinalis.

Plate V, fig. 183-186, opercula; and 187-192, tubercles and spines of Culicoides pupae: 183,
travisi; 184, variipennis; 185, venustus; 186, villosipennis; 187, arboricola; 188, bergi; 189,
bermudensis; 190, bickleyi; 191, biguttatus; 192, crepuscularis.

Plate VI, fig. 193-202, d and 1pm tubercles and spines (except as noted) of Culicoides pupae: 193,
footei; 194, furens; 195, haematopotus; 196, hollensis; 197, loisae; 198, melleus; 199, nanus;
200, parapiliferus; 201, scanloni; 202, spinosus.

Plate VII, fig. 203-207, d and lpm tubercles and spines (except as noted); and 208-213, caudal
segments, of Culicoides pupae: 203, stellifer; 204, testudinalis; 205, travisi; 206, variipen-
nis; 207, venustus; 208, arboricola; 209, barbosai; 210, bickleyi; 211, floridensis; 212, gut-
tipennis; 213, insignis.

Plate VIII, fig 214-225, 226'-227', Culicoides larvae: 214, frontoclypeus; 215-225, head and
thorax; 226'-227', terminal segments and setae. 214, 225, venustus; 215, bickleyi; 216,
chiopterus; 217, crepuscularis; 218, furens; 219, 226', guttipennis; 220, haematopotus; 221,
227', hollensis; 222, melleus; 223, stellifer; 224, variipennis.

Plate IX, fig. 226-235, photographs of wings of female Culicoides: 226, venustus; 227, insignis;
228, alachua; 229, chiopterus; 230, pusillus; 231, hinmani; 232, loughnani; 233, baueri; 234,
haematopotus; 235, footei.

Plate X, fig. 236-245, photographs of wings of female Culicoides: 236, arboricola, 237, gutti-
pennis; 238, ousairani; 239, villosipennis; 240, bickleyi; 241, piliferus; 242, scanloni; 243,
snowi; 244, testudinalis; 245, stellifer.

Plate XI, fig. 246-255, photographs of wings of female Culicoides: 246, furens, 247, barbosai, 248,
debilipalpis; 249, paraensis; 250, biguttatus; 251, loisae; 252, spinosus; 253, melleus; 254,
niger; 255, tissoti.

Plate XII, fig. 256-265, photographs of wings of female Culicoides: 256, nanus; 257, travisi; 258,
crepuscularis; 259, knowltoni; 260, mississippiensis; 261, bermudensis; 262, hollensis; 263,
variipennis variipennis; 264, variipennis australis.







FOREWORD


Except for mosquitoes, probably no other in-
sects in Florida cause more human discomfort
than the tiny biting midges which in Florida
are commonly called sand flies. Especially
along some sea coast areas these little flies,
almost invisible to the unaided eye, can make
life intolerable in early morning, all day on
overcast days, around dusk, and all night on
warm, moonlit nights. So small that they can
easily pass through ordinary 16-mesh wire
screen and mosquito netting, they bring
misery to many a picnicker, bather, fisherman,
hunter, and camper. In addition to the con-
siderable discomfort which they can cause,
many species are intermediate hosts and vec-
tors of filarial worms, protozoans, and viruses
which affect man and domestic animals, as well
as many wild animals. To many people the le-
sions produced by their bites last longer and
are more painful than those of most mos-
quitoes. Some species of Culicoides which oc-
cur about saltmarsh and brackish water areas
are extremely abundant at certain times of the
year and are difficult to control. Localities at
fresh water inlets of the seacoasts and on
streams leading to these inlets are places
where sand flies are most numerous.
Sand fly annoyance is affected by
temperatures and wind. Different species of
sand flies occur during different seasons when
there are different atmospheric temperatures,
but all are most annoying to man when there is
no breeze. Animals having a high body
temperature attract large numbers of the
blood-sucking females. Only female sand flies
take blood, most females requiring this for egg
maturation. In subtropical southern Florida
one may encounter some of them during any
month, although they are generally fewer in
numbers during the comparatively dry winter
months. Some species have 2 generations a
year, while others have only 1.
Sand flies have played a major role in delay-
ing the development of some parts of Florida
as tourist attractions, recreational areas, and
residential areas. Control of the numerous
species depends upon a thorough basic
knowledge of the taxonomy, morphology,
biology, ecology, and the geographic and
seasonal distribution of the various species.
The hematophagous habit of female Culicoides
incriminates them as possible vectors of many
pathogens. An accurate knowledge of the
feeding habits of these flies is an essential part
of the study of their bionomics, since only
those which feed both on the animal reservoirs
of the parasites and on the victim of the


disease can be involved in pathogen transmis-
sion. Drs. Blanton and Wirth present here a
comprehensive treatment of those species
known to occur in Florida. Each species is
described in detail, and keys are presented for
identification of adults, pupae, and larvae oc-
curring in this area, plus diagnostic tables for
quick identification. Key characters for every
species are illustrated, and there is a Florida
distribution map for every species.
Culicoides are recorded from all major land
masses of the world. The genus is large; Wirth,
Ratanaworabhan, and Blanton (1974) recorded
924 valid species, and more recent contribu-
tions of many workers have increased this
number. Wirth (personal communication), in
April 1979, stated that there are 122 species of
Culicoides described from North America and
1,007 from the world; 47 species are known to
occur in Florida.
Dr. Franklin Sylvester Blanton, known to
many of his friends as "Syl," was born in Jones
Mill (now Frisco City), Alabama, on 12
December 1902, son of Franklin H. and Martha
Ada Blanton. In 1942 he married Audrey
Quevedo and sired 2 daughters, Virginia and
Martha, and 1 son, Frank T. Blanton. His
degrees include the Bachelor of Science in
Agriculture, University of Florida (1929), with
a major in entomology and plant pathology;
Master of Science, Cornell University (1941),
with a major in medical entomology and
minors in economic entomology and ecology;
and Doctor of Philosophy, Cornell University
(1951), with a major in medical entomology and
minors in parasitology, ecology, and taxonomy
of Diptera. He began his professional career in
May 1929 with the United States Department
of Agriculture at the laboratory in Orlando,
Florida, as an agent on the Mediterranean
Fruit Fly Control Program. In April 1931 he
was transferred to Babylon, Long Island, New
York, to work on the control of insects and
nematodes affecting ornamentals and
greenhouse plants; he was in charge of the
laboratory from 1936-42. He was drafted into
the United States Army in October 1942, and
in November 1942 was commissioned 1st Lt.
at the Medical Field Service School at Ft. Joe
T. Robinson. During World War II he served 3
tours as Chief Entomologist in the Surgeon
General's office and spent a year (August
1945-June 1946) on duty in New Guinea and
the Philippine Islands and later in Japan and
Korea working on control of scrub and
epidemic typhus. While assigned to the USDA
Laboratory in Orlando during 1948-49, he






spent 2 summers in Alaska field testing insec-
ticides and repellents. During parts of 1949-54
he served as Chief Entomologist in the
Panama Canal Zone and Puerto Rico working
on malaria control and studies on ecology of
jungle yellow fever. He received the USA
Typhus Commission Medal Army Commenda-
tion with 3 oakleaf clusters and other military
citations. He retired from military service in
1956 with the rank of Lieutenant Colonel. He
worked most of 1954 at Cornell University
with Dr. B. V. Travis on punch card abstracts
of medicine and veterinary entomology. In
September 1956 Syl became a full Professor at
the University of Florida, teaching medical en-
tomology, a position which he held until his
retirement 30 June 1974, when he became a
Professor Emeritus of the University of
Florida.
Dr. Blanton is a Fellow in the American
Association for the Advancement of Science,
and during his career he has been a member of
the American Medical Association, En-
tomological Society of America (on original
committee to organize Section D Medical and
Veterinary Medicine), Washington En-
tomological Society, New York Entomological
Society, Canadian Entomological Society, Na-
tional Geographic Society, American Mosquito
Control Association, and The Florida En-
tomological Society. He is or has been a
member of Phi Sigma, Gamma Sigma Delta,
and Sigma Xi honorary societies and Alpha
Gamma Rho social fraternity. He is a Research
Associate of the Florida State Collection of Ar-
thropods and the (United States) National
Museum of Natural History. He is author or
coauthor of 99 publications in the field of en-
tomology and public health (taxonomy of
Diptera of medical importance; virus transmis-
sion; control of insects by chemical; thermal
treatments, fumigation; ecology of yellow
fever mosquitoes). In recent years he has
devoted much of his time to studying the
biting midges of the New World, which has in-
volved extensive collection and preparation of
material from Florida-and Latin American
countries. This study, made in cooperation
with Dr. Willis W. Wirth, is still being con-
tinued and has resulted in a series of publica-
tions and descriptions of many new species of
Ceratopogonidae.
Dr. Willis Wagner Wirth, better known as
"Bill," was born 17 October 1916 in Dunbar,
Nebraska. In 1966 he married Mabel Mercer
Petranek. He has 2 children by a previous mar-
riage, William Frederick and Katherine
Leilani. He pursued a major in biology at Peru
(Nebraska) State Teachers Cdllege (1933-38);
received his Bachelor of Science degree from
Iowa State (College) University (1940) with a


major in zoology and entomology and minors
in botany and chemistry; Master of Science
degree from Louisiana State University (1947)
with a major in entomology and minor in plant
pathology; and Doctor of Philosophy degree
from the University of California at Berkeley
(1950) with a major in systematic entomology
and minors in insect morphology, insect
ecology, limnology, and historical plant
ecology. His military career included the
following in the United States Public Health
Service: 1942-43, 1st Lt., Malaria Control in
War Areas (Louisiana); 1943-45, Aircraft
quarantine and disinsectization (Miami);
1945-46, Captain, Aircraft quarantine
(Honolulu); present, Major, inactive commis-
sion (Scientist Reserve). During 1941-42 he
was Instructor in Biology at Louisiana
Polytechnic Institute. During 1947-49 he was
Teaching Assistant in insect systematics and
forest entomology at the University of Califor-
nia at Berkeley, and during the summers of
1947-48 worked on mosquito surveys for the
California Department of Health, with special
reference to encephalitis virus isolations. From
1949 to date he has served as a Research En-
tomologist in the USDA Systematic En-
tomology Laboratory, located at the (United
States) National Museum of Natural History,
Washington, D. C.
Dr. Wirth is a Life Member and Fellow of the
Entomological Society of America, having
served as Secretary of Section A in 1960 and
on the Editorial Board, Thomas Say Founda-
tion (1960-64), and is or has been a member of
the Entomological Society of Washington,
Pacific Coast Entomological Society,
Hawaiian Entomological Society, Florida En-
tomological Society, Kansas Entomological
Society, American Entomological Society,
Association for Tropical Biology, Biological
Society of Washington, Washington
Biologists' Field Club, and Society for
Systematic Zoology. He is an Adjunct Pro-
fessor of Entomology of both the University of
Florida and the University of Maryland and is
a Research Associate of the Florida State Col-
lection of Arthropods. He is a member of
several national honorary societies: Beta Beta
Beta, Lambda Delta Lambda, Phi Kappa Phi,
and Sigma Xi; in 1947 he received the $100
Williams Prize from the Louisiana State
University Chapter of Sigma Xi. He has receiv-
ed several USDA citations for superior work
performance, and in 1956 he was a Fulbright
Reseach Scholar at the School of Public
Health, University of Sydney, Australia, work-
ing on the biology and taxonomy of Australian
biting midges. He has given numerous invita-
tional papers and participated in symposia at
professional society meetings and served in







special assignments at the University of
Florida in 1951 (studied biology of saltmarsh
sand flies), USDA Laboratory in Kerrville,
Texas in 1953 and 1955 (established the vector
of bluetongue disease of sheep), Museum of
ComparativeZoology in 1961 (acted as consul-
tant to MCZ in rearranging taxonomic collec-
tions of Diptera), New York in 1963 (assisted
Dr. Hugo Jamnback in preparation of an
authoritative manual on the classification and
biology of the New York species of Culicoides),
Smithsonian-Archbold-Bredin Biological
Survey of Dominica (3 months of field work in
spring of 1965 in the course of which the
largest single collection of West Indian
Diptera was collected), Panama in 1967 (a
study of breeding places of Neotropical biting
midges indicated that rotting fruits and other
plant materials are more important than soil
habitats in the rain forest), the Great Plains in
1969 (studied biting midges and brine flies
breeding in saline and alkaline lakes and mar-


she's, particularly in waterfowl refuges), Euro-
pean museums in 1957 and 1973 (studied type
collections of early taxonomists in the impor-
tant collections in London, Paris, Vienna and
Brussels, and several lesser collections in other
cities). Dr. Wirth is author or coauthor of 253
scientific publications, mostly in the fields of
public health and taxonomy of several groups
of Diptera, including Ephydridae, Dolichopodi-
dae, and Ceratopogonidae. He is generally
recognized as the world's foremost authority
on the family Ceratopogonidae.

Howard V. Weems, Jr.
Editor

Bureau of Entomology
Division of Plant Industry
Florida Department of Agriculture and
Consumer Services
29 April 1979







ACKNOWLEDGEMENTS


Our study was initiated as a project of the
University of Florida in 1961 with the
assistance of research and development grants
from the U. S. Army Medical Department. We
have received much additional support from
many individuals and institutions whose help
we acknowledge with sincere thanks.
We were given free access to the Culicoides
collections made by the Florida Department of
Health and Rehabilitative Services since 1948
in connection with its routine light trap
surveillance of mosquito populations. We are
greatly indebted to Elisabeth C. Beck and
John A. Mulrennan for this courtesy and for
their continued interest and assistance.
We have received much help and advice from
John R. Linley of the Florida Medical En-
tomology Laboratory at Vero Beach. Dr.
Linley generously allowed us to use several
photographs. Nina Branch of the Vero Beach
Laboratory furnished us many ceratopogonid
collections including Culicoides over a period
of years.
We are grateful to Andrew J. Rogers and B.
W. Clements of the West Florida Arthropod
Research Laboratory of the Florida Depart-
ment of Health and Rehabilitative Services at
Panama City for habitat photographs and for
advice and information concerning the biology
and control of saltmarsh Culicoides.
As Research Associates of the Florida State
Collection of Arthropods in the Florida Depart-
ment of Agriculture and Consumer Services we
have kindly been given free access to the im-
portant natural habitats found in the Florida
State Parks. For this courtesy we are grateful
to Howard V. Weems, Jr., Curator of the
Florida State Collection of Arthropods, and to
Roy Brooks, Deputy Director of the Division
of Recreation and Parks, Florida Department
of Natural Resources.
The junior author began intensive work on
Florida sand flies in August 1951 during a 4
week visit to the Lantana Field Laboratory of
the Cooperative Sand Fly Research Unit, then
under the direction of R. L. Goulding. Much
help was received from Dr. Goulding, from E.
L. Seabrook of the Palm Beach County Mos-
quito District, and from W. C. McDuffie and J.
C. Keller of the Orlando Laboratory of the U.
S. Department of Agriculture during that
visit.
We are also greatly indebted to Hugo Jamn-
back, Director of Science Service, New York
State Museum, Albany, for his generous loan
of the original plates of figures of larvae and
pupae from his 1965 bulletin. Many of our illus-
trations on plates I-VIII were redrawn from
his plates.
We are grateful to David G. Hall, Arlington,


Virginia, for permission to use his drawings of
Culicoides furens (Poey) for figures 1 and 2. For
use of photographs showing symptoms of
Bluetongue disease in sheep and cattle we are
indebted to John G. Bowne, Location Leader of
the Animal Disease Research Laboratory of
the U. S. Department of Agriculture in
Denver, Colorado. Ellis C. Greiner, Memorial
University of Newfoundland, furnished the
photographs of the Bennett trap operation in
fig. 18. We also wish to thank Wayne A.
Rowley, Iowa State University, Ames, Iowa,
for use of his scanning electronmicrographs
reproduced in fig. 33.
We are grateful to the late Richard Arch-
bold, James N. Layne, and the trustees of the
Archbold Biological Station, Lake Placid,
Florida, for courtesies and facilities extended
during a week's visit in April 1970. We wish to
thank Joe L. Kennedy, acting superintendent
of the Everglades National Park, for permis-
sion to collect Culicoides in the park during
April 1970.
We are indebted to E. V. Komarek, Director,
and Ross H. Arnett, Jr., of the Tall Timbers
Research Station, Tallahassee, for courtesies
and facilities extended during our stay in May
1973. We wish to thank Robert L. Barber,
manager of the J. N. "Ding" Darling National
Wildlife Refuge for assistance in our studies on
Sanibel Island in May 1973. F. C. Craighead of
Naples was also helpful in orienting us in
Everglades and Sanibel Island ecology. We
also acknowledge with thanks the generous
assistance of Dr. and Mrs. William L. Peters,
Florida A. & M. University, Tallahassee, dur-
ing our visit to the Blackwater River
Biological Station in May 1973.
The list of individuals who aided our study
by the collection and submission of specimens
is almost never-ending, but we would like to
acknowledge the following: R. M. Baranowski,
E. C. Beck, W. M. Beck, A. H. Boike, A. Broce,
J. F. Butler, H. A. Denmark, G. B. Fairchild, S.
W. Frost, D. H. Habeck, G. K. Hicks, E. M.
Irons, K. L. Knight, F. W. Mead, A. S. Mills, J.
E. Porter, R. I. Sailer, W. W. Smith, G. M.
Stokes, T. J. Walker, H. V. Weems, Jr., W. H.
Whitcomb, R. E. Woodruff and D. G. Young.
For the preparation of illustrations we are
pleased to acknowledge the assistance of
Gloria Gordon, Molly Griffin, Dorothy
Habeck, Linda Heath and Keiko Moore.
In our Florida distribution lists for each
species we use the following abbreviations to
save space: ERC, Entomological Research
Center personnel, Vero Beach; FSB, F. S. Blan-
ton; LT, light trap; SBH, State Board of
Health personnel*; WWW, W. W. Wirth.


*Now the Florida Department of Health and Rehabilitative Services.

xv


* At






THE SAND FLIES (CULICOIDES) OF FLORIDA

(DIPTERA: CERATOPOGONIDAE)1'2
Franklin S. Blanton3
Emeritus Professor, Department of Entomology and Nematology
University of Florida, Gainesville, Florida 32611
and
Willis W. Wirth3.4
Systematic Entomology Laboratory, IIBIII, USDA,
c/o United States National Museum of Natural History
Washington, D. C. 20560


INTRODUCTION


HISTORICAL

The people of Florida have always contended
with a serious sand fly problem because of the
long coastline, extensive beaches, and tidal
waterways. Accounts of early explorers and
settlers speak of the sand fly pest. Bernard
Romans (1775:152) refers to the "uncommon
swarms of flies, gnats, and other insects,
which attend putrid air, and unhealthy places
covered with wood;... gnats, here called mus-
quitos, are vastly numerous in some spots, but
after we have passed some distance the
brackish waters in every river, they diminish,
and at last we find none;... and on Dolphin-
island, likewise in Santa-Rosa bay, and in the
bays of St. Andrews, St. Josephs, and at St.
George's sound and islands, they are in-
tollerable, and all the sea coast of both pro-
vinces is exceedingly pestered with them. I
travelled across the peninsula in 1769, in the
months of June, July, August, and September,
but found none at all in the interior part; I
never heard these being attended by any fatal
disease except their troublesome bites, which
sometimes cause inflammations, specially in
the legs; but the inhabitants of the western
province so effectualy fortify themselves

This investigation was supported in part by
United States Army Medical Department Con-
tract No. DA-49-193-MD-2177. Approved as
Florida Agricultural Experiment Station Journal
Series No. 900.
2Contribution No. 424, Bureau of Entomology,
Division of Plant Industry, Florida Department
of Agriculture and Consumer Services.
3 Research Associates, Florida State Collection of
Arthropods, Division of Plant Industry, Florida
Department of Agriculture and Consumer Ser-
vices.
4 Adjunct Professor, Department of Entomology
and Nematology, University of Florida.


against those vermin with musqueto nets,
tents, and Baires, that whether at home or
traveling, they are not in the least danger from
the attacks of this terrible and bold, though
diminutive enemy, whose destruction gradual-
ly takes place as the woods daily diminish; in
very dry hot summers, scarce any of these ver-
min are seen; a very dry hot air causing the
deaths of numberless animaculae of every
kind, their effluvia, even of those that are im-
perceptible to the naked eye, arising or exhal-
ing from ponds, marshes, swamps, &c. must
spread a great quantity of noxious vapours
through the atmosphere, and consequently cor-
rupt the air, and spread disease throughout
their vicinity; this misfortune will likewise
cease on opening the country, till then let me
advise every new comer, particularly a person
of a gross habit of body, to be careful of his
constitution, a wine-bibber, or rum guzzler,
with such a plethoric habit, can hardly avoid
falling a prey to this bad air."
Williams (1837:71), in his account of the
natural history of Florida, listed 4 kinds of
gnats (Culex): 1) the gallinipper; 2)
mosquitoo"; 3) "Sand Fly; this insect con-
tinues but a short period, and is confined to the
coast"; and 4) "the Mite Gnat, is common to
every part of America, and we have our share
of them." His third kind, the sand fly, no doubt
refers to one of the saltmarsh sand flies of the
genus Culicoides, and the mite gnat could also
possibly refer to this genus.
Woods (1893) recounted that during a march
down the Florida East Coast the Confederate
soldiers were forced to walk along the beaches
and, when resting, to cover their bodies in the
sand and use clothing to protect their heads.
Swanton (1922), in a history of the Creek In-
dians, reported that they placed smudges
beneath their beds to repel the biting midges.
Dove et al. (1932) were of the opinion that sand
flies and mosquitoes were largely responsible






for the lack of early development of the coastal
areas of the South Atlantic States. They re-
counted an instance in which an escaped con-
vict successfully evaded the police but had to
give himself up because he could not bear the
sand fly bites.
The 1st studies of the biology of the most
pestiferous of the American coastal species of
Culicoides and of possible methods of control
were in 1932 at Fort Pierce by the United
States Department of Agriculture and the St.
Lucie County Sanitary District. These and
later studies resulted in the recommendation
that the saltmarshes be diked for water man-
agement to control the sand flies (Hull et al.,
1934, 1939, 1943; Platts et al., 1943).
Shortly after World War II, a Cooperative
Sand Fly Research Unit was set up at Fort
Lauderdale, Florida, under the joint sponsor-
ship of the United States Department of
Agriculture (supported financially by the
Department of Defense), the Florida State
Board of Health, and the mosquito abatement
districts of Dade, Broward, Palm Beach, and
Indian River counties. This laboratory made
considerable progress in defining the larval
habitats of Culicoides furens (Poey), C. bar-
bosai Wirth and Blanton, and C. melleus (Co-
quillett), species found on the lower Florida
East Coast, and in testing the efficiency of in-
secticides as control agents. As a result Curran
and Goulding (1950) reported methods of
sampling the larvae, Wirth (1952b) described
the immature stages of C. furens and C.
melleus, and Goulding et al. (1953) and Labrec-
que and Goulding (1954) discussed insecticides
that could be used to control adults and larvae.
In 1954 the Florida State Board of Health
established the Entomological Research
Center at Vero Beach where extensive studies
were made of the biology and control of sand
flies. Since 1967 J. R. Linley has carried out ex-
tensive studies of Culicoides biology and
physiology at Vero Beach. In 1965 the control
methodology research activity was transferred
to a new West Florida Arthropod Research
Laboratory in Panama City where A. J. Rogers
and B. W. Clements continued research on
biology and control of saltmarsh sand flies.
In Alachua County W. W. Smith of the
University of Florida, and several students,
conducted studies of the biology of Culicoides
from 1964 to 1969, principally with the species
breeding in tree holes.
According to Edwards et al., (1939), the 1st
historical account of the life history of a
Culicoides was given in 1713 by Derham (in
Edwards et al., 1939:26) in England: "For an
Instance of Insects endued *ith a Spear, I
shall, for its peculiarity, pitch upon one of the
smallest, if not the very smallest of all the


Gnat-kind, which I call Culex minimus
nigricans maculatus sanguisuga. Among us in
Essex they are called Nidiots, by Mouffet
Midges. It is about '/o of an inch, or somewhat
more long with short antennae, plain in the
female, in the male feather'd, somewhat like a
Bottle-brush. It is spotted with blackish spots,
especially on the wings, which extend a little
beyond the Body. It comes from a little slender
Eel-like Worm, of a dirty white Colour, swim-
ming in stagnating Waters by a wriggling Mo-
tion, as in fig. 5.
"Its Aurelia is small, with a black Head, lit-
tle short Horns, a spotted, slender rough Belly
Vid. Fig. 6. It lies quietly on the top of the
Water, now and then gently wagging itself this
way, and that.
"These Gnats are greedy Blood-suckers, and
very troublesome where numerous, as they are
in some places near the Thames, particularly in
the Breach-waters that have lately befallen
near us, in the Parish of Dagenham; where I
found them so vexatious, that I was glad to get
out of those Marshes. Yea, I have seen Horses
so stung with them, that they havJ had drops
of blood all over their Bodies, where they were
wounded by them."
Edwards et al. (1939) then pointed out that
"Exactly two hundred years elapsed before a
more precise description than that of Derham
was published of the larva and pupa of any
species of Culicoides. In the year 1913 Patton
published an illustrated account of the life-
history of an Indian species, and at the same
time Lutz published some account of his obser-
vations on Culicoides in Brazil."


ECONOMIC IMPORTANCE

The tremendous numbers of Culicoides
midges that are present in coastal areas can
make life almost unbearable in some places by
their annoying attacks. These small gnats are
called many names in many places, some of
them unprintable: punkiess" and "no-see-
ums" in the United States and Canada,
"moose-flies" in Alaska, "sand flies" on the
Atlantic and Gulf coasts of the United States
and in the West Indies, "jejenes" in Latin
America, "maruins" in Brazil, and "no-nos" in
Polynesia.
Reye (1964) wrote most aptly: "Most people
know the Ceratopogonid midges as the 'sand-
flies' met with on the seaside holiday or fishing
trip. The abrupt fall in morale and the strong
desire to be elsewhere which they engender, are
difficult to convey to those who have not ex-
perienced them; as Kettle (1962) puts it briefly
'one midge is an entomological curiosity, a
thousand can be hell'. ".







The tourist business is greatly affected by
sand fly problems along many Florida beaches
where there are significant populations of
Culicoides furens (fig. 1, 2), C. barbosai, C.
hollensis (Melander and Brues), C. melleus, and
C. mississippiensis Hoffman. In some inland
localities, particularly in the "Highland" and
"Crystal Springs" counties, C. floridensis
Beck and C. tissoti Wirth and Blanton may
become a nuisance. Linley and Davies (1971)
presented an excellent review of the problem of
sand flies versus tourism, a summary of the
biology of the most important species, and
recommendations for control. The main thrust
of their article, however, is directed toward
avoiding a sand fly problem rather than trying
to control a problem that already exists. For
example, they "advise that intending
developers should fully investigate the sand fly
situation before commencing their projects,
since control measures forced upon them later
may be very expensive."
There is a growing literature on the effects of
Culicoides bites on livestock. Allergic der-
matitis of horses has been reported as
"Queensland itch" in Australia (Riek, 1954)
and as "Kasen" in Japan (Yamashita et al.,
1957). McMullan (1971) reported a dermatitis
of the withers, tail, mane, and ears of horses in
Texas due to Culicoides bites, and recommend-
ed stabling at night for control.


DISEASE TRANSMISSION
Although all blood-sucking arthropods in-
cluding Culicoides must be suspect as poten-
tial vectors of disease, experimental work with
ceratopogonids as disease vectors has been
hindered and delayed because the insects are
so small and so difficult to colonize. Never-
theless, discoveries in recent years are increas-
ingly incriminating Culicoides as vectors of
pathogens of man and of wild and domestic
animals.

PROTOZOA
Fallis and Wood (1957), the first to in-
criminate Culicoides as vectors of protozoan
blood parasites of birds, discovered that
Haemoproteus nettionis (Johnson and
Cleland), a malaria parasite of ducks, would
develop in an unidentified species of
Culicoides. Fallis and Bennett (1960) studied
the sporogeny of Haemoproteus canachites
Fallis and Bennett, a parasite of the spruce
grouse, in C. sphagnumensis Williams in On-
tario. Bennett and Fallis (1960) and Bennett
(1960) found that heavy parasitaemia of
Haemoproteus spp. in immature birds was
preceded by peak abundance of Culicoides spp.


Bennett (1961) noted developing stages of
trypanosomes in Culicoides fed on infected
birds but did not consider this insect to be the
important vector in Ontario. Akiba et al. (1959)
and Akiba (1960) studied the transmission of
Leucocytozoon caulleryi Mathis and Leger, an
important pathogenic parasite of poultry in
Japan, by C. arakawai (Arakawa).
Garnham et al. (1961) worked with
Hepatocystis kochi (Laveran), a Plasmodium-
like malaria parasite of Cercopithecus
monkeys in Kenya. They traced its develop-
ment in C. adersi Ingram and Macfie, where
the sporocyst (= oocyte) lies free in the
haemocoel and is not attached to the gut wall
of the insect as with most malaria parasites.

FILARIAL WORMS
Sharp (1927) reported that Culicoides
grahami Austen could transmit Acan-
thocheilonema perstans Manson, a non-
pathogenic parasite of man in Africa, and in
1928 he showed that C. austeni Carter, In-
gram, and Macfie also was a vector. Henrard
and Peel (1949) and Chardrome and Peel (1949)
believed that Sharp had confused his parasite
with Dipetalonema streptocerca Macfie and
Corson, whose microfilariae occur in the skin
(those of A. perstans are found in circulating
blood); they obtained a 23% experimental in-
fection rate with D. streptocerca in C. grahami,
but could not get this vector to ingest A.
perstans. Subsequently Hopkins and Nicholas
(1952) obtained a 41% infection of reared C.
austeni with A. perstans. Also they found that
C. grahami collected in the field ingested fewer
A. perstans microfilariae than would be ex-
pected from their density in circulating blood,
but that C. austeni ingested more than ex-
pected. Duke (1954) confirmed the results with
C. grahami and showed that this species also
could be infected easily with D. streptocerca,
however he later (1956) found that many more
larvae of A. perstans reached the infective
stage in C. inornatipennis Carter, Ingram, and
Macfie than in C. grahami.
Steward (1933) was able to infect C. ob-
soletus (Meigen) and C. nubeculosus (Meigen)
with Onchocerca reticulata Diesing (= cer-
vicalis Raillet and Henry), a filaria that causes
fistulous withers in horses in Europe. When he
followed the entire development of the filaria in
C. nubeculosus, he found that the cycle took 25
days from ingestion of the microfilaria until
the infective stages were ready to leave the
head of the fly. Moignoux (1952) reported
transmission with C. nubeculosus in France.
McMullan (1972) reported cases of onchocercal
filariasis in Texas horses with symptoms
characterized by recurrent seasonal pruritis,









































































Fig. 1. CulicQides furens (Poey), the little gray saltmarsh sand fly, adult female (from Dove, Hall, and Hull, 1932).
Cover illustration.














-C





















/






D E





Fig. 2. Culicoides furens (Poey). a, eggs; b, larva; c, head of larva; d, male genitalia; e, pupa (from Dove, Hall, and
Hull, 1932).








mullms, chest,
i. i dditi.. to -dic. to kill
,he Mrofillrias he recommended stabling the
the = ,no gotctin i.'nt C.U-Id..r bit..
during yths.
Dkpf (1938" "'l Wd th, dppoiblp t of
mic-fffmao of what he believed to be 0,
eho. -o.i Ii ..d O -1-i- Louckardt
f.o.g. Hoffman in Mexico. However,
(195 Gibs- a 1d A-11 (1952) attempted ax
,fierietally to infect the four principal au-
ook.p,, hill. Colpoido spade. of the o.-
ch ...... .one of the Guatemala highlands with
0.ululs their results war.ngtie They
... eluded that ru.7oft-d. probably we, at in-
=Ivd nnaua tr-roaissa., which reur
-JaLly hyhle of Si-uliid-e Presumably
pso-how,, dealing with o th flra
BuckLley (1938) studied the possible vectors
of 0-h.-erc gibs-ni (Cleland and Johnston)
in cattle in Malaya. This worm causes ag
-odul. to ban, on the brisker ud ffi. ga
cattle. Of 20 species of CallrafdaE -11.vtZ c.






'path, chuan ii" .... orll oladi
Manson, He found that infection rawe of C
f,,,n collected in the field mrs nexesed it=m







iu a wooded regioo whr C pom ... i breM,
f. t..e holes and is aot found a, the treeless

Robins- 'IMP5) f ...d !C'redC




infected starting


fnoa hl and And rso, of Taiwan
= kayays Developm at of the mlcrofilariae



the meekey,


u j.











16th and 1 ttbaa, (1956) P apo~fed that U
. P. U Coqrpllett was the commaPp st
and most widespread biting midga the














































them into 2 h..ep that lat., developed elitti-I



tmieimg -rpl e h lny of C.apmpdiF
F-tver et 1 (11963) -vr able to Vchisve
bielogmeal transmission ,p,,Im,,ntally under
.t-gsget laboratory pr...Ios. Let., Jusima
.pd Joasp (1966) d.--trllpd eslliplipppi..











ar aspPtsdI ectom, but tee equard in
"'.iinti'g "giresmetttion has net koes
do...


E-plehlitis. K.,stad et .1. (195r7) isalabsd
eastern equine smoephalitle virus from an
,ed-mmied Ckft-d- -r soeth... Georgia.
They Is. tspttrtd toress-i-ti.. from
Lavi-C-still, to the affect that Venezuslan
qpin. troophahDa had been i-Ilted frma
C.Ureoids. i E-adar. We sed W. (1957)
recovered the virus of Japanese B encephalitis
f-es F-rip-myi. Lroro tnii...
(Shimaki) in China (L,,iohelea is another irreu



asperiment.Bly by usmg C. _taup s- .l







south Texas in 1971. They considered C. CONTROL MEASURES
or a prime suspect forthe biological Becoms te annoying f emale ieid may
transmission of the vrus from horse to horse ,.
in that area and along he Central bAm-.an "
oasts. temporary and must be repeated as new
Mivallaeh-us Arbovirses. Reeves et al females emerge and movements occur.




970 found m tha Crohlnais trans mitso a f i
arbIvirus in Californa which they med d
RButtonwllow." Jackrabblts and cottontail


reported that 2 additional viruses, Loka- and .. .... .... ..
Main Drain, e transmitted als toa the same ,- . '
hosts by C. aHipns, in California.
iAt Helerau, Bra Pinbelro et al. (1976 but ldo inexpensive treatmei
isolated OrpouV h iru in 2 out of 40 pools ?
totaling about 15,000 Cuhcotdes, of which 96%
were C para-nsis {Graldi), Oropouche virus
caused 5 epidemics among humans in the


-h,, ',', .. ,.. '.
. tuachs, my ,lgi, ,rthralgi., ad di.... a



to the vir-s, idicating that they may serve as
reservoir Iots.
The Arhovirus Resenreh L.aboratory at the
University of Ibedan, Nigeria, reported (1967,
unpublished) that 377 pools totaling 103,500
Culieotdes o., ected in November were in-
oculated for virus isolation attempts, and 44
viral agents were isolated. One agent was a
Simbu group virus, one a strain of Congo virus
that is also found in ticks and cattle, 2 were However, in pest control, the more effective
identical with AR1792 found in ticks and cat- the pestiide, and the higher proportion of the
Ile, and 37 were identified as a distinct virus" C. I .
(AR22888) created to the hletongue group.
Doberty t al. (19721 isolated 3 arboviruses

remains that cannot be chemically eliminated,
herds of cattle affected by ephemeral fever No
ephemeral favor virus strains we.. isolated I; ",.
from mosquito or biting midge collections .. .. ".
Prestwood et al. (19741 reported hemorrhagic
disease (HD) caused by bluetongue virus -,j.h r i , '



pre,,d in South C arohana and spread ,almost .


from peracute, acute, to chronic, and mortality 7


the EHDV is unknown









t-Chtt~A


(1969), ,,rking in Jamaica. margroTe





reduced the ta,,mlni... of C. b-b-osi to
10th of that in the shaded area. but had little
effect on C fin'_ and C' -sig-.s
-Di.. Najera (1970) sa.-i..ed the p.o
do ... ...d m the cration of ,program for
the enme.1l of mosquitoes and C.Ii-ode asn



of aintauia 'Roo, and the Hotel Owaaas'




3 1








,.ospouainsfsn fyI, u h
uslimi .-i
rt a aanoedbeasei
ranypemisbypupig
Blno t. 15)ama redcini


.3








the dry -imso




..d to control breeding of' c faruse is
magaesap.The great reduction in





On SambO Warlad estmn of dikea -it
canals was doeised to separate and protect a
zsLt-tc, sanctuary and !raaebwwatar sima-





Mi. '






GENERAL ACCOUNT


BIOLOGY


BITING HABITS
Snow (1955) compared the biting activity of
Culicoides on the ground and in the tree
canopy during the day and night in a bot-
tomland forest in western Tennessee. Pickard
and Snow (1955) reported 8 species of
Culicoides biting man in the Tennessee Valley;
also 2 species were found feeding on horses.
Snow et al. (1957) summarized the feeding
habits of Culicoides in the Tennessee River
Valley.
Judd (1959) found engorged C. haemato-
potus and C. biguttatus (Coquillett) in a cat-
bird nest in Ontario. Ryckman (1961) found
engorged C. ryckmani Wirth and Hubert in a
nest of a house finch (Carpodacus) in a cactusin
the desert of southern California; he also
recorded Leptoconops herteszi Kieffer and C.
variipennis feeding in ears of jackrabbits.
Jones (1961a) exposed sheep to attacks of
biting midges in Colorado and took many C.
variipennis. He also took smaller numbers of C.
hieroglyphics Malloch, C. haematopotus, and
C. stellifer (Coquillett) and L. kerteszi.
Messersmith (1965) operated a New Jersey
light trap in poultry houses in Virginia where
an outbreak of infectious synovitis had occur-
red. He took significant numbers of engorged
females of 6 Culicoides species. C arboricola
Root and Hoffman, C. crepuscularis, and C.
villosipennis Root and Hoffman were most
abundant and the most suspect potential vec-
tors.
Jamnback (1965) gave biting records for
New York Culicoides. Of the 37 New York
species, 17 were taken on man, 10 on birds, 6
on mammals other than man, and 16 with no
records.
Atchley (1967) summarized biting records of
New Mexico Culicoides and gave records for C.
obsoletus and C reeves Wirth biting man in
that state.
Hair and Turner (1968) reported host
preferences of 15 Culicoides species in Virginia
when given a choice of 10 caged birds and
mammals, and human bait. All fed to some ex-
tent on man, but C. arboricola, C.
crepuscularis, and C. travisi Vargas were
decidedly ornithophilic; C furens, C. guttipen-
nis (Coquillett), C. hinmani Khalaf, C. melleus,
C. sanguisuga (Coquillett), C. stellifer, and C.
variipennis preferred mammalian hosts.
Likewise Humphreys and Turner (1973) in-
vestigated the host preference of 12 species of


Culicoides in Virginia. They found only 2
species, C. guttipennis and C. sanguisuga, in
sufficient numbers for statistical analysis of
environmental and host factors. Size of host
was a factor in attracting these species, but
color of host was not. Optimum conditions for
adult feeding activity of C. sanguisuga were
16-18 C and 85-95% R.H. Jorgenson (1969)
reported on feeding precipitin tests of 3 of the
13 Culicoides species found in southeastern
Washington: C. varipennis fed heavily on
swine and also on cattle and horses; C. freebor-
ni Wirth and Blanton fed on these same hosts
plus dogs; and C neomontanus Wirth and
Blanton fed on cattle. In Denmark, when
Neilson (1971) collected biting midges on
various body parts of grazing cattle, he found
that C obsoletus, C. punctatus (Meigen), and
C. subfascipennis Kieffer preferred the belly
region, C. obsoletus and C. punctatus fed on
the back and flanks, and C. chiopterus (Meigen)
fed on the legs. Anderson (1964, personal com-
munication) studied the biting midges found
inears of deer and jackrabbits in Mendocino
County, California. In jackrabbits he repeated-
ly found C. utahensis Fox, C. tenuistylus
Wirth, C. neomontanus, and C. variipennis. In
deer he found many C. utahensis and C.
tenuistylus and fewer C. freeborni and Lep-
toconops kerteszi.
Some Culicoides species are known to prefer
to bite during the day, even in bright sunshine,
but other species prefer dusk or nighttime
feeding periods. Snow (1955) and Snow et al.
(1958) made detailed observations of the time
of feeding of Culicoides in Tennessee and
South Carolina. Also, Humphreys and Turner
(1971) tested the effect of various light inten-
sities on the feeding habits of laboratory-
reared C. guttipennis in Virginia, and deter-
mined that the greatest feeding activity occur-
red at light intensities between 0.1 and 9.0 fc
(bright light = 19.0 fc).
Such studies of the feeding preference of
Culicoides have been greatly hindered because
the females of most species are reluctant to
enter any kind of enclosure or cage to attack
their host. Indeed, advantage is taken of this
habit in South Africa where sheep are penned
in a fold under a roof at night to prevent
Culicoides attacks and control bluetongue (R.
A. Alexander, personal communication). Ap-
parently Culicoides are very sensitive to their
light horizon for orientation so bait traps of the
type normally used for mosquitoes (wire or
gauze cages with entrance baffles) have not
been of much value. However, Bennett (1960)

0







in Ontario developed an ingenious drop trap
that could be used at various elevations, even
.... -h ..... ,., -,.,




canopy in a southwestern Virginia locality.
They found that height above ground level and
sie of the host animal often werea important
factors m the host preference of some species,
and were perhaps more 1ror than the faet
that the host was a bird or a mammal.


MATING HABITS




unusual species mate without swarming: both ::
sexes .n or crawl on the soil or vegtabton




i western North Amrica) Most peiep e mate
only once, but Jones (1966b) found that C
_Pn"'i' ... .ma rep ately and that


more than once. Sperm is transferred to the
female inp asp ratophore in C nubpeculosus







th -,- ..,.I l- f th. ,, ta.k bl.. ..d..I.I


.. ... ..p-.. is i..d ft.-. .PPpPP






time. They are deposited an i n ist substrate




pdbpeh.
evaria, cycle, ,old be ,,,ert,!,,d by the


batches of eggs. James (1967) f ...d that C.
than 1 ish," body difficult to obse- 1,i C ... iip-i females would deposit so egg
f__ ad C brb-L Some species with
reduced me hpart, 1d- number of -i -
dibf~d. th) a e or it-. to take blood I- ~.~;



females = ligt attempting to bit, in mt-,-e
Florida A Pat...rou, i..., had relict rrol ,
btions f t b previous opposition. and con








emerging in Match situr developing during the


b,-p d,,, had no mile' and reproduced pard
thriogenitslly Females were gravid whso
they emerged from pupae and produced at
least I batch of mible egg, mt act a blood

meproduc s an the American mainland where
mal.. of the species ..our. Lar (1968) observed
pa-th .. g,.Tsi, and antogemy in C b-m

i.g the ataei.. th-gh av- population
depletion.




ovipoit' but decapitation can remove
hibiti.. (Lmayy 1965,), -nd agga will be
rpoaited freely for -poerimental studies.






debri, the bottom of Us, catch wr rmt hole
(gtppppp g-pppp C p-pp, Wirth and
p-7 ,,,i Wirth ad Jones, C fh P p


a ... s) Many speciess b ...d am the sh...s of


.L pr isppdo FredWh ", P96') r.'
exist at dopth of _'se the. a fmw in hea
belmw the air-ater Interface. 1, tropical main
f-xets, lagept oorc o of species may




pphppppp, ,pd
uncepeteretm a c-lpetit-o o to -mdi
'I "ctsol coding (Wirth at 0l., 1968).
Suitable food f pr the ,p,ophagoas and


may fced el small pcy p pismp such as


-lely to fall-w th. k-tcte intaffac. and rn




1 5-54 of Cut, p.. h e larval habitats in the


Tp ... d 1:,,l habitats of 9 species of Imma
Pcoides d an .. t the pH, sodium It.
conhenhratiopP and organic after P t '
ths habitats. 7 spaciws -er found in ..asheda
habitats, 5 in hatb..u...-mud habitat., 4 1n
harbaccoms-sand habitats, and 4 in alkaline
Zt he,,, t".
Smith (19660l found slightly rcid pH
(6.5-7.0) favorable f I-asa of several soil-



L- c -m, f .. d 1, Us, hole, h-er the pH



,,lu ths. ?H we th-a8.7, and grea I to









'.ctmx o the special fthportanc. of


Floda sPnd flies, a more detaed revw
should be rode of the characteristics of th.se
habitats in Florida. For this account -e have
drawn fely from the public ations of Davis
.. , ,,i ,i : -i ,








.P~p~ppppppd p .ppppppp
studed refully in order to appreciate the
complexity of intertidal habitats and the dif-
ficulty of characperppng them on physical fac-
tors alone. Provost concluded that tp e key op


m a species alysis of the vegetation.
Ocean tides am governed by many factors,
but the most important are the phase of the
moon, distance of the moun from the earth. and
declination of the moon with respect to the
each's equator. When the moon is in full or
n- phase its U'avitatlonal pull is augmented
by that of the aun resulting in maximum pull



















NEAP SPRING
T16ES


Fig. 12. Schematic drawing showing how, in Florida, the
same tide intervals can flood the high marsh continuously
at one time of the year and not at all at another. MSHW,
mean spring high water; MHW, mean high water;
MNHW, mean neap high water; MSL, mean sea level
(from Provost, 1973).


on the ocean and spring tides with large tide in-
tervals (fig. 12). The opposite or neap tides oc-
cur on either of the quarter moons. When the
moon is closest to the earth at perigee, it
creates maximum pull and extra large or
perigean tides, and when it is farthest from the
earth, its pull is minimal with extra small or
apogean tides. When the moon's axis is at its
greatest declination from the earth's equator
(230 30' north or south) it exerts maximum pull
creating large tide intervals called tropic tides,
and when it is in line with the earth's equator,
it creates minimal or equatorial tides. These
factors come into coincidence with extreme
maximum or minimum tides according to a
definite astronomical cycle with a predictable
schedule of other peaks and minimums in-
terspersed.
Tide intervals (amplitude or vertical distance
from low to high tide of any 1 cycle) also vary
geographically. In Florida the mean tide inter-
val may be as much as 6 feet on the Atlantic
coast near the Georgia state line to as little as
1 foot on the lower Florida Keys or in extreme
western Florida.
An additional factor is the seasonal change
in mean sea level (fig. 13). In Florida the
monthly mean sea level rises to more than a
foot above the annual mean for about 3 months
from August through October on the Atlantic
coast, but on the Gulf coast it rises much
earlier (in May) and lasts half the year.
Florida's tidelands usually co'itain 2 mean
high water marks (fig. 13). The outer mark
separates the low marsh, flooded by
tidesalmost daily, from the high marsh, flood-
ed only by the higher tides. The high marsh in
Florida has a unique flooding pattern whereby


for several months from December to May it
can remain unflooded by even most spring
tides, while during another time of the year
(from May or September to November) it may
remain continuously flooded because even
neap tides reach the mean high water level.
Thus the amplitude, the pattern of daily
flooding, and the pattern of flooding over
monthly or annual cycles have almost infinite
variations depending on the geographic loca-
tion of the marsh. Tidewater plants have
evolved in response to these patterns over
many years so that the presence of some of
these plants can very conveniently be used as
indicators of a certain type of marsh with a cer-
tain tidal pattern and a corresponding biota.
In Florida the low marsh usually is
characterized by extensive growth of smooth
cordgrass (Spartina alterniflora Loiseleur) in
the northern counties of the Atlantic coast,
while black needle rush (Juncus roemeriana
Scheele) is dominant on the northern parts of
the Gulf coast. The high marsh is characterized
by salt meadow hay (Spartina patens (Aiton)
Muhlenberg), spike grass (Distichlis spicata
(Linnaeus) Greene), etc. (Cooper, 1974).
In coastal North Carolina Kline and Axtell
(1977) studied the distribution of the 3
Culicoides species, C. hollensis, C. furens, and
C. bermudensis, in relation to plant cover.
They found a correlation between the species of
plants, as well as the midges, and the percen-
tage of time a marsh is flooded and the
characteristics of the soil. Tide-elevation rela-
tionships affected many soil physical and
chemical properties, which in turn influenced
plant growth. C. hollensis was most abundant
in tall growth of S. alterniflora that was
flooded an average of 19-36% of the time. C.



S -MNHW






MSHW--
M" m,,_,- __ MHW






MLW LOW4 IH- MARSH O-MNHW
MS, /7 "- pud" b" *
I _


I - -V ----- ---- ----



MSLW.o.. .
MOSTL.Y CORDGRASS (Spalna alternl oraT ) SALT MEADOW Hn A (Sit rtiinap n
OFTEN BLACK RUSH (Juscus rSofiana) SPIKE ORASS (DfiicNIs avicata)



G RASS SALT MARSH Ei T

MANGROVE SWAMP




Fig. 13. Intertidal profiles in Florida, with plants
characterizing low and high marsh. MSHW, mean spring
high water; MHW, mean high water; MNHW, mean neap



high water; MLW, mean low water; MSLW, mean spring
low MANter (from Provost, 1973).CK MANROVE (A
SALTWORT (Sati maritime)
GLASSWOfTS (Saleornda we.)
ETC.
MANGROVE SWAMP

Fig. 13. Intertidal profiles in Florida, with plants
characterizing low and high marsh. MSHW, mean spring
high water; MHW, mean high water; MNHW, mean neap
high water; MLW, mean low water; MSLW, mean spring
low wNater (from Provost, 1973).






almt~eedn. ad re .. tea air
tamp .. t ... .. lee, s 25F -y kill the t ..
i, al! but the most protected locations as far











ma..h, other Ppeci,, such as black cosew ee
ocvica-a untida Jacq.), white manfrove





t lis thus this.. (fig. 16).al
Jpp I





the J.-sr m ..ihe of the Gulf coast,
f.p .ctha Flo ida the g ...y y-Darsh,,
of the ..'th give way to luxuriant evergreen






S-o- IST IUTIO







ChppkPp, SouthC G-.,
'-'n






P;.t C. forpplarvaeppppmpmpabundant ip
W -OW.I--I .-R A/ 1 pe, I -









gph-Ip hd Ipeakin JuIy. JanonbPPI






(1965) summarized the information on the
seasonal occurrence of the New York
Culicoides, and Battle and Turner (1971) did
the same for the Virginia species.
Khalaf (1969) reported in detail on the
phenology of the Culicoides occurring around
New Orleans, Louisiana, by separating species
into 4 groups as follows: 1) Aestival species,
with high incidence in spring, summer, and
fall. Florida species with this seasonal pattern
include C. crepuscularis, C. furens, C.
haematopotus, C. stellifer, and C. venustus
Hoffman. 2) Vernoautumnal species, with high
populations in spring and fall, but virtually ab-
sent in summer. C. hollensis has this type
ofannual cycle. 3) Pseudovernal species, with
high spring populations but low incidence in
both fall and summer. C. spinosus Root and
Hoffman is an example. 4) Vernal species with
high populations only in spring. C. biguttatus,
C. bickleyi Wirth and Hubert, C. niger Root
and Hoffman, and C. travisi are listed, but
many rare or uncommon species appear to
have this type of distribution.
Frost (1964) published a summary of insects
taken in black light traps at the Archbold
Biological Station, Highlands County, Florida,
from 1958 to 1963. His records of six
Culicoides species are included under their
respective species headings below.
Detailed records were kept of the species in
succession in light trap collections made by W.
H. Whitcomb at Monticello in Jefferson Coun-
ty, Florida, from January through October,
1969. Significant populations of Culicoides ap-
peared in April; the dominant species was C.
niger (500 daily), and there were lesser
numbers of C. scanloni Wirth and Hubert (50),
C. biguttatus (25), C. venustus (25), and C.
haematopotus (10). In May and June the
Culicoides populations gradually decreased
and July collections consisted of C.
crepuscularis (100), C. niger (50), C stellifer
(25), C. venustus (10), C. haematopotus (5), C.
guttipennis (5), and C. variipennis (5). By
September the species.composition had shifted
noticeably so the predominant species was C.
crepuscularis (200), and there were lesser
numbers of C. stellifer (50), C. haematopotus
(20), C. venustus (10), C. variipennis (5), and C.
debilipalpis (5).

NATURAL ENEMIES
Little is known about the natural enemies of
sand flies in North America, and a comprehen-
sive study is badly needed. Bacon (1970) sum-
marized what is known of the subject
throughout the world. Weiset (1963) reviewed
our knowledge of pathogens of insects (in-
cluding ceratopogonids) of medical importance


in Europe and suggested their potential value
in biological control. Becker (1958) published
the most comprehensive study of natural
enemies, working with the Scottish species.
Chapman and his co-workers in Louisiana have
devoted more attention to this field than
anyone else in the United States. Wirth (1977)
reviewed the published literature on parasites
and pathogens of biting midges up to 1976.
Viruses: According to Chapman et al. (1968),
an iridescent virus has been found in Louisiana
that causes death of larvae of C. arboricola and
C. guttipennis breeding in tree holes.
Bacteria: Hertig and Wolbach (1924)
reported cocci (Rickettsiae (?)) infecting the fat
bodies of adult C. sanguisuga in
Massachusetts. Becker (1958) reported infec-
tion by Pseudomonas sp. in the fat bodies of C.
salinarius Kieffer in Britain.
Protozoa: Becker (1958) reported ciliates of
the genus Perezella infesting the haemocoel of
several species of Culicoides larvae in Britain.
A ciliate, Balantidium knowlesii, was describ-
ed by Ghosh (1925) from the haemocoel of
adult C. peregrinus Kieffer in India. The
pathogenicity of these ciliates to their host is
not well known but is thought to be very im-
portant.
Several Microsporidia have been found to
cause mortality in Culicoides larvae. Chapman
et al. (1968) found Pleistophora species causing
milky white discoloration and death of C.
nanus larvae in Louisiana. They also found
another species of Pleistophora in Culicoides
sp. larvae from a ground pool.
Nematoda: Mermithid worms are common
parasites of Culicoides throughout the world
and cause varying degrees of intersexuality of
the adults if they attack the developing gonads
of the mature larvae (Beck, 1952; Callot, 1959;
Dzhafarov, 1960; Curtis, 1962; Glukhova,
1967; Smith and Perry, 1967; Chapman et al.,
1968; Nickle, 1969). The mechanism was
described in detail by Rempel (1940) who
studied mermithid parasitism in Chirono-
midae. Although the parasites do not cause im-
mediate death, reproduction is effectively
prevented. In our Florida collections we have
occasionally seen large numbers of C.
crepuscularis intersexes caused by mermithid
parasites (fig. 17).
Also, Smith and Perry (1967) reported a high
rate of mermithid parasitism and intersexes in
3 species of soil-dwelling Culicoides in Alachua
County, Florida. They stated that mermithids
parasitized approximately equal numbers of
male and female Culicoides, but that only the
males developed intersex features of antennae,
palpi, etc. Rates of parasitism in the species
sampled ranged from 0% in C. insignis, to 35%
















Iro





in C hMosssspotus, to 52% in
155% in o crepuseularis. Ldke


that earmlthid parasites offer O
natural control of insert pests
culture and hossemination of
should be at-nmpted.











Predators: Be5ker (1958) rep

only regular predators observe











predaSeous ad probably co


numbers of early inste mosquito laae and
)L various sized Larvae of Cuiicoides opp. that
breed to tre holes."
Becker 19581 observed adult Culcoides in


females of C. m.ss-ppensas collected on

T. .. . .

mangroves at waters edge, amid sparse Cen



COLONIZATION
As noted, basic biological re rch on
C Cufliod s an d critical studies oI disease
cbllifo, to a H
ar eb the ra te
... in large, sustaining laboratory colonies. The
contrast Is m1ked with the grat suress in
ds rm loniing and studying mosquitoes, lice, ticks,
and that p e a tes, and other arthropods of medical impor-
the parates tanes. This neglect of Culioides has sustained
thn lack of information that has promoted fur-

to attempt laboratory rearing of Culcoids. He
kept engorged female C oxystoma (now C
achul5sei) in a test tube on moist paper. Three
days after they engorged, the females
oviposited on the paper, and several of the
subsequent larvae were reared to the adult


hachrng, ... transferred to tap water that
turned green on standing, and the larvae were
-rted that in i.. -.. .....

d feedingon - i "-




adult stage,

,, u, ., years. He mainta ined a small c ony of C
,,. nubeulosus from 1947 to 1955 when it core-
pletely died out. MegaId (1956) tk1 re.


,',me ]a1 larval food,. Attempts to roar the t,' r, h







species, C. halophilus Kieffer and C. cir-
cumscriptus Kieffer, by this method, however,
were unsuccessful.
Jones (1957, 1960) was the 1st to develop a
large, successful colony by using mass-rearing
techniques. After bluetongue virus was
transmitted to sheep (Price and Hardy, 1954)
by using field-collected C. variipennis, Jones
(1957) established a colony of this species at
Kerrville, Texas. His stock of C. variipennis
sonorensis Wirth and Jones was especially
adaptable to colonization because the adults
fed readily on rabbits and other laboratory
animals and mated readily in the small
laboratory containers, and the larvae
developed readily on a medium consisting
mainly of soil, fresh cow manure, and driest
yeast. In 1972 this colony was transferred to
the Denver, Colorado, USDA laboratory.
Jones (1966a, 1966b) and Jones et al. (1969)
subsequently reported successful mass produc-
tion at a level of 1,000 flies per day.
Hair and Turner (1966) colonized C. guttipen-
nis, a tree hole species, by using a larval
medium of simulated tree hole water made by
soaking leaf mold collected from local woods in
distilled water. Larval rearing was done in
darkened aquaria where the larvae fed on
microorganisms developing in the leaf mold in-
fusions. Adults were held in /z-gallon paper
containers in semidarkness in constant
temperature and humidity cabinets and were
fed on rabbits daily. This colony was main-
tained for nearly a year, through 12-15 genera-
tions, and produced about 1,000 adults daily.
Linley (1968, 1969) described methods used
in colonizing C. furens in Florida. About 500
adults were trapped and released in a mating
cage about 2 feet on a side. Larvae were fed in
metal pans on a mud substrate to which free-
living nematodes and baby cereal were added.
The colony was maintained to the 4th genera-
tion. Eggs were deposited autogenously
without adult blood meals.
Morii and Kitaoka (1968) maintained a col-
ony of C. arakawai for 4 years by placing larvae
in pans on mud and charcoal with flowing
water and feeding them yeast and mouse food
pellets. Adults were kept in vials on rice straw
where they mated readily. Blood meals were
supplied with chickens. Sun (1969) colonized C.
arakawai and C. schultzei in Taiwan for 6
months through 6 generations. Adults were
collected in light traps and kept in 11 by 15 cm
cages. Larvae were reared in fingerbowls and
fed yeast-blood-agar base cake. Adults were
fed on chickens.
Federici (1967) reared 3 Florida Culicoides
species through complete life cycles in small
laboratory colonies by using commercial
animal diet as the larval medium. The larvae


fed mainly on the microorganisms growing on
the medium but to a lesser extent on the
medium itself. A rabbit provided blood for the
females of C. insignis; C arboricola and C.
haematopotus were fed on a chicken.

METHODS OF COLLECTION AND STUDY
ADULT SURVEYS
Females of Culicoides may be collected while
they are feeding on man and animals by
touching them lightly with a finger, forceps, or
brush moistened with alcohol, by using an
aspirator, or by collecting them in a fine-mesh
insect net. Bait traps are not satisfactory
because Culicoides are reluctant to enter an
enclosed space. However, Bennett (1960), Hair
and Turner (1968), and Humphreys and Turner
(1973) have had good success with baited drop
traps (fig. 18). Sommerman and Simmet (1965)
made excellent collections of Culicoides in
Alaska with an interception trap mounted on
an auto that patrolled roadways during the
hours of adult activity. Nevertheless the most
satisfactory method of determining species in-
cidence and relative abundance of Culicoides in
a general area is by light traps. We have had
steady success with ordinary New Jersey mos-
quito traps, particularly if the hood is designed
to allow the light to shine, at least slightly,
above the horizon. A small portable AC
generator may be used for operation in
habitats not provided with electric power. A
suction fan greatly increased the efficiency of
our light traps in collecting these small
midges, even in such weakly powered portable
traps as the CDC models. Light traps or vehi-
cle traps also are useful in collecting males,
which are extremely helpful in identification.
Unfortunately some species are not readily at-
tracted to light or do not travel far from their
breeding places.
Most of our specimens of Culicoides were col-
lected in New Jersey light traps (fig. 19), many
of which have been operated for years
throughout Florida by the Florida State Board
of Health. Likewise, Mrs. Beck's extensive
reports (1951-1958) were based on light trap
collections made in conjunction with the state
mosquito abatement program. We have now
operated New Jersey traps and black light
traps over much of the state for more than 10
years and have covered nearly all the counties
at least for short periods.
Dove et al. (1932) described a wood and cloth
"recovery cage" or emergence trap used in
their early studies of saltmarsh sand flies in
the southeastern United States. Subsequent
workers have effectively used boxes like these,
or variously modified, in sampling adult
emergence from suspected breeding areas.







i5
c





































,,




































amd, mass,
I placed in


7 lderi o ohe dl o
at ..11!






clipped t o yelmdu, f-t sed at .ale


coastal salmaar she. in oar h Cerebra, and in

(fig. M, but the ...It. for C.U-ide hav
20I


and pi-nd as soo.1 m -me. plus.

e-tst ideatifi-ati.. ..do, the hlghkr
magnification of a compo,,d mi ..... p,
Vmuasmr ,atr, ,Ibl,, chlo,,1-9=m media such
= 1"", o r ,-' ae oneient and sek
erelntmoraly rent. bu!. we a vis







moist cotton in small (1 x 5 cm) vials stoppered
with cotton plugs. If numerous pupae are
found, larger quantities of the substrate can be
collected and taken to the laboratory for mass
rearing or recovery of the larvae.
The larvae are recovered easily if they are
sieved and floated with magnesium sulphate,
or with ordinary sugar or salt in nearly
saturated solution. They rise to the surface
where they may be counted, pipetted off, and
preserved. Or they can be rinsed in clean water
and introduced into culture pans or study
chambers. Also, Berlese funnels are useful in
extracting larvae and pupae from semi-aquatic
media, especially species of the subgenus
Avaritia whose pupae do not readily float in
water. If larvae are to be preserved for study,
they may be killed by quick submersion in hot
water and preserved in 70% alcohol.




GEOGRAPHY AND LIFE ZONES OF
FLORIDA

A short discussion of the present physi-
ography and climate of Florida and a
reconstruction of its geological history are
helpful in explaining the distribution patterns
of Culicoides species.



PHYSIOGRAPHY

Florida may be divided conveniently into 3
parts; the narrow western coastal strip
sometimes called the "Panhandle," the long
peninsula extending nearly 400 miles
southward from the main continental mass of
North America; and the numerous lowlying
islands reaching toward the Cuban Antilles
that'comprise the Florida Keys. Although each
part has its own characteristics of climate,
flora, and fauna, the boundaries are indistinct
so that a person passes imperceptibly from one
into the other.
Northern Florida extends 384 miles from the
Perdido River on the Alabama state line to
Jacksonville on the Atlantic Ocean. Eastern
Florida extends 392 miles from the Georgia
border to the tip of the peninsula at Cape
Sable, but the extreme length from the 31st
parallel to the southernmost Keys is about 444
miles. The state has 1221 miles of tidal shore
line, 714 miles bordering the mainland and 507
miles surrounding islands. The boundaries of
the state enclose an area of 54,861 square miles
of land and 3,805 square miles of water, a total
of 58,666 square miles (Cooke, 1939). ,


CLIMATE
The climate of Florida is oceanic because of
its proximity to the Atlantic Ocean and the
Gulf of Mexico. The summers are long and
warm, the winters are short and mild, and
there are no long periods of extreme heat or
cold. North Florida falls in K6ppen's zone of
Warm Temperate Rainy Climate, but south of
Lake Okeechobee the climate is classed as
Tropical Rainy Climate.
The climate of northern Florida is
characterized by mild, comparatively dry
winters and long summers with a pronounced
summer rainy season. The seasonal contrast in
rainfall is more pronounced as one travels
south down the peninsula but becomes less
distinct as one reaches the Panhandle, where
the rainfall is more uniform throughout the
year. Rainfall varies from year to year from fre-
quent wet years when lakes, marshes; and
shallow basins fill up and flood to dry years
when the water table recedes and the peaty
marshes catch fire. Tropical storms occasional-
ly bring high winds and extremely heavy rains.
Because of the tropical marine exposure, the
coastal areas are warmer than the interior in
winter and cooler in summer. Frequent after-
noon thundershowers prevent the frequent oc-
currence of extremely high temperatures dur-
ing the summer, and sea breezes temper the
summer heat. The presence of warm water,
especially that of the Gulf Stream along the
east coast, ameliorates the cold waves of
northern air that occasionally sweep down in
winter. Thus the zone of tropical climate and
vegetation extends far northward along the
coasts, especially on the east coast, but the
center of the peninsula is relatively temperate
as far south as Lake Okeechobee. Since no
point in Florida except the southernmost Keys
escapes occasional killing frost, the climate of
southern Florida can only be classed as sub-
tropical. A summary of climatic data for
selected Florida stations is given in table 1.


GEOLOGY
The geologic history of the Florida peninsula
was reviewed by Cooke (1939) and White
(1970). The following account is summarized
from their publications. Present-day Florida
occupies only a fraction of a much larger
geographic unit, the Floridian Plateau (fig. 21).
This plateau is a partially submerged platform
250 to 400 miles wide that extends from the
deep water of the Atlantic Ocean to the deep
water of the Gulf of Mexico. It is attached to
the continent of North America on the north
and extends southward nearly 500 miles to the







Table 1. Summary of climatic data for selected
Florida localities from Climate and Man, USDA Yearbook of Agriculture 1941

Temperature OF Growing Rainfall
Average Season %
Jan. July Max. Mi. Days Annual May-Oct.
Marianna 52.8 81.0 106 13 264 54.51 55
Tallahassee 54.5 80.8 104 -2 282 54.89 59
Gainesville 57.6 80.8 103 6 285 49.10 66
Jacksonville 56.8 81.4 104 10 299 48.21 67
Orlando 61.4 82.0 103 18 314 52.45 -
Tampa 61.8 81.6 98 19 348 48.35 73
Fort Myers 64.5 81.1 98 24 killing 52.06
frosts
Fort Lauderdale 68.9 81.4 99 28 62.98 -
Homestead 66.8 80.5 98 26 61.88 85
Key West 69.9 83.2 100 41 no k.f. 38.36 -


Straits of Florida, which separate it from
Cuba.
The Floridian Plateau consists of a core of
metamorphic rocks that are continuous with
the Georgia Piedmont. The core lies buried
under more than 4,000 feet of sedimentary
rocks, mostly limestone, the seaward extension
of the Coastal Plain of Georgia and Alabama.
The Floridian Plateau has been relatively
stable for millions of years, alternately dry
land or covered by shallow seas, and has not
been subjected to violent crustal movements
or earthquakes. The only deformations have
been a gentle doing that has elevated the nor-
thern part of the state to a little more than 325


Fig. 21. Outline of the Floridian Plateau and
Topographic Divisions of Florida (after Cooke, 1939).


feet above sea level and a gentle downward dip
of the older strata in the south.
Cooke divided the portion of the Floridian
Plateau that lies above sea level, the present-
day State of Florida, into 5 natural
topographic regions: (1) The Coastal Lowlands,
the most recently emerged region, which lies
generally less than 100 feet above sea level,
and 4 interior, generally higher, hilly regions;
(2) the Western Highlands; (3) the Marianna
Lowlands; (4) the Tallahassee Hills; and (5) the
Central Highlands. Berner (1950) and Beck and
Beck (1966) presented maps based on Cooke's
5 topographic regions.

Biogeographic Divisions of Florida.
Harper (1910, 1914, 1921, 1927) divided
Florida into natural divisions that he
characterized mainly on the basis of vegeta-
tion: (1) West Florida Coast Region; (2) West
Florida Pine Hills; (3) West Florida Limestone
Region; (4) Middle Florida Hammock Belt; (5)
Lime-sink Region; (6) Middle Florida Flat-
woods; (7) Gulf Hammock Region; (8) Lake
Region; (9) East Florida Flatwoods; (10) East
Coast Strip; (11) South Florida Flatwoods; (12)
Miami Limestone Region; (13) Coast Prairie;
and (14) The Keys.
Howell (1932) preferred a more ecological
grouping of the prominent physiographic
features of the state, which he described as (1)
Flatwoods, (2) High Pineland, (3) Hammocks,
(4) Sand Scrub, (5) Swamps, (6) Everglades, (7)
Salt Marshes, (8) Sea Beaches, (9) Prairies, and
(10) The Keys. He felt this classification more
adequately reflected the distribution of bird
life in Florida.







Rogers (1933) described 19 major habitats in
northern Florida that affected the distribution
of crane flies (Tipulidae):
Sea Coast Habitats: (1) Juncus flats of tidal
marshes.
Inland Habitats:
Aquatic and Semi-aquatic Situations:
A. Streams: (2) seepage areas and small
rills; (3) small streams; (4) swamp and
bog streams; (5) larger calcareous
streams; (6) lower streams.
B. Ponds, lakes, marshes, and swamps:
(7) fluctuating ponds; (8) sinkhole
ponds; (9) lakes; (10) marshes; (11)
swamps.
Terrestrial Situations:
A. Hammocks or hardwood formations:
(12) low hammock; (13) magnolia-
laurel oak-holly-white oak hammock;
(14) high hammock.
B. Pine lands: (15) pine flatwoods; (16)
high pine; (17) turkey oak on cut-over
high pine lands; (18) sand scrub.
(19) Restricted Areas of Particular Interest:
(a) Devil's Mill Hopper, Alachua County; (b)
Torreya ravines of Liberty County; (c) Red clay
hills of northwestern Florida; (d) Ash and gum
swamps of Jackson County.
Berner (1950) followed Cooke's physio-
graphic divisions in his monograph on the
mayflies (Ephemeroptera) of Florida, and
described the following types of mayfly
habitats: (1) Permanent creeks: (a) sand-
bottomed creeks with little vegetation; (b) silt-
bottomed creeks with little vegetation; (c)
sand-bottomed creeks choked with vegetation;
and (d) silt-bottomed creeks choked with
vegetation. (2) Rivers: (a) stagnant rivers; (b)
slow-flowing, deep rivers; (c) larger calcareous
streams. (3) Ditches and puddles: (a) roadside
ditches; (b) puddles. (4) Ponds: (a) sinkhole
ponds; (b) fluctuating ponds; (c) temporary
woods ponds; (d) sporadic ponds; (e) Jerome
Sink. (5) Lakes: (a) sand-bottomed lakes; (b)
silt-bottomed lakes; (c) disappearing lakes. (6)
Marshes. (7) Swamps: (a) cypress swamps; (b)
bayheads. (8) Springs.
Young (1954) regrouped Cooke's 5 topo-
graphic regions and discussed them in relation
to the numerous but more discrete geographic
regions proposed earlier by Harper (1914,
1921) and Davis (1943) for north, central, and
south Florida, respectively. Thus, Young sub-
divided Cooke's Coastal Lowlands into 3
categories: (1) the West Florida Lowlands or
Flatwoods; (2) various Peninsular Florida
Lowlands or Flatwoods; and (3) the South
Florida Lowlands. Moreover he recognized 3
adjuncts to the South Florida Lowlands, which
he called: (a) the Rim of the Everglades or East
Coast Ridge Area; (b) the Everglades Keys


Area; and (c) the Florida Keys. Also, instead of
"Highlands," Young preferred "Uplands" for
Florida's hilly regions, which he called (4) the
West Florida Uplands, and (5) the Peninsular
Florida Uplands. He considered the (a) Marian-
na Redlands Area and (b) the Apalachicola
Ravines and Bluffs Area as adjuncts to the
West Florida Uplands.
Neill (1957) recognized 7 biogeographic areas
of Florida, of which he said some were much
better defined than others. He listed: (1) the
Western Panhandle; (2) the Apalachicola
Drainage; (3) the remainder of the panhandle,
not markedly different from adjoining portions
of the Gulf Coastal Plain in Alabama and
Georgia; (4) Northeastern Florida, but slightly
differentiated from the Lower Atlantic Coastal
Plain of Georgia; (5) the Central Florida
Highlands; (6) the Gulf Hammock; and (7) the
Subtropical Tip of the peninsula. He noted that
these areas usually were not clearly defined
and that fairly wide zones of transition exist,
perhaps because many of the Pleistocene bar-
riers to distribution are no longer extant.
Beck (1965) classified the streams of Florida
as habitats controlling the distribution of
aquatic invertebrates. He recognized: (1) sand-
bottomed streams; (2) calcareous streams; (3)
larger rivers; (4) swamp and bog streams; and
(5) canals of southeastern Florida.
McPherson et al. (1976) published an
elaborate review of the ecosystems of south
Florida as a summary of 51 separate reports
made by 5 bureaus or former bureaus of the
United States Department of Interior under
phase 1 of a project entitled the "South Florida
Environmental Study." In response to
ecological concerns about the proposed con-
struction of an international jetport in the Big
Cypress Swamp, the South Florida En-
vironmental Study was begun in 1971 under an
agreement among the State of Florida, the
Department of Interior, and the University of
Florida Center for Wetlands. Their review
broke down the south Florida regional eco-
system into 3 components with the following
subsystems:
Freshwater and terrestrial ecosystems:
Canals and lakes; Ponds and sloughs;
Sawgrass marshes; Wet prairies; Pine forests;
Cypress forests; Mixed swamp forests; Bay
heads; Hardwood hammocks; and Palmetto
and dry prairies.
Coastal ecosystems: Sandy beaches;
Mangrove and saltmarshes; Shallow estuaries
and bays; and The reef tract.
Man-dominated ecosystems: Agricultural;
and Urban.
We now will discuss some of the geographic
and ecological peculiarities of these divisions
in relation to the Culicoides fauna.


































Fig. 22. Wicomico and pre-Wicomico shore lines in the
Southeastern States (after Cooke, 1939).




Coastal Lowlands
The Coastal Lowlands border the Florida
coast from Georgia to Alabama and extend in-
land from 30 to more than 125 miles since the
inner margin lies approximately at the 100-foot
contour line. They originally were plains
representing the 4 marine terraces (Wicomico,
earliest and highest; Penholoway; Talbot; and
Pamlico, the latest and lowest (fig. 22-25).
These terraces formed the bottoms of the seas
during 4 high water levels of the late
Pleistocene interglacials. Because drainage is
poor and the climate is warm and rainy, the
soils of the Coastal Lowlands have developed
into ground-water podzols and half-bog soils
from marine sands that rest unconformably on
clay or rock. The resulting "hardpan" or
organic material and iron oxides in the subsoil
prevents effective water movement in the soils,
which do not percolate during heavy rains and
prevent subsurface water from moving upward
during dry periods.
Most of the Coastal Lowlands are covered by
forests of longleaf and slash pine with
undergrowth of saw palmetto, wire grass,
runner-oaks, and gallberry. Cypress, gums, red
maples, wax myrtles, cabbage palms, and


hollies are found in the swamps, around ponds,
and along streams. Most of the plants are ex-
tremely hardy, adapted to withstand alternate
and prolonged flooding and drying, and highly
resistant to recurrent fires.
The Everglades lie south of Lake
Okeechobee and consist of large expanses of
saw-grass savannas. Extensive mangrove
swamps are found along the salt and brackish
coastal waters south of 200 latitude, and
grassy saltmarshes prevail farther north.
1. West Florida or Apalachicola
Flatwoods. The Apalachicola Flatwoods,
1st characterized by Harper (1914), extend
from the Wakulla to the Choctawhatchee
rivers. This poorly drained pineland is noted
for its profusion of pitcher-plants (Sarracenia)
and other bog plants; a high percentage (15%)
of Ericaceae characterizes the shrubs, % of
which are evergreen. Since this area has been
cut off from the eastern flatwoods for a long
period by a high ridge in central Wakulla and
southern Leon counties, there is considerable
endemism in its plants and animals. Hobbs
(1942) described an endemic group of crayfish,
the Procambarus rogersi complex, whose
species are confined to the Apalachicola Flat-
woods. Young (1954) speculated that this
region may have served as a "bay of refuge"
where pre-Pleistocene forms have been concen-


Fig. 23. Penholoway shore line in the Southeastern
States (after Cooke, 1939).







treated. The Culicoides fauna of the
Apalachicola Flatwoods is characterized by
high populations of C. niger; at the
Ochlockonee River State Park, large numbers
of C. biguttatus, C. haematopotus, and C.
scanloni also were taken. In the tidal salt-
marshes near the coast (fig. 10) which are
characterized by growth of Juncus
roemerianus Scheele, C. mississippiensis
predominates, and there are lesser numbers of
C. furens, C. melleus, and C. bermudensis.
2. Peninsular Florida Flatwoods. The flat-
woods of most of Florida are much the same as
those of eastern Georgia, southern Alabama,
and Mississippi, which are sometimes termed
the Sabalian Biotic District. In peninsular
Florida, the outlines of the region are rather
irregular and patchy, and the region is subject
to some variation because of elevation and
minor climatic differences. Surface drainage is
slow and inadequate and the "hardpan" is
resistant to drainage erosion. Water collects in
characteristic shallow ponds, but some are
deep and permanent, usually underlain with
Bayboro and Portsmouth sandy soils, and sup-
port vegetation of cypress, gums, Juncus, and
Sphagnum. Streams (fig. 26) are sluggish,
shallow, mud-bottomed, and dark with pro-
ducts of organic decay. Occasionally they have


Fig. 24. Talbot and post-Talbot shore lines in the
Southeastern States (after Cooke, 1939).


Fig. 25. Pamlico and pre-Pamlico shore lines in the
Southeastern States (after Cooke, 1939).


sand bottoms for short distances. They usually
are choked with vegetation such as Juncus,
Eleocharis, Persicaria, Pontederia, waterlilies,
and water hyacinths (fig. 27). The more perma-
nent streams support luxuriant growths of
hardwood trees and shrubs, many evergreen,
that are called "bays" or "bayheads." Toward
the south, the flatwoods merge gradually into
the Everglades where prairie and marshy (fig.
28) conditions become dominant. Culicoides
species characteristic of the Peninsular Flat-
woods include C. arboricola, C. crepuscularis,
C. floridensis, C. niger, C. scanloni, C. spinosus,
and C. stellifer.
3. South Florida Lowlands. South of Lake
Okeechobee the Peninsular Flatwoods gradual-
ly merge into an extensive savanna that
becomes the Everglades (fig. 29) and the Big
Cypress Swamp region. The bottom of Lake
Okeechobee and of most of the northern
Everglades was laid down by the Pamlico Sea,
the last of the interglacial seas of the Ice Age.
Much of the floor of the remaining Everglades
was formed of more recent freshwater marl
that forms an impervious layer over the
Pamlico sands. Originally the marshy
sawgrass glade land was covered with shallow
water most of the time, and deposits of peaty


-- - - - -- - - - -








-- - - - - - - -


-- - - - -- - - - -



















Fig. 26. Small stream in hardwood hammock in Fig. 28. Muddy cow pasture at the Ona Range Experi-
Highlands Hammock State Park, Highlands County, ment Station in Hardee County, Florida; larval habitat of
Florida; larval habitat of Culicoides niger, C. .scanlo C Culicoides insignis, C. knowlton and C. pusillus.
spinosus, and C. stellifer.


soil were built up, but surface water gradually
trickled off into low channels extending toward
the coasts. In the northwestern part of the
Everglades area, large areas of cypress
characterize the Big Cypress Swamp. To the
south, in Dade and Monroe counties, the
sawgrass marsh of Mariscus is replaced by a
wet marl prairie interspersed with low ham-
mocks of bays, willows, wax myrtles, palms,
and other trees. Where the ground is rocky and
higher, hammocks of tropical hardwoods have
developed. However, as southern Florida has
been settled, extensive systems of canals and
ditches have been constructed to allow
.agricultural development of the deeper
Everglades muck soils, especially near Lake
Okeechobee. As the water table has lowered,
the glades themselves have suffered more fre-
quent fires, and much of the remaining muck
and peat has been destroyed, along with exten-
sive wildlife habitats (Craighead, 1971).
Culicoides species characteristic of the
Everglades include C edeni Wirth and Blan-
ton, C. floridensis, C. insignis, C. knowltoni

Es dBSL, rillM


Beck, and C. stellifer.
3a. Rim of the Everglades or East Coast
Ridge Area. From Fort Lauderdale to below
Homestead a rocky ridge with as much as 20
feet elevation supports pinelands of Pinus
elliottii Engelmann, var. densa Little and Dor-
man, with undergrowth of saw palmetto, silver
palm, dwarf oaks, poisonwood, wire grass, etc.
Over the bed rock of Miami oolite, which is
usually exposed and weathered into jagged
pockmarks, soils are alkaline and restricted to
residual marls and sands of marine origin.
Hammocks of tropical hardwoods have
developed around sinks or along stream
courses and are prevented from spreading by
the frequent fires that sweep through the
pinelands.
3b. Everglades Keys. The Everglades
Keys are isolated extensions of the East Coast
Ridge that were formerly separated by exten-
sive wet arms of the Everglades and so provide
extensive, isolated habitats of tropical ham-
mock and pinelands on which some tropical
American Culicoides have established


Fig. 29. Everglades with sawgrass and cabbage palms in
Fig. 27. Small branch of the Myakka River in Sarasota eastern Collier C.. Fl :,] i.- al habitat of
Co., Florida, choked with water hyacinth; larval habitat of Culcoides edeni C ... .. ..a C. knowltoni
Culicoides insignia, C. edeni C. knowlton and C. stellifer. and C. stellifer.


' .'.-.aT4 .







footholds in North America.
3c. Florida Keys Like the Everglades
Keys, the islands of the Florida Keys are
basically extensions of the East Coast Ridge.
Generally the Keys are dry, sandy, or rocky
islands with little soil or fresh water. The
southern Keys are larger and have more pot
holes and freshwater pools and extensive
pinelands and hammocks.
3d. Southwest Coast and Ten Thousand
Islands. From Cape Sable to Cape Romano
there is an unbroken expanse of dense
mangrove swamps and saltmarshes. Tidal
ranges up to 4 feet bring salt water far inland
and form a maze of islands, water courses, and
mangrove swamps that are virtually
uninhabited. Fishermen venturing into these
passages meet tremendous populations of
hungry C. furens and C. barbosai.
Davis (1943) outlined and discussed the
natural features of the South Florida
Lowlands, with particular emphasis on the
physiographic regions (of which he recognized
11), geology, soils, topography and hydrology,
vegetation, and fish and wildlife. He recogniz-
ed, described, and mapped 9 vegetational
types in this area: (1) Pine forests; (2) Ham-
mock forests; (3) Inland swamp and cypress
forests; (4) Coastal mangrove swamp forests;
(5) Beach and dune vegetation; (6) The
Everglades and other freshwater marshes; (7)
Salt-water marshes and salt- or brackish-water
prairies; (8) Wet prairies; and (9) Dry or saw-
palmetto prairies.
South Florida Lowland Culicoides with
special habitats are as follows: C. loughnani
Edwards, which breeds in the cacti on the Keys
and coastal sand areas (fig. 9); C. barbosai and
C. furens, which breed in the coastal mangrove
swamps (fig. 15, 16); C. melleus, which is found
on sandier shores of bays and inlets (fig. 10); C.
mississippiensis and C. bermudensis,which are
found in open saltmarshes (fig. 11); and C. in-
signis, which appears as the water becomes
less saline.

Florida Highlands
4. West Florida Uplands. The hilly West
Florida Uplands lie between the Perdido and
Apalachicola rivers and extend from the
Coastal Lowlands north into Alabama and
eastward along the southern edge of the
Marianna Redlands. In the north, the hills are
as much as 300 feet above sea level, southward
there is a broad, gently rolling upland. The
remnants of 3. marine terraces (Brandywine,
Coharie, and Sunderland) (fig. 30) that emerged
during the early Pleistocene seas are found in
the areas below 270 feet. The West Florida
Uplands are deeply trenched by narrow, steep-


walled valleys of the Escambia, Blackwater,
Yellow, and Shoal rivers, and by several
smaller rivers. The heads of the valleys of
many streams are strikingly steep, the valleys
are narrow because the underlying, sand has
drained off most of the rainfall, and surface
erosion has been slight. Numerous lakes and
sinks have resulted from solution of the
underlying rocks by percolating rainwater dur-
ing the Pleistocene glacial periods when sea
levels were lower and the water table was ac-
cordingly depressed. Soils of the West Florida
Uplands belong for the most part to the red-
yellow podzol group of lateritic coastal plain
soils, of which the Norfolk, Ruston, and
Orangeburg sands are the most widespread.
The streams are usually small and clear and
have only poorly developed marginal swamps.
Ponds, lakes, and marshes are not common ex-
cept in local areas where solution has formed
many lime sinks and cypress ponds. The
vegetation of the West Florida Uplands is
more varied than that of the flatwoods, and in-
cluded a greater variety of deciduous trees and
conifers, of which the beech is characteristic.
Common Culicoides species of the West
Florida Uplands are C. arboricola, C. bigut-
tatus, C. haematopotus, C. mulrennani Beck, C.
niger, C. scanloni, C. stellifer, and C. venustus.


I',
Ti


i:
w
~3 i
" :






I






,


SUmERLAND


/
S ... . i ,
U ".-:-- rl


Fig. 30. Early Pleistocene shore
Southeastern States (after Cooke, 1939).


lines in the


I







4a. Marianna Redlands.-The Marianna
Redlands is a limestone area in Jackson,
Holmes, and Washington counties. It consists
of low, rolling hills and hollows and is dotted
with sinks, ponds, and lakes caused by solution
of the underlying limestone. Soils are heavy
red clay and loam for the most part, some of
the richest in Florida. Farming has altered
most of the native forest cover, and what re-
mains is a sparse 2nd growth of Pinus taeda
Linnaeus and P. echinata Miller intermixed
with deciduous trees. Aquatic situations are
scarce and usually are muddy or clayey tem-
porary ponds and puddles. The most common
Culicoides taken in our collections at Florida
Caverns State Park were C. arboricola, C.
baueri Hoffman, C. husseyi Wirth and Blan-
ton, C. mulrennani, C. spinosus, C. travisi and
C villosipennis. Macropeza blantoni Wirth and
Ratanaworabhan is an interesting ceratopo-
gonid taken at this locality; this endemic
species is the only species of Macropeza found
in the Western Hemisphere. The genus is com-
mon and widespread in Africa and Asia, and
several species occur in Europe.
4b. The Tallahassee Hills form a 100-mile ex-
panse of gently sloping hills with rounded sum-
mits adjoining the Marianna Redlands on the
east. Part of this area is a high plain lying at
more than 300 feet above sea level. This plain
is now being dissected into valleys and ridges
by rivers such as the Chattahoochee and its
tributaries. Sweet gum, live oak, and water oak
are abundant because of high phosphorus soils.
Loblolly pine, dogwood, red oak, hickory, post
oak, black-jack oak, and black oak thrive on
high iron or aluminum soils or both. Pinus
taeda, Magnolia grandiflora Linnaeus, beech,
and Parthenocissus are common because of the
abundance of humus. W. H. Whitcomb ran
light traps extensively at Monticello where the
following Culicoides species were common: C.
arboricola, C. crepuscularis, C. niger, C.
stellifer, C. variipennis, C. venustus, and C.
villosipennis.
4c. Apalachicola Ravines and Bluffs
Area.--Northwestern Liberty County and
western Gadsden County contain an unusual
area of high relief with high bluffs and deep
ravines along the Apalachicola River. The
region contains some of the most interesting
endemics in Florida and many plants with
northern affinities that occur nowhere else in
the state. The conditions along these bluffs ap-
parently have remained suitable for the sur-
vival of many forms that inhabited Florida
during the Pleistocene glaciations but have
died out elsewhere in the state. Other northern
forms probably followed the rivers from nor-
thern Georgia and Alabama and found suitable
habitats. Cooler temperatures and/or higher


rainfall during glacial times brought many
plants and animals with northern distributions
into Florida. Indeed, during Wisconsin times
Florida was too cool to support a tropical flora,
but cool and wet enough to support some
organisms that now occupy the Transition or
even more northerly life zones (Neill, 1957).
Two yews, the torreya (Torreya taxifolia Ar-
nott) and Florida yew (Taxus floridana
Nuttall), head the list of notable plant and
animal endemic species in this area that have
been discussed by Neill.
Neill (1957) also stated that many
peculiarities of east-west distribution in the
Florida Panhandle can be explained by the
deep embayments of the Gulf of Mexico along
3 river systems during the Pleistocene in-
terglacials: (1) the Escambia-Blackwater-
Yellow River basin; (2) the Choctawhatchee-
Alaqua basin; and (3) the Apalachicola River
and its tributaries. Many organisms find their
eastern or western limit of their range at one or
another of these Pleistocene embayments. C.
guttipennis and C. hinmani, for example have
been taken in Florida only in northern Florida
east of the Apalachicola River.
At Torreya State Park we consistently
trapped more species of Culicoides than at any
other Florida locality. The commonest species
were C. husseyi, C. mulrennani, C. niger, C.
spinosus, and C. villosipennis. C. footei was
found in Florida only at this location.
5. Peninsular Uplands. The crest of the
Floridian Plateau forms the Peninsular
Uplands of Florida which extends about 250
miles from Georgia south to Glades County
where it tapers to a blunt point. In the north
the Peninsular Uplands includes the nearly
level plain of the Sunderland Terrace at about
150 feet above sea level down to 120 feet where
it forms the southern edge of the Okefenokee
Swamp. Hilly areas at 100 to 200 feet elevation
extend southward, and are interspersed with
some broad low plains (parts of the Wicomico,
Penholoway, and Talbot Terraces) that contain
lakes such as Lake Panasoffkee and Tsala
Apopka. The Lake Region adjoins this area on
the east and extends southward to the end of
the Uplands where the relatively high relief of
sandy hills attains a height of 325 feet at Iron
Mountain near Lake Wales. Streams are scarce
because the underlying limestone carries off
the rainwater underground. As a result many
sinks and lakes have been formed by dissolu-
tion. During glacial times when the sea level
was lower, rainwater was actively eroding the
limestone to form large underground rivers
and caverns. Many of the caverns have col-
lapsed forming numerous springs, sinks, and
lakes (fig. 30).






The soils fall in 3 main groups: (1) the
Fellowship, Hernando, and Gainesville sands
and sandy loams of the middle Florida ham-
mock belt; (2) Norfolk or Lakeland and Blanton
sands of the peninsular lime sink region; and
(3) Norfolk or Lakeland, Eustis, Lakewood and
St. Lucie sands in the Peninsular Lake Region.
Since the soils are impoverished, they add few
minerals to the surface water except calcium
and some phosphorous and so limit the produc-
tivity of the lakes and streams. Plants
characteristic of the Peninsular Uplands fall in
2 groups: the high pinelands have open forma-
tions of longleaf pine and a saw palmetto
understory; the sandier and more xerophytic
ridges have been taken over by a dense scrub
comprised of sand pine (Pinus clausa
(Engelmann) Vasey), small evergreen oaks,
saw palmetto, huckleberry, and rosemary. The
presence of many endemic Florida species of
plants and animals in the high pineland and
rosemary scrub of the Peninsular Uplands is
thought to reflect the isolation of these areas
as island land masses in the Miocene and dur-
ing the Pleistocene sea level fluctuations (Neill,
1957).
Characteristic Culicoides species of the
Peninsular Uplands are C. alachua Jamnback
and Wirth, C. baueri, C crepuscularis, C. edeni,
C. floridensis, C. knowltoni, C. scanloni, C.
spinosus, C. stellifer, and C. tissoti. This region
is the center of endemism for Florida
Culicoides, with C. alachua, C. edeni, C.
floridensis, C. knowltoni, and C. tissoti found
elsewhere only rarely.
The following is a summary of the Florida
Culicoides species characteristic of major
geographic divisions.
1. Widespread, found over the entire state:
C. arboricola, C. crepuscularis, C.
debilipalpis, C. paraensis. In coastal
saltmarshes over the entire state: C. ber-
mudensis, C. furens, C. melleus.
2. Entire North Florida: C. haematopotus, C.
hinmani C. niger, C. piliferus Root and
Hoffman, C. spinosus, C. stellifer, C.
villosipennis.
3. Entire South Florida: C. edeni, C. floriden-
sis, C. insignis, C. pusillus Lutz.
4. Florida Uplands (West and Peninsular):
C. baueri, C. bickleyi, C chiopterus, C.
juddi Cochrane, C. nanus, C. scanloni, C
torreyae Wirth and Blanton.
5. West Florida Uplands: C. beckae Wirth
and Blanton, C. biguttatus, C. guttipen-
nis, C. husseyi, C. mulrennani, C.
ousairani Khalaf, C. testudinalis Wirth
and Hubert, C. travisi, C. variipennis, C.
venustus.
6. Peninsular Uplands: C. alachua, C. snowi,
C. tissoti.


7. Apalachicola Ravines and Bluffs Area: C.
footei, C. parapiliferus Wirth and Blan-
ton.
8. West Florida or Apalachicola Flatwoods:
C. loisae Jamnback.
9. Peninsular Florida Flatwoods: C.
knowltoni (prairies).
10. South Florida Lowlands: C. barbosai
(coastal saltmarshes), C. loughnani
(coastal cactus).


MORPHOLOGY
ADULT
Good descriptions of the structure of adult
Culicoides (fig. 31) have been given by Carter
et al. (1920), Jobling (1928), Gad (1951),
Tokunaga (1937), Wirth (1952a), Wirth and
Blanton (1959), Jamnback (1965), Atchley
(1967, 1970), Ortiz (1969), and Rowley and
Cornford (1972), to whom the reader is referred
for a fuller treatment. The following brief ex-
planations are given to define the most impor-
tant characters used in classifying Culicoides.
Head (fig. 32a): The head is subspherical,
with the anterior surface more or less flattened
and in line with the anterior surface of the pro-
boscis. The compound eyes are large and reni-
form and more or less contiguous above the
bases of the antennae; they may be bare or
with short pubescence between the ommatidial
facets. The degree of separation or contact of
the eyes is often useful in distinguishing
species; in the angle between the eyes on the
frons is an interocular seta, and above this is
often a transverse suture which marks the
separation of the frons and the vertex.
The antenna has 15 divisions which for con-
venience in this paper are termed segments,
although it is recognized that the 13 divisions
of the flagellum are not true segments in the
morphological sense. The basal scape is
ringlike, submerged in the head capsule, and
hidden by the greatly enlarged pedicel; the 1st
flagellar segment is slightly enlarged and
always bears a number of small sensory pits
(sensilla coeloconica), each surrounded by
minute setae; some or all of the distal segments
also bear these distal sensilla (fig. 33d), the
number, shape, and distribution of which are of
great importance in classification. In the
female the 1st 8 flagellar segments are shorter
and bear long verticils basally, and the 5 distal
segments are more elongated and without ver-
ticils; the ratio of the combined lengths of the 5
elongated distal segments divided by the com-
bined lengths of segments 3-10 forms the
antennal ratio (abbreviated AR). In the male
the pedicel is more enlarged and contains the
important Johnston's organ. The transition in






















2ND RADIAL CELL_
R1
1ST RADIAL CELL
SUBCOSTA
R-M CROSS-VEIN
M1+2
RADIUS
COSTA
BASAL ARCULUS
LATERAL SUTURE
ANEPISTERNAL CLEFT
PSEUDOSUTURAL FOVEA \ _'_
HUMERAL PIT_
ANTERIOR SPIRACLE --. \
SCAPE
PEDICEL


Ml
..______" CELL M1
M2
'h-.'.'. ____ __ CELL M2
,', -.;, ' i M3+4

CELL M4
"' ,, '" MEDIAL FORK
____MEDIO-CUBITAL FORK

Cu2
/__ IST A
/ / ."1- ANAL CELL
,' .-____-.HALTER
S.' SCUTELLUM
S- _____________POSTSCUTELLUM


i-
-


WING BASE (EXCISED)
POSTERIOR SPIRACLE
TPLEURAL MEMBRANE
,\ i SPERMATHECAE
.'CERCI

-~,i~~u~uu~u~~u~uu~u~~u~uu~u..,,.,; RC
%' ..
. .:~
'."1 .. . "


CLYPEUS

MAXILLARY

PROBOSCIS


;ITARSUS


Fig. 31. Adult female of Culcoides furens, lateral view with left king and right legs removed, with parts labeled (cx,
coxa; em, epimeron; ep, episternum; m, meron; pn, pronotum; pp, propleuron; px, precoxale; tr, trochanter).












VERTEX--- -------
MEDIAN BRISTLE BASE------
OCELLUS -------
COMPOUND EYE
FRONTO-CLYPEUS ---. f


---------------SCAPE
----------PEDICEL
,,-OLFACTORY PITS







_--MAXILLARY PALPUS
-----SENSORY PIT
Y -


I
c-MANDIBLE
------LACINIA
\-----HYPOPHARYNX


A. FEMALE HEAD


9TH STERNUM--------------
CAUDOMEDIAN EXCAVATION----
BASI STYLE----------
---- ^r ir


VENTRAL ROOT OF BASI


\ ------- --------------AEDEAGUS

S'\

\ \ \
\ \ \--------------9TH TERGUM
S\----- APICOLATERAL PROCESS
-----------------------CERCUS

B. MALE GENITALIA ._e-


Fig. 32. a, head and mouthparts of Culicoides female, schematic, parts labeled; b, male genitalia of Culicoides,
schematic, parts labeled (from Arnaud, 1956).


DISTISTYLE---


\J
\






length of the flagellar segments occurs be-
tween segments 12 and 13 in the male; each of
segments 3-12 has a whorl of greatly
elongated, erectile verticils forming a more or
less dense plume.
The mouthparts are well developed, often as
long as the head capsule, better developed in
the female than in the male, and in females of
most species are adapted for piercing and


a-c from Rowley and Cornford, 1972).


blood-sucking. They consist of 6 slender,
distally toothed blades of subequal lengths, in-
cluding a strong upper labrum-epipharynx, a
pair of maxillae, a pair of strongly toothed
mandibles, and a median tubular
hypopharynx. These parts are enclosed in a
proboscis formed by the fleshy part of the
labium; the length of the proboscis is of value
in classification and is expressed as the pro-


showingg deal of
ft A Rowley; fig.






boscis/head ratio (P/H Ratio), which is ob-
tained by dividing the distance from the end of
the labrum-epipharynx to the tormae, by the
distance from the latter to the interocular seta
base. The maxillary palpus is 5-segmented.
The 3rd segment is more or less swollen and
bears on the distal part of the medioventral
surface a specialized sensory pit or group of
sensilla which forms an important taxonomic
character (fig. 33). In measuring the palpal pro-
portions the line separating segments 1 and 2
is arbitrarily selected as halfway along the in-
distinct diagonal division between these
segments. The papal ratio (PR) is a useful in-
dex obtained by dividing the length of the 3rd
papal segment by its greatest breadth.
Thorax: The thorax is moderately broad and
convex above, arched anteriorly, and projec-
ting slightly over the head; it bears dorsally a
pair of small depressions behind the humeri,
known as the humeral pits, which have been
assumed to be sensory but whose exact func-
tion is unknown. In various species the disc of
the mesonotum is ornamented with a distinc-
tive pattern, best seen in fresh or dried
specimens, but usually visible to some degree
in slide-mounted material.
The legs are slender without special ar-
mature, but the fore-tibia bears apically a
small spur and tuft of modified hairs, and the
tip of the hind tibia bears an anterior spur and
2 transverse rows of modified spines. These
combs are used as grooming organs in both
sexes. Linley and Cheng (1974) gave an ex-
cellent account with Scanning Electron
Microscope photographs of the structure of the
tibial combs of Culicoides and their function as
grooming organs. The "hind tibial comb" in-
cludes only the longer spines in the distal row
and is of some value in classification. The 4th
tarsomere usually is cylindrical but in a few
groups is cordiform; the claws are small and
equal on all legs, simple in the female but divid-
ed at the apices in the male. The empodium is
vestigial.
The wings bear dense microtrichia, the size
and pigmentation of which give rise to the
characteristic pattern of dark and light spots,
and more or less abundant macrotrichia or
longer hairs. The color pattern of spots or
bands is characteristic for each species and is
of primary importance in classification;
however in some groups or species it is poorly
developed or even absent. In the males the
wing is longer and narrower than in the female,
and the color pattern is less contrasting. The
wing length is measured from the basal arculus
to the wing tip; the costa extends usually to
more than V2 the wing length; the'costal ratio
(CR) is the value obtained by dividing the
length of the costa by the wing length. There


are, with rare exceptions, 2 complete radial
cells formed by the more heavily sclerotized
radial branches; usually the 2nd or distal one is
broader and longer than the 1st which is often
slit-like. We use the Tillyard modification of
the Comstock-Needham system of wing vena-
tion, in which the branches of the anterior fork
are called M1 and M2, and those of the
posterior fork are M3+4 and Cul, from front to
back. The color of the halter is also useful in
species separation.
Abdomen: The female abdomen is relatively
stout, with the apex somewhat tapered and a
pair of small rounded cerci visible below the
9th tergum. Internally the female possesses
1-3 sclerotized spermathecae, which usually
are oval to pyriform in shape, with the slender
bases of the ducts sclerotized a short distance.
The spermathecae are joined by hyaline ducts
to a common duct, and at the juncture there
usually is a small sclerotized ring. In most
species with 2 sclerotized spermathecae there
is also a small rudimentary 3rd one. The
number and shapes of the functional sper-
mathecae and the presence or absence of a ring
are important in classification. The length of
the spermatheca is measured in the axis of the
base of the duct and includes the sclerotized
portion of the duct or neck.
The male abdomen is slender and bears ter-
minally the prominent genitalia (fig. 32b, 34)
which are of primary importance in group


i


n y).
thony).







classification and species identification. The
9th segment is in the form of an irregular
sclerotized ring consisting of the fused tergum
and sternum. The 9th tergum forms an expand-
ed plate which is convex externally and hollow-
ed out mesally, and bears the anus flanked by a
pair of membranous cerci on the ventromesal
surface. The hind corners of the 9th tergum are
frequently expanded as a pair of apicolateral
processes. The 9th sternum is much shorter
than the tergum; usually with a caudomedian
excavation. The base of the aedeagus ar-
ticulates with the lateral ends of this excava-
tion. The forceps-like genital appendages or
gonopods arise laterally at the base of the
tergum and are 2-segmented. The enlarged
basal segment or basistyle bears 2 internal pro-
cesses at the base, a mesally directed ventral
root and an anteriorly directed dorsal root, the
latter articulating directly with the base of the
paramere. The distal segment (dististyle or
clasper) is setose and slightly swollen at the
base, is slender and nearly bare distally with
an incurved point, and when not extended is
folded mesad across the 9th tergum. The
aedeagus usually is a Y-shaped structure with
a median process directed ventrocaudad, form-
ing a sclerotized support on the ventral surface
of the male genital duct. The parameres usual-
ly are a pair of selerotized internal rod-like
sclerites with knobbed bases and ventrally
directed distal points, but they are subject to
great modification in the shape and direction of
the basal knob, the middle stem, and the distal
point. In some groups of species the parameres
may fuse mesally in part or completely in a
plate-like structure. Although morphologists
suggest that it is more appropriate to use the
term paramere for the primary gonopod here
called the basistyle and dististyle, and the
term claspette for the internal sclerite here
called the paramere, we prefer to follow tradi-
tional usage.
Internal Anatomy: Details of the internal
structure of adult Culicoides are incompletely
known. Jobling (1928) included the internal
anatomy and musculature in his study of the
head and mouthparts of C. pulicaris.
Pomerantzev (1932) published a good
anatomical and histological account of the
male and female reproductive systems of C.
nubeculosus. Megahed (1956) studied the
alimentary tract of this species in detail.
Bowne and Jones (1966) published an excellent
account of the anatomy and histology of the
salivary glands of C. variipentis under normal
conditions and when infected with bluetongue
virus.


PUPA
The pupa of Culicoides (fig. 35) is the usual
type found in the Diptera Nematocera. The lar-
val exuviae are not retained on the posterior
segments of the pupa. The length ranges from
2-5 mm, and the color varies from pale
yellowish in some species, usually pale brown,
sometimes the abdomen paler; but the pupae of
some species are dark brown. The body con-
sists of a stout unsegmented cephalothorax
bearing a pair of elongate anterior respiratory
horns, and a narrower, 9-segmented abdomen
tapering to a pair of pointed caudal
apicolateral processes. The antenna, wing and
leg cases are closely appressed to the sides and
venter of the cephalothorax.
The surface of the cephalothorax bears a
definite number of seta-bearing tubercles,
whose position and development are of value in
classification, and more or less granular or
spiny development of the integument, also
characteristic of the species. The abdominal
segments also bear certain fixed tubercles of
characteristic shape and armature, and the in-
tegument also is often provided with
characteristic spinules or reticulations. The
armed and seta-bearing tubercles of the pupa
have been named by Carter et al. (1920) from
which figs. 35 a and b have been reproduced,
and their system of chaetotaxy has fortunately
been followed rather uniformly by later
workers who have made extensive descriptions
of the pupae (Lawson, 1951; Kettle and
Lawson, 1952; Wirth, 1952b; Jones, 1961b;
Linley and Kettle, 1964; Jamnback, 1965; and
Linley, 1965a, 1970a, 1970b).
Respiratory horn: The respiratory horns
arise from tubercles situated on the
antcrolateral region of the thorax, the
tubercles tapering to a short narrow stalk, at
the end of which the base of the horn ar-
ticulates freely. The main tracheal trunk of the
pupa runs through the middle of the horn, and
gives off small branches to a series of minute
round spiracular papillae, some usually occur-
ring singly along the side and the majority
clustered in a fan-like arrangement around the
apex of the horn. The surface of the horn often
bears characteristic spines or transverse
wrinkles or convolutions.
Operculum: The head region of the pupa
forms a convex anterior region, the midportion
of which lies between the arms of the molting
suture, and on eclosion of the adult separates
as an anteriorly-hinged plate called the oper-
culum (fig. 35a op). The anterior border of
the operculum bears 2 large anteromarginal
tubercles (fig. 35b, c- am), each bearing a
large articulated seta. The surface of the oper-
culum is beset with various small spinules or
papillae (fig. 35i) of value in species recogni-
tion.



















































VENTRAL LATERAL

I.a.s.m.


DORSAL
d as m. d.pm.



















N


VENTRAL


DORSAL

/ d. .s.m.\ \ o.s.m.


32 2~ A23 1 3 5 43
2 1 2 3

A M D i


i.2




d.

G


:2jS-'e :a

K-0u r

.2 ~"'. -


od.

J

1. p. m.


Q__Y:~


(


d. a. s. m.


ARBORICOLA


MELLEUS


Fig. 35. Culicoides pupa (a, b from Carter et al., 1920; c-d, f-1 from Jones, 1961b; e from Linley, 1970b) (for key to ab-
breviations see text): e, C. arboricola; f-l, C. melleus.


." P. M.


.P. IH.


V. P. '.


: :
i:



1


1

..
d
.--~ .:
i-:
I I':






Tubercles of the thorax: Two groups of
thoracic tubercles are of special taxonomic
value and are used in the descriptions. The
anterodorsal tubercles (fig. 35b, c, j ad) are a
pair of tubercles located about midway be-
tween the am tubercles of the operculum and
the base of the respiratory horn. Each tubercle
has a pair of setae which are usually very une-
qual or rarely subequal in length. The dorsal
(fig. 35b, d, g d) tubercles are a series of 5
seta-bearing tubercles on each side of the dor-
sum. They are numbered 1 to 3 from front to
back in a series of 3 near the midline, and 2
others (4 and 5) nearby but located laterad of 1
and 3. In most species, tubercles 1 to 3 are in
line and equidistant or nearly so, but in a few
species tubercles 1 and 2 are located side by
side. The relative length of the setae on
tubercles 1 and 2 provides another useful
character.
Tubercles of the abdomen: The tubercles of
abdominal segments 3-7 are well developed and
form characteristic groups which have been
given designations as shown in fig. 35b, d, e.
The lateral posteromarginal tubercles (lpm) are
the most useful and are a group of 3 seta-
bearing tubercles on each side of the segment;
the seta on the middle of these is longer and
more slender than those on the tubercle above
and below, and serves as a convenient land-
mark when examining pupal exuviae, which
are usually distorted in mounting on the slide.
The dorsal posteromarginal (dpm) tubercles
form a posterior row of 5 pairs across the dor-
sum of each segment in line with the 1pm
tubercles, and are usually much shorter and
broader with shorter setae than the Ipm
tubercles. Similarly the ventral postero-
marginals (vpm) form a row of 3 pairs of
shorter, broader tubercles across the venter of
each segment.

LARVA
Larvae of Culicoides (fig. 36g) are slender
and wormlike, 2-5 mm long when mature. The
head capsule is yellowish to brownish and the
body is translucent whitish, usually with some
subcutaneous pigment in a characteristic pat-
tern on the thoracic segments. There is a short
neck segment, 3 thoracic segments, and 9 ab-


dominal segments quite similar to those of the
thorax. The setae of the head and body are
quite inconspicuous, except that in a few
species there are groups of long erectile
perianal bristles near the apex of the last seg-
ment. Four membranous anal gills or papillae
(fig. 36f) can be extruded from the anus or
retracted into thin-walled pouches in the rec-
tum of the larva.
The head capsule (fig. 36 a-c) is comprised of
2 sclerites which are separated during molting
by the molting suture. The median dorsal
sclerite lying between the arms of the molting
suture is the frontoclypeus (fig. 36b, 214)
which makes up about 95% of the length of the
head capsule, and because dissections for the
pharyngeal apparatus often make it inconve-
nient to measure the length of the entire head
capsule, the index of the head length is given
by the measurement of the frontoclypeus in the
larval descriptions (following Jamnback,
1965). A simple, or usually double, pigmented
eyespot is located on each side of the head at
about midlength. The surface of the head bears
a definite number of setae and sensilla which
were given lettered designations by Saunders
(1924), followed by Lawson (1951), Kettle and
Lawson (1952), Linley and Kettle (1964), and
Linley (1970b). Another system of lettering
was used by Tokunaga (1937) for C. cir
cumscriptus and these letters were given
names by Wirth (1952b) for C. melleus and C.
furens, but Tokunaga's system must be aban-
doned in favor of the prior one of Saunders.
The antennae are located dorsally on each
side of the head near the anterior end between
the bases of the labrum and the mandibles. The
antennae are much reduced, and like the
labrum, maxillae, and mandibles, do not pro-
vide enough variation between species for tax-
onomic use. The arrangement of the mouth-
parts was figured in a diagram by Lawson
(1951) from which fig. 36e has been reproduced.
The pharyngeal apparatus consists of a ventral
hypopharynx and a dorsomedian epipharynx,
the latter composed of a series of usually 4
overlapping combs. The epipharynx (fig. 36d)
is of considerable taxonomic value in
Culicoides, especially the number of teeth and
the total width of the conspicuous dorsal comb
(Jamnback, 1965).

























U 8

PLJ











































Fig. 36. Culicoides larva and egg: a-d, g, ,C. melleus larva; a, head, ventral view; b, head, dorsal view; c, head, lateral
view; d, hypopharynx, ventral view; g, larva, lateral view; e, diagrammatic reconstruction of skeletal structures of
Culicoides head, capsule not shown except for strip joining dorsal ends of subgenal band; f, lateral view of extruded anal
papillae of C. nubeculosus; h, Culicoides egg (for lettering of setae and for abbreviations see text) (e,f,h redrawn from
Lawson, 1951).
epi

11, L hyp

E F H




Fig. 36. Culicoides larva and egg: a-d, g,,C. nelleus larva; a, head, ventral view; b, head, dorsal view; c, head, lateral
view; d, hypopharynx, ventral view; g, larva, lateral view; e, diagrammatic reconstruction of skeletal structures of
Culicoides head, capsule not shown except for strip joining dorsal ends of subgenal band; f, lateral view of extruded anal
papillae of C. nubeculosus; h, Culicoides egg (for lettering of setae and for abbreviations see text) (e,f,h redrawn from
Lawson, 1951).







SYSTEMATIC ACCOUNT


CLASSIFICATION

The Ceratopogonidae can be distinguished
from other Diptera, or 2-winged flies, by their
long, usually 15-segmented antenna, the
mouthparts are usually well developed and fit-
ted for piercing and blood-sucking, the body
usually is rather short and stout, the scutellum
lacks a median furrow or keel, and the wing
venation is characteristic. In the
Ceratopogonidae the radial veins usually are
thickened and shortened and the branches
form 1 or 2 radial cells on the anterior wing
margin, the posterior veins are less con-
spicuous, forming 2 forks, the anterior com-
prised of Ml and M2, and the posterior fork
made up of M3+4 and Cul. In the genus
Culicoides the wing is more or less hairy, usual-
ly with a conspicuous pattern of pale spots on a
dark background, 2 radial cells usually are pre-
sent and more or less of the same length, the
costa usually ends at slightly past midlength
of the wing, the tarsal claws are small and
equal in both sexes; there is no hairy em-
podium, and large thoracic humeral pits are
present.
Attempts to subdivide the large genus
Culicoides into subgenera have been only par-
tially successful. Root and Hoffman (1937) and
Edwards et al. (1939) first proposed division of
the genus into 2 series based on the male
genitalia and certain external features. Fox
(1948) erected the subgenus Hoffmania for 12
Neotropical species, and later (1955) proposed
Macfiella and A varitia. Khalaf (1954) made the
1st major attempt to group Culicoides species


according to their phylogenies, and recognized
4 subgenera: Culicoides s. str., Monoculicoides
Khalaf, Selfia Khalaf, and Oecacta Poey.
Vargas (1953) placed the crepuscularis group
in the new subgenus Beltranmyia; in 1960 he
proposed 5 new subgenera, Anilomyia,
Diphaeomyia, Drymodesmyia, Glaphiromnyia,
and Mataemyia, and resurrected
Haematomyidium Goeldi for some species
formerly placed in Oecacta; and in 1973 pro-
posed Wirthomyia to include 1 North
American species.
In the Old World fauna, the subgenera
Haemophoructus Macfie, Trithecoides Wirth
and Hubert, Meijerehelea Wirth and Hubert,
Pontoculicoides Remm, and Callotia Vargas
and Kremer are recognized. Most of the
subgenera are well founded and useful,
especially in the regions where their type
species occur. Difficulties arise, however, when
trying to separate the species of Culicoides s.
str. from Hoffmania in the Old World, and
assigning limits to Oecacta,
Haematomyidium, etc. At present Oecacta is
still a dumping ground for many species that
cannot be assigned to other subgenera, and
many groups such as the debilipalpis, bigut-
tatus, and piliferus groups may, when they are
better known, be assigned to additional
subgenera.
The subgeneric and group classifications of
the Florida Culicoides are outlined in table 2,
along with a summary of the mean values of
certain useful numerical characters of the
females. These subgenera and species groups
may be briefly characterized as follows:


Table 2. Systematic arrangement of Florida species of Culicoides
with mean values of certain numerical characters (females)

Wing Antennal
Legh, Costal Antennal Se y Palpal P/H
S Ratio Ratio rn Ratio Ratio

Subgenus Hoffmania
insignis 1.11 0.65 1.32 3,5,7,9,11-15 2.8 1.05
venustus 1.45 0.65 1.10 3,11-15 3.2 0.83

Subgenus A varitia
alachua 0.93 0.60 1.33 3,11-15 2.8 0.80
chiopterus 0.91 0.59 1.15 3,11-15 1.9 0.65
juddi 0.98 0.56 1.27 3,11-15 1.7 0.60
pechumani 0.63 0.57 1.11 3,11-15 1.5 0.44
pusillus 0.64 0.53 1.18 3,13-15 2.6 1.08

38








Antennal
Costal Antennal Sensory Palpal
Ratio Ratio Pattern Ratio


Subgenus Drymodesmyia
hinmani 0.73 0.63
loughnani 1.21 0.55
Subgenus Diphaeomyia
baueri 1.03 0.56
edeni 1.12 0.58
footei 0.90 0.61
haematopotus 1.15 0.60
Subgenus Oecacta
Guttipennis Group
arboricola 1.19 0.60
beckae 1.15 0.61
guttipennis 1.32 0.60
ousairani 1.21 0.57
villosipennis 1.62 0.61
Piliferus Group
bickleyi 1.07 0.60
husseyi 1.20 0.58
parapiliferus 1.15 0.59
piliferus 1.12 0.62
scanloni 0.90 0.62
snowi 1.04 0.61
testudinalis 1.04 0.60
Furens Group
barbosai 0.88 0.58
furens 0.91 0.58
stellifer 1.05 0.59
Debilipalpis Group
debilipalis 0.80 0.65
paraensis 0.78 0.59
torreyae 0.80 0.59
Biguttatus Group
biguttatus 1.30 0.64
loisae 1.00 0.60
mulrennani 0.93 0.65
spinosus 1.20 0.64
Unplaced Oecacta
floridensis 0.77 0.65
melleus 1.05 0.60
nanus 1.00 0.60
niger 1.24 0.58
tissoti 1.02 0.58
travisi 1.40 0.62
Subgenus Beltranmyia
bermudensis 0.97 0.60
crepuscularis 1.30 0.56
hollensis 1.40 0.61
knowltoni 1.35 0.58
mississippiensis 1.20 0.58
Subgenus Monoculicoides
variipennis s.s. 1.75 0.63
ssp. sonorensis 1.26 -


1.41 3,11-15
1.17 3-15

1.12 3,7-10
1.78 3,10-15
1.14 3-10
1.70 3-15


1.48 3,5,7,9,11-15 2.8
1.53 3-15 2.5
1.54 3,5,7,9,11-15 3.0
1.00 3-15 2.5
1.64 3,5,7,9,11-15 2.4

1.20 3,13-15 2.3
1.42 3,5,7,9,11-15 2.3
1.22 3,5,7,9,11,13-15 2.5
1.87 3,5,7,9-15 2.2
1.47 3,5,7,9-15 2.2
1.20 3,5,7,9,11-15 2.1
1.24 3,5,7,9,13-15 2.3

0.93 3,7-10 2.2
1.28 3,7-10 2.4
0.97 3,8-10 2.7

0.83 3,8-10 2.2
0.77 3,8-10 2.1
0.99 3,8-10 2.3

1.15 3,11-15 2.7
1.21 3,11-15 2.1
1.06 3,11-15 2.3
1.10 3,11-15 2.6

1.21 none 2.0
1.10 3,10-14 2.7
1.40 3-15 1.7
1.05 3-15 2.5
1.04 3,7-14 2.3
1.50 3-15 2.7

1.00 3,13-14 2.2
1.40 3-15 2.2
1.16 3,11-15 3.0
1.48 3-15 2.5
1.30 3,11-15 3.2

0.94 3,8-10 3.0
- 3,7-10 2.2


Wing
Length,
mm







Subgenus Hoffmania Fox
Wing with 2nd radial cell ending in a pale
spot; veins at base of mediocubital fork usually
pale-bordered in cell M4; small dark spot often
present over r-m crossvein or at end of R4 5;
female antennal sensory pattern usually
3,11-15; eyes usually contiguous; sper
mathecae 2 plus rudimentary 3rd and sclero
tized ring, ovoid with short necks.
Male genitalia rounded caudad with
apicolateral processes minute to small;
dististyle usually with rounded tip; basistyle
with ventral root absent, dorsal root short and
slender; aedeagus with basal arch low and
usually forming a transverse sclerotized bar,
distal portion with internal sclerotized peg and
apical papilla; parameres with short basal arm,
often joined in a mesal bridge, short and
slender distally with minute fringing setae.
Pupa with respiratory horn unique in having
no lateral spiracular openings and having
tracheal rings extending i2 length of horn;
operculum with short, stout spines, a unique
elongate process near posterior end; ad setae
long, subequal; caudal segment with a
V-shaped patch of spines on disc.
Larva unique in having brown head capsule;
thorax with faint purplish pigmented spots,
most abundant on prothorax, often faint; no
lateral pigmented spots.
Florida species: C. insignis Lutz, C. venustus
Hoffman.

Subgenus Avaritia Fox
Small species with broad, nearly hare wings.
Female with distal part of 2nd radial cell usual-
ly ending in a pale spot; eyes contiguous;
antennal sensory pattern usually 3,11-15; sper-
mathecae 2 plus rudimentary 3rd and sclerotiz-
ed ring, ovoid with short necks; tibial comb
with 5 spines.
Male genitalia with 9th tergum lacking
caudomedian notch, usually rounded caudad,
with apicolateral processes lacking or in form
of blunt rounded lobes; basistyle with ventral
and dorsal roots long and simple, subequal;
dististyle usually with rounded tip; aedeagus
with high basal arch, tip simple; parameres
separate, with simple anterolateral arm, short
slender body, and slender, microscopically
seLose or bare tip.
Pupa with respiratory horn stout, without
spines or transverse convolutions, lateral open-
ings not borne on protuberances; operculum
with long hair-like spines; ad setae long, sub-
equal; d tubercles 1 to 3 in line and 1 and 2
quite close together; caudal segment with a
transverse band of spines across disc.
Larva with unique small eyes made up of a
single pigmented spot on each side; head very


short; thorax unpigmented dorsally, with
lateral pigmented spots on each segment at
midlength.
Florida species: C. alachua Jamnback and
Wirth, C. chiopterus (Meigen), C. juddi
Cochrane, C. pechumani Cochrane, and C.
pusillus Lutz.

Subgenus Drymodesmuia Vargas
Female with wing hairy and conspicuously
ornamented with rounded pale spots including
pale spots straddling base of vein Ml and mid-
portion of vein M2, 2nd radial cell dark to tip;
3rd palpal segment swollen with deep pit
usually opening by a smaller pore; antenna
sensory pattern 3,11-15 and usually some on
proximal series; tmesonoLal pattern usually
with long median and shorter lateral dark
brown vittae narrowly bordered by grey
pollinosity; spermathecae 2 plus rudimentary
3rd and sclerotized ring, shapes variable; tibial
comb with 4 spines.
Male genitalia with long, slender, api-
colateral processes on 9th tergum; basistyle
with ventral and dorsal roots simple; dististyle
slender with bent tip; aedeagus usually short
and broad with low basal arch and stout basal
arms; parameres separate, short with basal
knob directed anterolaterad, stem swollen at
base without ventral lobe, apex slender, sim-
ple, and often twisted.
Immature stages not described.
Florida species: C hinmani Khalaf and C.
loughnani Edwards.

Subgenus Diphaeomyia Vargas
Female with eyes narrowly separated; wing
moderately hairy, 2nd radial cell dark to tip;
pale spot over r-m crossvein often lying entire-
ly distad of vein; pale spot straddling middle of
vein M2 and pale spot straddling or lying im-
mediately anterior to base of vein Ml; palpal
pit usually round and shallow; antennal sen-
sory pattern variable; tibial comb with 4
spines; spermathecae 2 plus rudimentary 3rd
and sclerotized ring, ovoid with long slender
necks.
Male genitalia with apicolateral processes of
9th tergum short and pointed; basistyle with
foot-shaped ventral root and simple dorsal
root; dististyle slender with bent tip; aedeagus
with broad basal arch, slender basal arms, arch
often bearing a pair of spurlike posterior pro-
cesses on shoulders; parameres separate, with
stout basal knob, slender on midportion with
long pointed tip bearing lateral fringing spines.
Pupal respiratory horn with lateral
spiracular openings on raised protuberances,
transverse convolutions on miidportion, spines
absent; d tubercles 1 to 3 not in line, 1 and 2







almost touching.
Larva with head capsule strongly tapering
anteriorly.
Florida species: C. baueri Hoffman, C. edeni
Wirth and Blanton, C footei Wirth and Jones,
and C. haematopotus Malloch.

Gullipennis Group
Usually large, dark species with hairy wing
bearing prominent pattern. Female with eyes
narrowly to broadly separated; antenna with 5
distal segments rather elongated; sensory pat-
tern 3,11-15 with sensoria also present on some
of proximal segments; 3rd palpal segment
usually moderately sdvollen with definite pit;
mesonotum usually with prominent pollinose
pattern; legs usually dark with prominent
banding; tibial comb usually with 5 spines;
spermathecae 2 plus rudimentary 3rd and
sclerotized ring, ovoid with short necks. Wing
pattern with 2nd radial cell dark to tip; pale
spots straddling midportions of veins M1 and
M2, apices of veins Ml, M2, M3+4, and
sometimes Cul pale-margined; anal cell usually
with 3 separate pale spots arranged in a
triangle distad of the proximal pale area.
Male genitalia with prominent, usually
stout, apicolateral processes on 9th tergum;
basistyle with simple ventral and dorsal roots;
dististyle slender and tip bent except in C. gut-
tipennis; aedeagus with basal arch usually ex-
tending to /2 of total length, distal portion
usually slender with specific apical modifica-
tions; parameres separate, with strong basal
knob, basal constriction on stem, latter usually
sinuate, swollen proximally and tapering to
simple distal point bent or curved ventrad.
Pupa respiratory horn with surface at least
in part with appressed broad scaly spines;
operculum densely covered with short stout
spines; a setae very unequal; d tubercles 1 to 3
in line with I and 2 closer than 2 and 3; all ab
dominal segments with dense scalelike con-
fluent spines giving reticulated appearance.
Larva with head long and narrow; thorax un-
pigmented dorsally, with faint lateral
pigmented spots present or absent on meso-
and metathorax; last segment with long
perianal bristles.
Florida species: C. arboricola Root and Hoff-
man, C. beckae Wirth and Blanton, C. gut-
tipennis (Coquillett), C. ousairani Khalaf, and
C. villosipennis Root and Hoffman.

Piliferus Group
Moderate size grayish to dark species with
hairy wing bearing variable pattern. Female
with eyes contiguous to moderately separated;
antennal sensory pattern usually 3,5,7,9,11-15,


but often with less or more sensoria; palpus
usually with distinct round pit; wing pattern
typically with more or less distinct pale spots
straddling base of vein M1 and midportion of
vein M2, second radial cell dark to tip, rarely
pale at apex; distal pale spot in cell R5 when
present elongate and more or less filling distal
portion of cell; tibal comb with 4-5 spines; sper-
mathecae 2 plus rudimentary 3rd and sclero
tized ring, spermathecae usually oval without
necks.
Male genitalia with moderately long, pointed
apicolateral processes on 9th tergum; basistyle
with ventral root foot-shaped, dorsal root
slender; aedeagus with basal arms slender,
basal arch rounded, tip simple, usually slender;
parameres separate, with round basal knob,
stem slender, sinuate, with at most slight in-
dication of a ventral swelling; apex tapering to
fine tip with lateral fringe of fine spines.
Pupa with respiratory horn lacking
transverse convolutions but bearing some
sharp spines, lateral spiracular openings not on
raised protuberances; operculum with spines

papillate, but some with short spines on disc;
ad setae very unequal; d tubercles 1 to 3 in line,
1 and 2 closer than 2 and 3; caudal segment
with or without spines on disc.
Larva with thorax unpigmented dorsally ex-
cept for transverse bars sometimes present
near anterior segment margins with well
developed lateral spots, especially on meso-and
metathoracic segments.
Florida species: C. bickleyi Wirth and
Hubert, C. husseyi Wirth and Blanton, C.
parapiliferus Wirth and Blanton, C. piliferus
Root and Hoffman, C. scanloni Wirth and
Hubert, C. snowi Wirth and Jones, and C.
testudinalis Wirth and Hubert.

Furens Group
Small to moderate size species with promi-
nent thoracic and wing pattern. Female with
eyes moderately separated; antennal sensory
pattern 3, 7-10 or 3,8-10; 3rd palpal segment
moderately slender with round shallow pit;
wing moderately hairy; 2nd radial cell dark to
tip; wing pattern variable; cell M2 with 2 pale
spots in distal portion; tibal comb with 4
spines; spermathecae 2 plus rudimentary 3rd
and sclerotized ring, ovoid with long slender
necks.
Male genitalia with well developed
apicolateral processes; basistyle with foot-
shaped ventral root; aedeagus with high,
rounded, basal arch and slender basal arms,
distal process slender with variable tip;
parameres separate, with stout basal knob,
stem slender and sinuate, often with distinct







ventral lobe, apex slender with distal fringing
spines.
Pupa with respiratory horn bearing
transverse convolutions in midportion, lateral
spiracular openings on distinct raised pro-
tuberances; operculum with short to moderate-
ly long spines; ad setae very unequal; d
tubercles 1 to 3 in line (C. stellifer) or not
aligned.
Larva with thoracic pigmentation varying
from extensive over most of dorsum to faint
lateral spots only.
Florida species: C. barbosai Wirth and Blan-
ton, C. furens (Poey), and C. stellifer (Co-
quillett).

Debilipalpis Group
Small dark brown species with small,
definite wing markings. Female with eyes nar-
rowly to broadly separated; antenna sensory
pattern 3,8-10; wing moderately hairy, 2nd
radial cell dark to tip, with pattern of small
round pale spots usually not meeting wing
margin; no pale spots straddling veins Ml and
M2, cell M2 with 1 pale spot in distal portion;
tibial comb with 4 spines; spermathecae 2 plus
rudimentary 3rd and sclerotized ring, ovoid
with long slender necks.
Male genitalia with well developed
spicolateral processes on 9th tergumi basistyle
with footshaped ventral root; aedeagus with
high basal arch and slender basal arms, distal
process short and tapering with tip simple or
modified; parameres separate, with stout basal

without ventral lobe, apex slender and pointed
with distal fringing spines.
Pupa (of Neotropical C. hoffmani Fox) with
respiratory horn bearing coarse spines,
without transverse convolutions, lateral
spiracular openings on protuberances; oper-
culum with short blunt spines on margins and
a median band; abdominal segments with
dense spinules.
Larva (of Neotropical C. hoffmani) with 8
long perianal bristles.
Florida species: C. debilipalpis Lutz, C.
paraensis (Goeldi), and C. torreyae Wirth and
Blanton.

Bigullalus Group
Small to moderate size species with poorly
marked, usually hairy wing. Female with eyes
narrowly separated to contiguous; antenna
with sensory pattern usually 3,11-15,
segments usually slender; palpus usually with
sensory pit irregular; wing with 2nd radial cell
usually dark to tip, sometimes slightly pale at
apex, pale markings definite over r-m crossvein
and past tip of 2nd radial cell, indistinct on


posterior and distal portion of wing if present;
tibial comb usually with 4 spines; sper-
mathecae 2 plus rudimentary 3rd, sclerotized
ring present or absent; spermathecae ovoid
with short necks.
Male genitalia with prominent apicolateral
processes; basistyle with ventral and dorsal
roots simple and slender, subequal; aedeagus
with high basal arch, distal portion often
rather stout; parameres separate, with base
bent at right angles to stem, the latter short,
simple, with apex rather stout and simple, or
crowned with short, sharp spines.
Pupa with respiratory horn rather stout,
with a few spines, transverse convolutions ab-
sent, lateral spiracular openings not on raised
protuberances; opercular spines short; ad setae
very unequal; d tubercles 1 to 3 in line, nearly
equidistant; caudal segment with or without
spines on disc.
Larva (of C. biguttatus) with dorsum of
thorax unpigmented, with faint lateral
thoracic spots.
Florida species: C. biguttatus (Coquillett), C.
loisae Jamnback, C. mulrennani Beck, and C.
spinosus Root and Hoffman.

Subgenus Beltranmyia Vargas
Female with eyes narrowly to widely
separated; antenna sensory pattern usually
3-15 or 3,11-15, often variable; palpus usually

prominent pattern; wing usually hairy, pattern
when present of large round to oval pale spots
centering between the veins, no pale spots
straddling veins Ml or M2; tibial comb with

rudimentary spermatheca and sclerotized ring
absent.
Male genitalia with well developed api-
colateral processes; ventral root of basistyle
absent, dorsal root short; aedeagus with high
rounded basal arch, tip usually slender and
simple; parameres usually separate, basal knob
small with distinct anterior process, stem
usually slender, tapering to fine, abruptly bent,
simple tip.
Pupa with respiratory horn without
transverse convolutions, surface with spines
(crepuscularis) or bare, lateral spiracular open
ings not on protuberances; opercular spines
short to moderately long; ad setae very une-
qual (except both long and equal in hollensis); d
tubercles 1 to 3 in line, 1 and 2 closer than 2
and 3; caudal segment with or without spines
on disc.
Larva with dorsum of prothoracic segment
largely covered with pigmentation, meso- and
metathoracic segments to a lesser extent;
meso- and metathorax with more heavily







pigmented round lateral spots.
Florida species: C bermudensis Willams, C.
crepuscularis Malloch, C. hollensis (Melander
and Brues), C. knowltoni Beck, and C.
mississippiensis Hoffman.

Subgenus Monocalicoides Khalaf

Large, stout-bodied species with 2nd radial
cell dark to tip and macrotrichia moderately
abundant. Female with eyes broadly
separated; antennal sensory pattern 3,8-10 or
sometimes 3-10; frontal tubercles present be-
tween frontal carina and mesal margins of 1st
antennal segment; mesonotum usually with
numerous small dark punctures at bases of the
hairs; wing pattern usually with pale and dark
streaks instead of round pale spots; tibial comb
usually with 5-7 spines; spermatheca 1, shape
various, frequently arcuate, opening to duct


large, rudimentary spermatheca and sclerotiz-
ed ring absent.
Male genitalia with prominent apicolateral
processes; basistyle with short, simple ventral
root, dorsal root long and slender; aedeagus
with short, broad basal arch and tip notched or
bifid; parameres broadly fused basally, with
slender distal processes.
Pupa with respiratory horn bearing pro-
nounced transverse wrinkles, lateral spiracular
openings not on protuberances; operculum
densely covered with short spines except on
posterior '/; ad setae very unequal; d tubercles
1 to 3 in line, equidistant; caudal segment
without patch of spines on disc.
Larva with head strongly tapering anterior-
ly; pharyngeal sclerites uniquely massive;
thorax with dorsal surface covered with red-
dish brown mottled pigmentation.
Florida species: C. variipennis (Coquillett).


KEYS TO SPECIES OF FLORIDA CULICOIDES


KEY TO ADULT FEMALES
1. One spermatheca present (fig. 47e) 2
1'. Two functional sclerotized sper-
mathecae present (fig. 38f) 7
2(1). Spermatheca elongate and C-shaped
(fig. 125f); wing with pattern of distinc-
tive gray streaks (fig. 125d); antenna
without sensoria on segments 11-15
(fig. 125a) ...... .
. variipennis (Coquillett), p. 161
2'. Spermatheca oval (fig. 73e) or if
elongate is straight (fig. 81e); wing pat-
tern of rounded pale spots (fig. 55c) or
pattern faint or absent; antenna with
sensoria present on some or all of
segments 11-15 (fig. 89a) ......... 3
3(2). Spermatheca elongate and saclike, at
least 4 times as long as wide (fig. 81e) ..
.... knowltoniBeck, p. 110
3'. Spermatheca oval, not more than twice
as long as wide (fig. 55e) 4
4(3). Wing with conspicuous pattern of oval
or rounded pale spots (fig. 55c) 5
4'. Wing without pattern or with indistinct
pale spots on distalportion (fig. 73c) 6
5(4). Second radial cell dark to tip (fig. 55c);
antennal sensory pattern 3-15 (fig. 55a)
.. crepuscularis Malloch, p. 75
5'. Second radial cell pale at tip (fig. 89c);
antennal sensory pattern 3,11-15 (fig.
89a) mississippiensis Hoffman, p. 119


6(4). Spermatheca lightly sclerotized (fig.
47e); mandible with 7 vestigial teeth;
small species with narrow wing, 0.97
mm long bermudensis Williams, p. 65
6'. Spermatheca strongly sclerotized (fig.
73e); mandible with 12-15 teeth; larger
species with broader wing. 1.00-1.40
mm long .
hollensis (Melander and Brues), p. 101
7(1). Wing with extensive pattern of inter-
connected, distinct, pale spots; veins
Ml, M2, and M3+4 pale-margined
nearly to bases; spermathecae elongate,
saclike, with large openings to ducts
........ .loughnani Edwards, p.114
7'. Wing patternenot soextensive, almost
with separate pale spots and pale
margins of veins Ml, M2, M3+4, if pre-
sent, do not extend nearly to bases;
spermathecae oval or ovoid, not saclike,
usually with slender necks ....... 8
8(7). Second radial cell with distal portion in
a pale area ....... ............ 9
8'. Second radial cell dark to tip .. 16
9(8). Cell M4 with pale lines bordering veins
M3+4 and Cul at base of fork ..... 10
9'. Cell M4 dark along veins at base of fork
. . . . . 1 1
10(9). Cell Ml with 2 pale spots in distal por-
tion; r-m crossvein not bearing an
isolated dark spot . ......
...... venustus Hoffman, p. 166







10'. Cell M1 with 1 pale spot in distal por-
tion; rm crossvein bearing an isolated
dark spot insignis Lutz, p. 106
11(9). Wing with extensive macrotrichia ex-
tending to base; costa longer (CR 0.64);
palpus with irregular shallow pit ....
spinosus Root and
Hoffman (part), p. 146
11'. Wing with sparse macrotrichia on
distal portion only; costa shorter (CR
0.53-0.60); palpus with deep round pit.
. . . . . . . 1 2
12(11). Small species, wing 0.64 mm long; wing
without macrotrichia; legs pale; anten-
nal sensory pattern various .... 13
12'. Larger species, wing 0.91-0.98 mm long;
wing with some macrotrichia near tip;
legs with dark markings; antennal sen-
sory pattern 3,11-15 ...... 14
13(12). Mandible with 14 distinct teeth; eyes
hairy; antennal sensory pattern 3,13-15
pusillus Lutz, p. 140
13'. Mandible without teeth; eyes bare;
antenna sensory pattern 3,11-15 .
......... pechumani Cochrane, p. 136
14(12). Mandible without teeth ......
.. ...... juddi Cochrane, p. 109
14'. Mandiblewith 6-13 teeth 15
15(14). Proboscis short, P/H Ratio 0.65; 3rd
palpal segment short, PR 1.9; mandible
with 6-9 teeth; wing with inconspicuous
markings .. .chiopterus (Meigen), p. 72
15'. Proboscis long, P/H Ratio 0.80; 3rd
palpal segment long, PR 2.8; mandible
with 13 teeth; wing with more con-
spicuous markings .....
... alachua Jamnback and Wirth, p. 54
16(8). Wing with pale spot straddling midpor-
tionofvein M2 ... .. .. 17
16'. Wing without pale spot straddling mid-
portion of vein M2 ...... ... 29
17(16). Anal cell with 3 prominent pale spots
arranged in a triangle in addition to any
pale area at base of cell; apices of veins
M1, M2, and M3+4 with pale spot at
wingmargin .. .. ........ 18
17'. Anal cell with 1 or 2 pale spots in distal
portion; apices of veins M1, M2, and
M3+4paleordark ...... 22
18(17). Apex of vein Cl pale at wing margin .
. arboricola Root and Hoffman, p. 56
18'. Apex of vein Cul dark ... ...... 19
19(18). Mesonotum without pattern of
pollinose gray patches dn anterior por-
tion; palpal pit deep; wing markings
restricted, pale spot over r-m crossvein


not extending into cell M2
..... ousairani Khalaf, p. 129
19'. Mesonotum with prominent pattern of
pollinose gray patches from anterior to
posterior margins; palpal pit shallow;
wing markings various 20
20(19). Wing markings extensive, pale spot
over rm crossvein extending caudad
well into cell M2; spermathecae
pyriform with swollen tapering necks;
antenna sensory pattern 3,5,7,9,11-15
. guttipennis (Coquillett), p. 92
20'. Wing markings restricted, pale spot
over r-m crossvein not extending into
cell M2; spermathecae ovoid with
slender necks; sensory pattern various
. . . . . . . 2 1
21(20). Hind femur with prominent subapical
pale ring; halter dark; antenna with sen-
sory pattern 3-15; distal segments not
greatly elongated, AR 1.53 ..... .
beckae Wirth and Blanton, p. 63
21'. Hind femur without subapical pale
ring; halter pale; antenna with sensory
pattern 3,5,7,9,11-15; distal segments
greatly elongated, AR 1.64 .......
villosipennis Root and Hoffman, p. 169
22(17). Cell R5 with a double transverse pale
spot in distal portion; veins Ml, M2,
M3+4, and Cul pale-margined nearly
their entire lengths; antennal sensory
pattern 3,7-10 baueri Hoffman, p. 61
22'. Cell R5 with a single round or elongate
pale spot in distal portion, at or near
wing tip; veins Ml, M2, M3+4, and Cul
dark (Ml with pale spot at wing tip in
C. hinmani); antennal sensory pattern
otherwise .. .... 23
23(22). Spermathecae with distinct necks; cell
R5 with small round pale spot in distal
portion ...... ..... 24
23'. Spermathecae without distinct necks;
cell R5 large, indefinite pale area filling
distal portion ......... 26
24(23). Pale spot straddling vein M1 at prox-
imal 1A of its length; distal pale spot in
cell R5 located at extreme tip of cell;
antenna sensory pattern 3-15 varying
to 3,11-15; spermathecae with long
slender necks; pale spot over r-m cross-
vein lies entirely distad of vein .... 25
24'. Vein M1 without straddling pale spot
at proximal '/A; distal pale spot in cell
R5 located subapically in cell; antenna
sensory pattern 3,11-15; spermathecae
with stout tapering necks; pale spot
over r-m crossvein lies centered on the
vein ..... ... hinmani Khalaf, p. 99







25(24). Body and legs dark brown with paler
markings; pale spot over r-m crossvein
extends anteriorly to coastal margin;
anal cell with proximal pale spot pres-
ent near wing margin.
....... haematopotus Malloch, p. 95
25'. Body and legs yellowish brown, legs
without dark brown bands; pale spot
over r-m crossvein not extending
anteriorly to costal margin; anal cell
without proximal pale spot near wing
m argin ....... ...... .
.... .. edeni Wirth and Blanton, p. 81
26(23). Antennal sensory pattern 3,13-15 . .
.... bickleyi Wirth and Hubert, p. 67
26'. Antennal sensory pattern 3,5,7,9-15
.. 27
27(26). Eyes broadly separated (fig. 101); pro-
boscis longer, P/H Ratio 0.92; dark
brown species .
parapiliferus Wirth and Blanton, p. 134
27'. Eyes narrowly to moderately separated
(fig. 103); proboscis shorter, P/H Ratio
0.71 to 0.78; pale brown species. 28
28(27). Proximal antennal segments short,
moniliform, AR 1.87; 3rd palpal seg-
ment greatly swollen; eyes narrowly
separated ........ . .
.. piliferus Root and Hoffman, p. 137
28'. Proximal antennal segments more
elongate, AR 1.47; 3rd palpal segment
moderately swollen; eyes moderately
separated .. ............... ..
.... scanloni Wirth and Hubert, p. 142
29(16). Wing with conspicuous pattern in-
cluding definite pale spots in distal por-
tions of cells R5 and M ... 30
29'. Wing without pale spots, or with 2 pale
spots, over r-m crossvein and at end of
2nd radial cell. and at most indistinct
pale marks in apices of cells R5 and Ml
. . . . . . 3 5
30(29). Cell M1 with 3 pale spots, the distal one
at wing margin rarely indistinct ... 31
30'. Cell Ml with only 2 pale spots ..... 33
31(30). Cell M2 and anal cell each with 2 pale
spots in distal portion; subapical pale
spot in cell R5 double or with anterior
side meeting anterior wing margin; 3rd
palpal segment with shallow or ir-
regular sensory pit; mesonotum with
prominent pattern ........... 32
31'. Cell M2 and anal cell each with 1 pale
spot in distal portion; subapical pale
spot in cell R5 rounded andnot meeting
anterior wing margin; 3rd palpal seg-
ment with small deep sensory pit;


mesonotum without prominent pattern
... paraensis (Goeldi), p. 131
32(31). Cell R5 with small round pale spot be-
tween poststigmatic pale spots and
distal pale spot; mesonotum with pat
tern of punctiform brown dots; anten-
nal sensory pattern 3,7-10 ..
furens (Poey), p. 87
32'. Cell R5 without small round pale spot
between poststigmaLic pale spots and
distal pale spot; mesonotum with pat-
tern of pollinose gray patches; antenna
sensory pattern 3,8-10 .........
. ... stellifer (Coquillett), p. 149
33(30). Cell M2 with 2 pale spots between level
of mediocubital fork and wing margin;
mesonotum with prominent pattern of
punctiform brown dots .........
barbosai Wirth and Blanton, p. 58
33'. Cell M2 with 1 pale spot between level
of mediocubital fork and wing margin;
mesonotum without prominent pattern
S. .. . . . . 3 4
34(33). Third palpal segment with deep sensory
pit opening by a smaller pore
. .. debilipalpis Lutz, p. 78
34'. Third palpal segment with large, ir-
regular sensory pit ............
... torreyae Wirth and Blanton, p. 156
35(29). Yellowish species; wing without pat-
tern ....... ... .... 36
35'. Brownish to blackish species; wing with
or without faint pattern ...... 37
36(35). Mesonotum with grayish dusting;
larger, stouter species, wing 1.05 mm
long; antennal sensory pattern 3,10-14;
palpus with scattered sensilla; sper-
mathecae dark brown
.. .. melleus (Coquillett), p. 116
36'. Mesonotum shining pale yellow;
smaller slender species, wing 0.77 mm
long; antenna without sensoria; palpus
with definite round sensory pit; sper-
mathecae pale yellowish
.. ... . floridensis Beck, p. 83
37(35). Wing whitish hyaline with whitish
macrotrichia; radial veins forming a
conspicuous blackish stigma; body
blackish; legs dark brown without pale
bands ...... .
Stissoti Wirth and Blanton, p. 154
37'. Wing grayish, at least some
macrotrichia dark, without blackish
stigma, pattern often present; legs
usually with basal bands on tibiae 38
38(37). Wing uniformly grayish, without pale
spots; mandible without teeth; antennal

15







sensory pattern 3,11-15; spermathecae
ovoid with tapering necks, sclerotized
ringabsent .loisaeJamnback, p. 112
38'. Wing with pale spots at least over r-m
crossvein and at end of 2nd radial cell;
mandible with distinct teeth ... 39
39(38). Wing with pale spots present only over
r-m crossvein and at end of 2nd radial
cell .. .. .... 40
39'. Wing with at least faint pale spots pres-
ent in addition to the two over r-m
crossvein and at end of 2nd radial cell..
. .. . . . . 4 4
40(39). Third palpal segment with irregular
sensory pit; antennal sensory pattern
usually 3,11-15; spermathecae with
tapering necks but without sclerotized
ring .. ... .. 41
40'. Third palpal segment with definite
round sensory pit; antennal sensory
pattern 3,5,7.9.11-15 usually; sper-
mathecae without necks but with
sclerotized ring present ....... 42
41(40). Body dark brown; wing with numerous
macrotrichia; spermathecae with stout
tapering necks; proboscis long, P/H
Ratio 0.85; larger species, wing length
1.30mm .. ....... ..
biguttatus (Coquillett), p. 70
41'. Body yellowish brown; wing with few
macrotrichia; spermathecae with
slender tapering necks; proboscis short,
P/H Ratio 0.63; smaller species, wing
length 0.93 mm ...... .. .. .
..... mulrennani Beck, p. 122
42(40). Wing creamy white; halter pale. ..
S.. snowi Wirth and Jones, p. 144
42'. Wing dark grayish hyaline; halter in-
fuscated .. .. ... ... .. 43
43(42). Eyes narrowly separated; larger
species, wing 1.20 mm long; wing
darker, macrotrichia coarser
Shusseyi Wirth and Blanton, p. 104
43'. Eyes broadly separated; smaller
species, wing 1.04 mm long; wing paler,
macrotrichia finer .. ...........
testudinalis Wirth and Hubert, p. 152
44(39). Third palpal segment short and greatly
swollen, with a deep sensory pit open-
ing by a smaller pore ........ 45
44'. Third palpal segment longer and
moderately swollen, with a shallow
round or irregular sensory pit ..... 46
45(44). Wing with distinct pale spots around
the margin; antennal sensory pattern
3-15; spermathecae without necks ..
.. .nanus Root and Hoffman, p. 124


45'. Wing without pale spots in apices of
cells R5 and Ml; antenna sensory pat-
tern 3-10; spermathecae with long
slender necks .
..... footei Wirth and Jones, p. 85
46(44). Third palpal segment with irregular
sensory pit; antennal sensory pattern 3,
11-15; spermathecae ovoid, tapering to
slender necks, sclerotized ring absent .
spinosus Root and
Hoffman (part), p. 146
46'. Third palpal segment with definite
round sensory pit; antennal sensory
pattern 3-15; spermathecae oval
without necks . ....... . 47
47(46). Pale spots present in apices of cells R5,
Ml, and M2; body yellowish brown;
distal antennal segments long and
slender, AR 1.50 ..........
....... travisi Vargas, p. 159
47'. Pale spots absent in apices of cells R5,
Ml, and M2; a broad characteristic
band of pale spots from end of 2nd
radial cell to cell M4; body dark brown;
distal antennal segments shorter, AR
1.05 niger Root and Hoffman, p. 126


KEY TO MALE GENITALIA'

1. Ninth tergum rounded caudad, with
apicolateral processes absent or in-
conspicuous 2
1'. Ninth tergum with conspicuous api-
colateral processes or lobes ........ 4
2(1). Ninth tergum with apicolateral pro-
cesses absent; aedeagus with a high
basal arch without transverse sclero-
tized bar, conspicuous subapical ven-
tral lobe, the distomedian process in-
conspicuous, internal sclerotized peg
absent ................. ...
alachua Jamnback and Wirth, p. 54
2'. Ninth tergum with distincL apicolateral
processes present; aedeagus with basal
arch low, in form of sclerotized
transverse bar, distomadian process
long and slender with papilliform tip,
ventral lobe absent, internal sclerotized
subapical peg present ... .. 3
3(2). Apicolateral processes widely spaced,
longer and slender; parameres joined by
basalbridge.... insignis Lutz, p. 106
3'. Apicolateral processes approximated,
short and papilliform; parameres
separate ... venustus Hoffman, p. 166

'Male of pechumani unknown.







4(2). Parameres broadly fused in a triangular
plate; aedeagus with deeply bifid
distomedian process; apicolateral pro-
cesses prominent, slender and di-
vergent
.. variipennis (Coquillett), p. 161
4'. Parameres separate or joined by a
slender, inconspicuous basal bridge;
aedeagus with distomedian process not
deeply cleft; apicolateral processes
various . .. .. . . 5
5(4). Parameres with apical crown of spines
or bare distally .. .... 6.
5'. Parameres with distal fringing spines.
27
6(5). Parameres with apical crown of 4-6
sharp spines 7..... ...
6'. Parameres bare distally .. ..... 8
7(6). Apicolateral processes long, would
touch if directed medially ...
spinosus Root and Hoffman, p. 146
7'. Apicolateral processes shorter, would
not touch if directed medially ....
loisae Jamnback,p. 112
8(6). Aedeagus with internal sclerotized
point directed cephalad at juncture of
basal arms; apicolateral processes in
form of low rounded lobe or blunt pro-
cess .. .. ... ....... 9
8'. Aedeagus without internal sclerotized
point at juncture of basal arms;
apicolateral processes bluntly or sharp-
ly pointed, usually elongate 10
9(8). Apicolateral processes in shape of low
rounded lobe; anterolateral arm of
paramere long and slender.
..... . pusillus Lutz, p. 140
9'. Apicolateral processes in shape of blunt
point; anterolateral arm of paramere
short and stout .......
chiopterus (Meigen), p. 72
10(8). Genitalia massive; dististyle abruptly
bent in midportion ......
Sguttipennis (Coquillett), p. 92
10'. Genitalia not massive, of normal size;
dististyle straight or curved in midpor-
tion ......... .. ........ 11
11(10).Ventral root of basistyle absent or
much shorter than dorsal root; basal
knob of paramere with distinct anterior
process; ventral membrane of 9th ster-
numspicolate ... .. 12
11'. Ventral and dorsal roots of basistyle
long and slender, subequal in length;
basal knob of paramere with or without
anterior process; ventral membrane of
9th sternum not spiculate .... 16


12(11). Parameres joined at base by a slender
sclerotized or membranous bridge 13
12'. Parameres separate .. . 14
13(12). Apicolateral processes short and
pointed; parameres with bridge
sclerotized, anterior process long, distal
portion shorter .. .........
.. knowltoni Beck, p. 110
13'. Apicolateral processes long and
slender; parameres with bridge mem-
branous, anterior process short, distal
portion longer .......
Shollensis (Melander and Brues), p. 101
14(12). Aedeagus with apex broad, bearing
strong lateral sclerotization forming a
distinct ventral channel ...... .....
... ... bermudensis Williams, p. 65
14'. Aedeagus with apex moderately
slender, without lateral sclerotization..
. . . . . . .. . ... 1 5
15(14). Gulf coast species in saltmarshes; refer
to fem ales ........ ..... ...
...I. mississippiensis Hoffman, p. 119
15'. Widespread species in freshwater: refer
to fem ales ..... ..... ......
....... crepuscularis Malloch, p. 75
16(11). Parameres with arm bearing basal knob
abruptly turned laterad at nearly right
angles to stem ..... ........ 17
16'. Parameres with arm bearing basal knob
extending obliquely anterolaterad 22
17(16). Apicolateral processes of 9th tergum
stout at bases or widely diverging;
parameres relatively short and stout
with apices bent laterad 18
17'. Apicolateral processes of 9th tergum
slender and not diverging; parameres
longer and slender with twisted tips .
S. . . . . . . . . . 2 0
18(17). Apicolateral processes widely diverg-
ing ......... ..... .. 19
18'. Apicolateral processes stout at bases,
not widely diverging, with deep mesal
notch between them; (aedeagus and
parameres moderately stout distally) .
biguttatus (Coquillett), p. 70
19(18). Aedeagus with tip broad and truncate;
parameres with expanded tips .....
... melleus (Coquillett), p. 116
19'. Aedeagus with distal portion tapering
to slender rounded tip .. ...
.... .. mulrennani Beck, p. 122
20(17). Aedeagus with tip elongate with sub-
parallel sides .... travisi Vargas, p. 159
20'. Aedeagus with distal portion tapering
to slender rounded tip ... 21






21(20). Aedeagus with basal arch entending to
0.4 of total length .
hinmani Khalaf, p.99
21'. Aedeagus with basal arch extending to
0.6 of total length
nanus Root and Hoffman, p. 124
22(16). Aedeagus with basal arch entending to
0.4 of total length; apicolateral pro-
cesses long and slender, would meet if
directed medially .
... loughnani Edwards, p. 114
22'. Aedeagus with basal arch intending to
0.5 or more; apicolateral processes
shorter and stouter .. .. 23
23(22).Parameres with constricted portion
near basal knob short, swollen midpor-
tion longer, sinuate and gradually
tapering to sharp distal point; aedeagus
tapering to slender distal point with or
without subapicalprocesses .. 24
23'. Parameres with constricted basal por-
tion rather extensive, swollen midpor-
tion short and bulbous, abruptly nar-
rowed to slender, elongate distal blade
niger Root and Hoffman, p. 126
tissoti Wirth and Blanton. p. 154
24(23). Aedeagus with 2 pairs of short, lateral
hyaline processes subapically .
villosipennis Root and Hoffman, p. 169
24'. Aedeagus with simple Lip .. 25
25(24). Paramere short and greatly swollen in
midportion .. ousairani Khalaf, p. 129
25'. Paramere more elongate, moderately
swollen in midportion 26
26(25). Paramere swollen on greater portion,
the slender tip not so long ....
arboricola Root and Hoffman, p. 56
26'. Paramere swollen less distally, the
slender tip longer .....
S. beckae Wirth and Blanton, p. 63
27(5). Aedeagus with a pair of long sclerotized
processes on shoulders of basal arch .
28
27'. Aedeagus without sclerotized processes
on shoulders of basal arch ... 29
28(27). Aedeagus with distal process tapering
to tip; parameres without ventral lobe,
distal portion not expanded
baueri Hoffman, p. 61
28'. Aedeagus with sides of distal process
subparallel; parameres each with
distinct ventral lobe and expanded, leaf-
like distal blade; (separate on wing pat
tern, see figures)
Shaematopotus Malloch, p. 95
S footei Wirth and Jones, p. 85
edeni Wirth and Blanton, p. 81


29(27). Parameres each with swollen ventral
lobe on distal portion . .. 30
29'. Parameres without ventral lobe or
swelling on distal portion (piliferus
group; separation difficult on color and
wing pattern)
bickleyi Wirth and Hubert, p. 67
husseyi Wirth and Blanton, p. 104
parapliferus Wirth and
Blanton, p. 134
piliferus Root and Hoffman, p. 137
scanloni Wirth and Hubert, p. 142
snowi Wirth and Jones, p. 144
testudinalis Wirthand Hubert, p. 152
30(29). Ventral lobe of parameres in form of a
low swelling .. .... . 31
30'. Ventral lobe of parameres in form of
pouch-like expansion .... 33
31(30). Paramere with ventral swelling slight,
often indistinct; lateral spines in com-
pact group, distal point stouter
.. stellifer (Coquillett), p. 149
31'. Paramere with ventral swelling low and
broad ..... 32
32(31). Apicolateral processes of 9th tergum
long, slender and divergent
.. ..... furens (Poey), p. 87
32'. Apicolateral processes of 9th tergum
short and pointed ...... . .
barbosai Wirth and Blanton, p. 58
33(30). Basal arch of aedeagus extending to V2
of total length; parameres with base of
stem short and curved, stouter .... 34
33'. Basal arch of aedeagus extending to %
of total length; paramere with base of
stem long, straight and slender .....
I debilipalpis Lutz, p. 78
paraensis (Goeldi), p. 131
34(33). Aedeagus with distal process short and
stout ..... .. .. ..
torreyae Wirth and Blanton, p. 156
34'. Aedeagus with distal process long and
tapering to slender tip ...
.. .floridensis Beck, p. 83


KEY TO KNOWN PUPAE OF
FLORIDA CULICOIDES

1. Respiratory horn without lateral
spiracular openings; operculum with
elongate, fingerlike process on midline
near posterior margin .... 2
1'. Respiratory horn with lateral spir-
acular openings; operculum without
process on midline near posterior
margin ...... .. ...... .. 3







2(1). Respiratory horn elongate, L/W Ratio
10.0, with 8-10 apical spiracular open-
ings; operculum with numerous dark
spines covering disc; caudal segment
without patch of spines on disc or on
apicolateral processes .
insigniss Lutz, p. 106
2'. Respiratory horn shorter, L/W Ratio
7.4, 11-13 apical spiracular openings;
operculum with spines confined to
lateral margins; caudal segment with
patch of spines on disc and spines pres-
ent on apicolateral processes ......
.......... venustus Hoffman, p. 166
3(1). Operculum with long hairlike spines,
about as long as maximum width of
operculum .chiopterus (Meigen), p. 72
3'. Operculum with spines or papillae not
more than I& as long as maximum
width of operculum 4
4(3). Respiratory horn with lateral spiracular
openings borne on prominent pro-
tuberances; surface of horn not
spiny . .. ... ... 5
4'. Respiratory horn with lateral spir-
acular openings at most on slightly
swollen areas; surface of horn spiny or
not spiny .. .. .. .. 14
5(4). Respiratory horn without definite
transverse wrinkles or convolutions in
midportion ... .. .. 6
5'. Respiratory horn with transverse con-
volutions in midportion ........... 7
6(5). Respiratory horn long and slender, L/W
Ratio 10.0; Ipm tubercles elongate, con-
ical ... nanus Root and Hoffman, p. 124
6'. Respiratory horn short and stout, L/W
Ratio 5.3; Ipm tubercles rounded or
blunt-pointed ........... ... . ...
hollensis (Melander and Brues), p. 101
7(5). Respiratory horn with 15-20 apical and
4-7 lateral spiracular openings, all
lateral openings on a single pro-
tuberance near base
.. melleus (Coquillett), p. 116
7'. Respiratory horn with not more than 8
apical and 4 lateral spiracular openings,
lateral ones each on a separate pro-
tuberance .........8
8(7). Dorsal d tubercles 1 and 2 side by side
and contiguous.. 9
8'. Dorsal d tubercles 1, 2, and 3 in line or
out of line but not laterally contiguous
. 11
9(8). Operculum disc bare, with a few strong
spines along lateral margin; convolu-
tions of respiratory horn extending


from proximal 4 to slightly past
midlength; 1pm tubercles strongly bifid .
footed Wirth and Jones, p. 85
9'. Operculum with spines on distal area as
well as along margins; convolutions of
respiratory horn extending from 0.45 to
0.75 of distance to tip; Ipm tubercles
not bifid, with rounded or blunt point
and subapical seta .. .... .. 10
10(9). Operculum between am tubercles bear-
ing moderate to distinct spines
haematopotus Malloch, p. 95
10'. Operculum between am tubercles
without spines
.... edeni Wirth and Blanton, p. 81
11(8). Respiratory horn with convolutions
confined to a narrow band just distad of
distal lateral spiracular opening; disc of
operculum with numerous sharp spines;
1pm tubercles rounded or with blunt
points, with a subapical seta ...
............ bauer Hoffman, p. 61
11', Respiratory horn with convolutions in a
broader band at least some of which are
located proximad of distal lateral
spiracular opening; disc of operculum
with or without spines; Ipm tubercles
bifid ......... ....... ....... 12
12(11). Operculum with long sharp spines on
disc; d tubercles 1 and 2 out of line 13
12'. Operculum with short blunt spines on
disc; d tubercles 1 and 2 in line ...
stellifer (Coquillett), p. 149
13(12). Apicolateral processes of caudal seg-
ment widely divergent, forming angle
of 75 with longitudinal axis of body
barbosai Wirth and Blanton, p. 58
13'. Apicolateral processes less divergent,
forming an angle of 50 with
longitudinal axis of body
furens (Poey), p. 87
14(4). Respiratory horn with transverse con-
volutions in midportion .. 15
14'. Respiratory horn without transverse
convolutions in midportion .... 17
15(14). Operculum with blunt nodules in
sublateral patches on disc; am setae
long and blunt tipped, 0.8 as long as
maximum width of operculum; small
yellowish pupa .
. floridensis Beck, p. 83
15'. Operculum with sharp spines on disc;
am setae short and sharp, less than 'A
as long as maximum width of oper-
culum; large brownish pupa 16
16(15). Respiratory horn with 6-8 apical
spiracular openings; caudal segment







with patch of spines on disc and spines
on apicolateral processes
crepuscularis Malloch, p. 75
16'. Respiratory horn with 11 15 apical
spiracular openings; caudal segment
without patch of spines on disc or
spines on apicolateral processes
variipennis (Coquillett), p. 161
17(14). Abdominal segments entirely covered
with confluent scalelike spines giving a
reticulated appearance; respiratory
horn short and stout, widest near tip,
L/W Ratio 4.5, with 11-15 apical spir
acular openings
.. guttipennis (Coquillett), p. 92
17'. Abdominal segments with spines not
confluent, not giving a reticulated ap-
pearance; respiratory horn more
elongate and slender, L/W Ratio 5.1-8.0,
with not more than 12 spiracular
openings 18
18(17). Operculum with disc bare, spines con-
fined to a few along lateral margins

18'. Operculum with spines on disc 20
19(18). Caudal segment without patch of
spines on disc; respiratory horn short,
L/W Ratio 6.9 ..............
S............. loisae Jamnback, p. 112
19'. Caudal segment with patch of spines on
disc; respiratory horn longer, L/W
Ratio 8.0
scanloni Wirth and Hubert, p. 142
20(18). Caudal segment without patch of
spines on disc 21
20'. Caudal segment with patch of spines on
disc 22
21(20). Operculum with longer spines exten-
ding only halfway from am tubercles to
posterior margin; respiratory horn with
few spines, nearly bare ..
bermudensis Williams, p. 65
21'. Operculum with numerous short spines
extending all the way to posterior
margin; respiratory horn with abun-
dant spines
arboricola Root and Hoffman, p. 56
22(20). Caudal apicolateral processes with
apices directed parallel to longitudinal
axis of body; respiratory horn with
abundant coarse spines .. 23
22'. Caudal apicolateral processes with
apices directed 20-45" to longitudinal
axis of body; respiratory horn with
coarse or fine spines .. ...... 25


23(22). Thoracic d tubercles 1 and 2 closer than
2 and 3; respiratory horn shorter, L/W
Ratio 5.0, with 8-9 apical spiracular
openings; apicolateral processes with
tips darkened
parapiliferus Wirth and Blanton, p. 134
23'. Thoracic d tubercles 1, 2, and 3 equidis-
tant; respiratory horn longer, L/W
Ratio 5.6-6.0, with 4-6 apical spiracular
openings 24
24(23). Apicolateral processes with apices not
darkened .
. biguttatus (Coquillett), p.70
24'. Apicolateral processes with apical 1
darkened traoeisi Vargas, p. 159
25(22). Respiratory horn with 6-7 apical
spiracular openings ... 26
25'. Respiratory horn with 3-5 apical
spiracular openings ........ 28
26(25). Surface of respiratory horn with promi-
nent coarse spines; d tubercles 1 and 2
closer than 2 and 3 .............. 27
26'. Surface or respiratory horn with in-
conspicuous spines; d tubercles 1, 2,
and 3 equidistant
spinosus Root and Hoffman, p. 146
27(26). Operrulum with numerous strong sharp
spines on disc
.. snowi Wirth and Jones, p. 144
27'. Operculum with fewer weak spines on
disc .
villosipennis Root and Hoffman, p. 169
28(25). Operculum with surface of disc weakly
papillate without definite spines;
apicolateral processes diverging at
angle of 20-30 ............. 29
28'. Operculum with surface of disc bearing
definite sharp spines; apicolateral pro-
cesses diverging at angle of 30 30
29(28). Respiratory horn longer, L/W Ratio 7.5;
4 apical spiracular openings; caudal
apicolateral processes diverging at
angle of 30.
lesludinalis Wirth and Hubert, p. 152
29'. Respiratory horn shorter, L/W Ratio
6.3; 5 apical spiracular openings;
apicolateral processes diverging at
angle of 20'
niger Root and Hoffman, p. 126
30(28). Respiratory horn longer, L/W Ratio 7.2
piliferus Root and Hoffman, p. 137
30'. Respiratory horn shorter, L/W Ratio
6.0 bickleyi Wirth and Hubert, p. 67







PARTIAL KEY TO KNOWN LARVAE
OF FLORIDA CULICOIDES
(MATURE LARVAE)
(From Jamnback, 1965)


1. Very large larvae, about 8 mm long;
frontoclypeus 0.25-0.28 mm long,
yellow, brownish posteriorly;
pharyngeal apparatus massive, heavily
sclerotized, dorsum of thoracic
segments uniformly brownish; head
capsule strongly tapered anteriorly .
........ variipennis (Coquillett), p. 161

1'. Smaller larvae, less than 6 mm long;
frontoclypeus less than 0.24 mm long
(except C. venustus); pharyngeal ap-
paratus not massive or heavily
sclerotized, as usual in genus .... 2

2(1). Head capsule brown, frontoclypeus
0.23-0.28 mm long; thoracic dorsum
with numerous discrete pigmented
spots ....... venustus Hoffman, p. 166

2'. Head capsule yellow, at most tinged
with brown; frontoclypeus less than
0.23 mm long (except C. guttipennis);
thoracic dorsum various ....... 3

3(2). Setae on last abdominal segment long,
longer than maximum width of seg-
ment .. 4
3'. Setae on last abdominal segment short,
less than A as long as maximum width
of segment ........ . .... 5

4(3). Frontoclypeus 0.23-0.25 mm long ....
... guttipennis (Coquillett), p. 92
4'. Frontoclypeus 0.22 mm long ......
villosipennis Root and
Hoffman, p. 169
5(3). Dorsum of thorax pigmented, pro-
thorax almost entirely covered with
pigmentation ....... 6

5'. Thoracic pigmentation absent, or
limited to lateral spots, if on dorsum
confined to transverse bars near
anterior margins of segments ...... 9

6(5). Distinct pigmented lateral bodies on
meso- and metathoracic segments in ad-
dition to pigmentation on dorsum ... 7

6'. Lateralpigmented bodies absent 8

7(6). All thoracic segments with dorsum
almost entirely pigmented
.... crepuscularis Malloch, p. 75


7'. Meso- and metathoracic segments with
pigmentation lighter and confined to
anterior 1/2 or less .
hollensis (Melander and Brues), p. 101
8(6). Pigmentation arranged in a distinct
and definite pattern on dorsum of pro-
and mesothorax (fig. 220); pigment red-
dish purple ...... ... . .
..... haematopotus Malloch, p. 95
8'. Pigmentation diffuse over entire dor-
sum, especially on pro- and mesothorax
(fig. 218); pigment dirty brownish
... ........ furens (Poey), p. 87
9(5). Venter of head capsule with 3
longitudinal selerotized brown lines
running the length of head capsule;
thoracic markings may be very faint...
.... melleus (Coquillett), p. 116
9'. Venter of head capsule without
sclerotized brown lines ....... 10
10(9). Thorax with lateral pigmented bodies
at midlength on each segment; eyes
small, consisting of a single spot on
each side chiopterus (Meigen), p. 72
10'. Prothorax with lateral pigmented
bodies near anterior margin present or
absent, those on meso- and metathorax
at midlength; eyes larger consisting of 2
spots on each side ......... .. 11
11(10). Small, glistening white larvae, fron-
toclypeus 0.13-0.15 mm long; lateral
pigmented spots on meso- and
metathorax faint or absent, none
elsewhere stellifer(Coquillett),p. 149
11'. Larger larvae, frontoclypeus more than
0.18 mm long: lateral pigmented spots
on meso- and metathorax distinct,
transverse bands often present on dor-
sum near anterior margins of pro- and
mesothoracic segments .a.. 67
.... bickleyi Wirth and Hubert, p. 67







DIAGNOSTIC TABLES FOR QUICK IDENTIFICATION


USEFUL SORTING CHARACTERS
FOR CULICOIDES FEMALES


1. Wings (see fig. 226-264):
a. Without markings: floridensis, loisae,
melleus, tissoti.
b. Two anterior spots only: biguttatus,
husseyi, mulrennani, testudinalis, (some
hollensis).
c. Two anterior pale spots plus diffuse
distal markings: bermudensis, footei,
niger, snowi, spinosus, travisi, (some
hollensis, testudinalis).
d. Debilipalpis type pattern: debilipalpis,
hinmani, torreyae.
e. Crepuscularis type pattern:
crepuscularis, knowltoni, mississippien-
sis.
f. Guttipennis type pattern: arboricola,
bechae, guttipennis, ousairani,
villosipennis
g. Insignis type pattern: insignis,
venustus.
h. Obsoletus type pattern:alachua,
chiopterus, juddi, pechumani, pusillus.
i. Pattern diagnostic for species: barbosai,
baueri, edeni, haematopotus, furens,
loughnani, nanus, paraensis, stellifer,
variipennis.
2. Spermathecae:
a. One spermatheca present (fig. 47e): ber-
mudensis, crepuscularis, hollensis,
knowltoni, mississippiensis, variipennis.
b. Two spermathecae present but sclero-
tized ring absent (fig. 87e): loisae,
melleus, mulrennani, spinosus.
c. Two spermathecae, rudimentary 3rd,
and sclerotized ring (fig. 39e): most
species
3. Antennal sensillar patterns:
a. 3-15: beckae, crepuscularis, haemato-
potus, knowltoni, loughnani, nanus,
niger, ousairani, travisi.
b. 3-10: footei.
c. 3,7-10: barbosai, baueri, furens
d. 3,8-10: debilipalpis, paraensis, stellifer,
torreya, variipennis.
e. 3,10-14: melleus
f. 3,10-15: edeni.
g. 3,11-15: alachua, biguttatus, chiopterus,
hinmani, hollensis, juddi, loisae,
mississippiensis, mulrennani, hollensis,
pechumani, spinosus, ven'ustus.
h. 3,13-15: bickleyi, pusillus.
i. 3,13-14: bermudensis.


j. 3,7-14: tissoti.
k. 3,5,7,9,11-15: arboricola, guttipennis,
husseyi, insignis, snowi, villosipennis.
1. 3,5,7,9-15: piliferus, scanloni.
m. 3,5,7,9,13-15: testudinalis.
n. 3,5,7,9,11,13-15: parapiliferus.
o. None: floridensis.
4. Eyes broadly separated (fig. 47d): ber-
mudensis, hollensis, mississippiensis,
parapiliferus, snowi, testudinalis, tissoti,
variipennis.
5. Eyes contiguous (fig. 37d): alachua,
chiopterus, hinmani, insignia, juddi,
pechumani, venustus.
6. Antennal segments 8-10 short, last 5
elongated, AR 1.60 or more (fig. 105a):
edeni, haematopotus, piliferus, villosipen-
nis.
7. Palpal pit deep and opening by smaller pore
(fig. 57b): debilipalpis, footei, hinmani,
loughnani, nanus, ousairani, paraensis.
8. Third palpal segment without definite pit
(fig. 51b): biguttatus, insignis, melleus,
mulrennani, stellifer, venustus
9. Mandibular teeth 10 or less or absent: ber-
mudensis, juddi, loisae, pechumani.


USEFUL SORTING CHARACTERS FOR
CULICOIDES MALES

1. Apicolateral processes on 9th tergum:
a. Absent: alachua, pusillus.
b. Very small: insignia, venustus.
c. Long and stout: arboricola, beckae,
biguttatus, guttipennis, knowltoni,
melleus, villosipennis.
d. Long and slender: bermudensis, bickleyi,
crepuscularis, furens, hinmani, hollensis,
husseyi, loughnani, mulrennani, nanus,
niger, ousairani, snowi, spinosus,
stellifer, tissoti, travisi, variipennis.
2. Ventral root of basistyle:
a. Absent: bermudensis, insignis,
knowltoni, venustus.
b. Foot-shaped: barbosai, baueri, bickleyi,
debilipalpis, edeni, furens,
haematopotus, paraensis, parapiliferus,
piliferus, scanloni, snowi, stellifer,
testudinalis, torreyae.
3. Aedeagus:
a. Basal transverse bar and internal peg:
insignis, venustus.
b. Long sublateral process on shoulder:
baueri, edeni, footei, haematopotus.
c. Subapical processes near tip: villosipen-
nis.







4. Parameres:
a. Fused or basally joined: hollensis, in-
signis, knowltoni, pusillus, uariipennis.
b. Microscopic hairs at tip: insignis,
venustus.
c. Apical crown of spines: loisae, spinosus.
d. Lateral fringing spines near tip: bar-
bosai, baueri, bickleyi, debilipalpis,
edeni, floridensis, furens, husseyi,
paraensis, parapiliferus, piliferus,
scanloni, snow stellifer, testudinalis,
torreyae.
e. Ventral lobe present; barbosai,
debilipalpis, edeni, floridensls, furens,
haematopotus, paraensis, stellifer, tor-
reyae.


USEFUL SORTING CHARACTERS FOR
CULICOIDES PUPAE


1. Operculum (see fig. 159-186):
a. Very long, hair-like spines (fig. 165):
chiopterus.
b. Moderately long spines (fig. 161): baueri,
bermudensis, furens, haematopotus,
hollensis, melleus (confined mainly to
lateral margins in melleus).
c. Short spines on lateral margin, disc
papillose or bare (fig. 168): footei, loisae,
nanus, niger, parapiliferus, scanloni,
testudinalis.
d. Short spines abundant over most of sur-
face posterior to anteromedian tubercles,
except at extreme posterior end (fig.
159): arboricola, crepuscularis, guttipen-
nis, snowi, uariipennis, villosipennis.
e. Short spines on lateral margins and disc
(fig. 163): bickleyi, biguttatus, piliferus,
spinosus, stellifer, travisi, venustus (few
or none on disc in venustus).
2. Caudal segment with patch of spines on disc
(fig. 210): bickleyi, biguttatus,
crepuscularis, footei, haematopotus, nanus,
niger, parapiliferus, piliferus, scanloni,
snowi, spinosus, stellifer, testudinalis,
travisi, venustus (present as transverse
band of spines in chiopterus).
3. Caudal apicolateral processes extending out
as right angles to longitudinal axis of body:
hollensis, melleus, variipennis.
4. d tubercles 1 and 2 closer than 2 and 3, not
in line (fig. 194): furens, haematopotus,
hollensis, melleus.
5. d setae 1 and 2 long, subequal (fig. 189): ber-
mudensis, bickleyi, crepuscularis, hollensis,
loisae, melleus, niger, piliferus, scanloni,
snowi, stellifer, variipennis.


6. d setae 1 and 2 short, subequal (fig. 187): ar-
boricola, biguttatus, furens, guttipennis,
haematopotus, nanus, spinosus, travisi.
7. d seta 2 much longer than seta 1:
chiopterus.
8. Respiratory horn without lateral spiracular
openings (fig. 145, 157): insignis, venustus.


USEFUL SORTING CHARACTERS FOR
CULICOIDES LARVAE


1. Dorsum of prothorax completely or almost
completely pigmented (fig. 217):
crepuscularis, furens, guttipennis (often
very faint), haemutopotus, hollensis,
variipennis.
2. Thoracic pigmentation confined to lateral
spots, often lacking on the prothorax (ex-
cept sometimes narrow transverse
pigmented bars are present near anterior
margins of the segments) (fig. 215): bickleyi,
chiopterus, melleus, (often very faint or ab-
sent), stellifer (often faint).
3. Thorax with numerous discrete pigmented
spots on dorsum (fig. 225): venustus.
4. Head capsule brown: venustus.
5. Head capsule strongly tapered anteriorly
(fig. 218): furens, haematopotus, variipen-
nis.
6. Head capsule with 3 longitudinal sclero-
tized bars on ventral side: melleus.
7. Head:
a. Very short (frontoclypeus 0.18 mm or
less) (fig. 216): bermudensis, chiopterus,
haematopotus, stellifer.
b. Very long (frontoclypeus 0.20 mm or
more) (fig. 219): guttipennis, hollensis,
variipennis, venustus, villosipennis.
8. Eyes unusually small, each composed of a
single round or U-shaped pigmented spot
(fig. 216): chiopterus.







SPECIES DESCRIPTIONS


Culicoides alachua Jamnback and Wirth
(fig. 37, 38, 228)


Culicoides alachua Jamnback and Wirth,
1963:187 (male, female; Florida; fig.).
Culicoides obsoletus (Meigen), misident.; Beck,
1952:102 (Florida record); Beck, 1956:133
(Florida); Beck, 1958:8 (Florida).

TYPES: Holotype, male, allotype, female,
Island Grove, Alachua Co., Florida, 22 March
1955 holotypee) and 18 March 1955 (allotype),
light trap (USNM 69114).

FEMALE: Wing length 0.93 mm.

HEAD: Eyes (fig. 37d) bare; contiguous.
Antenna (fig. 37a) with lengths of flagellar
segments in proportion of 28-25-25-25-26-26-
27-30-45-49-50-55-83; AR 1.33; sensory pattern
3,11-15. Third palpal segment (fig. 37b)


slender, with small, round, shallow, sensory pit
near tip; PR 2.8. Proboscis moderately long,
P/H Ratio 0.80; mandible with 13 teeth.

THORAX: Dark brown; mesonotum with pro-
minent pattern, disc pruinose bright gray, with
sublateral pair of dark brown vittae. Legs (fig.
37f) pale brown, knee spots blackish, femora
darker distally, all tibiae with indistinct, nar-
row, basal pale rings; tibial comb with 5 spines,
that nearest the spur longest.

WING (fig. 37c, 228): With moderately
distinct pattern of rather small pale spots; key
features include distal 1 of 2nd radial cell ly-
ing in a transverse distal pale spot extending
across cell R5 nearly to vein Ml; small round
pale spot lying over r-m crossvein not reaching
costa or into cell M2; elongated pale spots in
apices of cells R5, M1, and M2 reaching wing
margin; pale spots in cells M1 and M2 in line
transversely with poststigmatic pale spot in
cell R5 and large pale spot in cell M4; anal cell


ZD=IZTrZr300 0a0
a


*-7--


czzz-cD


Fig. 37. Culicoides alachua: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f, hind femur and tibia; g,
male parameres; h, male genitalia, parameres removed.







dark with small round pale spot in distal por-
tion; macrotrichia sparse but coarse, confined
to distal '/3 of wing; radial cells distinct, 2nd
short and broad; CR 0.60. Halter pale.

ABDOMEN: Brown, darker distally. Sper-
mathecae (fig. 37e) 2, subspherical to ovoid
with short, slender necks; subequal, each
measuring 0.045 by 0.033 mm.

MALE GENITALIA (fig. 37h): Ninth ster-
num with deep, narrow, median cleft on caudal
margin, with adjacent corners rounded; 9th
tergum rounded distally, without trace of
apicolateral processes. Basistyle stout, ventral
root slender, dorsal root short and angular;
dististyle slender and rounded distally, not
swollen at tip. Aedeagus with basal arch exten-
ding to 0.6 of total length; distal portion with
slender elongated posterior process, bearing a
broad, rounded, subapical, ventral lobe.
Parameres (fig. 37g) separate, each with long
lateral process from base; stem swollen at base,
tapering distally to slender filament lacking
distal fringing hairs.

DISCUSSION: C. alachua is closely related
to C. obsoletus (Meigen) and C. sanguisuga (Co-
quillett), which it replaces in Florida. The wing
of C. alachua has more restricted pale mark-
ings than in the other 2 species, and the body
color is darker and the 3rd palpal segment is
stouter than in C. sanguisuga. The male
genitalia of C. alachua has the median cleft on
the 9th sternum and rounded 9th tergum
similar to those of C. obsoletus and C.
sanguisuga, but the aedeagus is quite distinc-
tive, with a conspicuous, broadly rounded, ven-
tral lobe.

IMMATURE STAGES: Unknown.

LARVAL HABITAT: Unknown. Other
members of this group breed in wet leaves and
compost, strawy manure and similar moist
habitats associated more with decaying
organic matter than with wet soil or mud
(Jamnback and Wirth, 1963). Cattle manure
should be investigated as a likely breeding site
for this species, since it affords breeding
places for the closely related species, C.
chiopterus (Meigen).

FEEDING HABITS: Unknown. All our
material is from light trap collections.

SEASONAL DISTRIBUTION: Reported
by Mrs. Beck (1952, 1956, 1958) from 6 coun-
ties in central Florida (25 Jahuary to 3
August). In our records C. alachua was taken
in February and March in Highlands County,


January in Lake County, and from 4 April un-
til 3 August in northern Florida counties.


DISTRIBUTION: Northern Florida, South
Carolina.


FLORIDA DISTRIBUTION (fig. 38): C.
alachua apparently is restricted in Florida to
the Peninsular Uplands Region, our records
coming from Liberty, Jefferson, Alachua, Put-
nam, Marion, Lake, Orange, and Highlands
counties. We have examined 3 males and 69
females, as follows: Alachua Co., Gainesville,
IV-64, J. Spooner, LT, 2 females; 20-IV-67,
WWW, LT, 1 female; 29-VI-67, FSB, LT, 1
female. Alachua Co., Island Grove, III-55,
SBH, LT, 2 males, 1 female; 3-VIII-66, W. W.
Smith, LT, 16 females; 1-VIII-66, FSB, LT, 1
female. Highlands Co., Highlands Hammock
St. Park, 1-III-49, SBH, LT, 1 female.
Highlands Co., Sebring, 2-11-48, C. T. Parsons,
1 female. Jefferson Co., Monticello, 7-VI-61,
SBH, LT, 1 female. Lake Co., Goose Prairie,
9-III-54, SBH, LT, 1 male. Lake Co., Leesburg,
25-1-55, SBH, LT, 1 female. Liberty Co.,
Telogia, V-61, SBH, LT, 1 female. Marion Co.,
Citra, 5-IV-54, SBH, LT, 3 females. Orange
Co., Orlando, 19-VII-69, G. M. Stokes, LT, 2
females. Putnam Co., Lons Lake, VII-71, FSB,
LT, 1 female. Putnam Co., Redwater Lake,
27-V-67, FSB, LT, 5 females.


Fig. 38. Florida distribution of Culicoides alachua.






Culicoides arboricola Root and Hoffman
(fig. 36e, 39, 40, 131, 159, 187, 208, 236)


Culicoides arboricola Root and Hoffman,
1937:166 (male, female; Maryland; fig.);
Fox, 1942:416 (pupa; Maryland; fig.); Beck,
1952:103 (Florida); Foote and Pratt,
1954:15 (redescribed; habits; distribution;
fig.); Snow, Pickard, and Moore, 1957:22
(Tennessee; habits); Beck, 1958:8 (notes;
Florida); Snow and Pickard, 1958:3
(Alabama; breeding habits); Jones,
1961a:702 (larval habitats); Messersmith,
1965:321 (Virginia); Jamnback, 1965:40
(redescribed; all stages; biology; fig.); Hair,
Turner and Messersmith, 1966:195 (larval
habitat; Virginia); Messersmith, 1966:93
(Virginia; seasonal incidence); Wirth and
Blanton, 1967:210 (redescribed; fig.;
distribution); Smith and Varnell, 1967:519
(notes; Florida); Childers and Wingo,
1968:10 (Missouri; habitat; seasonal
distribution); Khalaf, 1969:1159 (Louisiana;
seasonal incidence); Gazeau and
Messersmith, 1970a:32 (Maryland; larval
habitat); Linley, 1970b:717 (immature
stages; fig.; Florida); Battle and Turner,


1971:22 (redescribed; fig.; Virginia); Hum-
phreys and Turner, 1973:82 (feeding
preference; Virginia).
TYPE: Holotype, male, Gwynns Falls Park,
Baltimore, Maryland, July 1931, tree hole, F.
M. Root and W. A. Hoffman (USNM 53305).

FEMALE: Wing length 1.19 mm.

HEAD: Eyes (fig. 39d) bare; narrowly
separated. Antenna (fig. 39a) with lengths of
flagellar segments in proportion of
22-17-17-17-18-18-18-18-39-39-39-41-57; AR
1.48; sensory pattern 3,5,7,11-15. Third palpal
segment (fig. 39b) moderately swollen, PR 2.8;
sensory pit broad and shallow. Proboscis long,
P/H Ratio 1.00; mandible with 15 teeth.
THORAX: Dark brown; mesonotum with pat-
tern of pruinose gray patches. Legs (fig. 39f)
brown, knee spots blackish; all femora with
subapical, and all tibiae with subbasal, promi-
nent pale rings, hind tibia with distal pale;
tibial comb with 5 spines, the 2 nearest the
spur longest, subequal.
WING (fig. 39c, 236): With prominent pattern
key features include double distal pale spot in


a




----------------- ~ -~~--~
b










d
.\J ..











f


Fig. 39. Culicoides arboricola: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f, hind femur and tibia;
g, male parameres; h, male genitalia, parameres removed.






cell R5, pale spots straddling base of vein Ml
and midportion of vein M2, anal cell with pale
base and 3 pale spots distally; apices of veins
Ml, M2, M3+4, and Cul pale at wing margin;
pale spot over r-m crossvein extending broadly
into cell M2; macrotrichia long and abundant;
CR 0.60. Halter pale.

ABDOMEN: Dark brown. Spermathecae (fig.
39e) 2, subspherical to slightly ovoid with
short slender necks; slightly unequal, measur-
ing 0.052 by 0.045 mm and 0.048 by 0.040 mm.

MALE GENITALIA (fig. 39h): Ninth ster-


tergum with moderately long, angulate,
apicolateral processes. Basistyle with ventral
and dorsal roots simple, lengths subequal;
dististyle nearly straight with slender, slightly
hooked, distal point. Aedeagus with basal arch
high, extending to 0.6 of total length; distal
process simple, tapering to slender, pointed
tip. Parameres (fig. 39g) separate; each with
large basal knob, base of stem constricted,
straight midportion somewhat swollen prox-
imad, tapering distally to slender pointed tip
abruptly bent anteroventrally.

DISCUSSION: C. arboricola is distinguished
from the other members of the guttipennis
group by its extensive pale wing markings
with pale apex of vein Cul in addition to those
at the apices of veins Ml, M2, and M3+4, and
by the subapical pale band on the hind femur.

IMMATURE STAGES: The pupa was
described and figured by Fox (1942) and Jamn-
back (1965). Linley (1970b) described and
figured the larva and pupa.

PUPA (fig. 36e): Respiratory horn (fig. 131)
uniformly light brown, with 9-12 apical and 2-3
lateral spiracular openings; moderately
slender, slightly wider near base, L/W Ratio
6.1. Operculum (fig. 159) almost entirely
covered with short, stout spines; am setae long
and slender, more than as long as maximum
width of operculum. The d tubercles 1-3 in line,
1 and 2 closer than 2 and 3, seta 1 and 2 very
short; thoracic surface between and adjacent
to tubercles covered with fine papillae. Ab-
domen entirely covered with fine sharp
spinules except for a narrow anterior ring
around each segment; Ipm tubercles (fig. 187)
each bifid, with seta unusually long; caudal
segment (fig. 208) with dorsum covered with
fine spinules; caudal apicolateral processes
covered with fine spinules to apiceS, apices not
darkened, tip directed posteriorly at an angle
of 25 from body axis.


MATURE LARVA: Head ambercolored, length
0.210 mm; comb with 5 spines. Thorax with
pigmented areas absent. Last abdominal seg-
ment with long perianal bristles, as long as
maximum width of segment.

LARVAL HABITAT: Wet tree holes, wet
wood debris, water and moist woody debris
(Hair et al., 1966). Snow et al. (1957) recorded
C. arboricola from water and moist debris in
tree cavities in the Tennessee River Valley.
Snow and Pickard (1958) recorded it from tree
cavities and tree stumps in Alabama, and
listed associated ceratopogonid species. Jones
(1961a) described tree hole breeding sites in
Texas and Wisconsin. Smith and Varnell
(1967) reared C. arboricola from tree holes in
Florida. Childers and Wingo (1968) found it
common in tree holes in central Missouri.
Gazeau and Messersmith (1970a) reared C. ar-
boricola from a tree hole in Maryland.

FEEDING HABITS: This species is
primarily ornithophilic. Snow et al. (1957)
found females in the Tennessee Valley resting
inside chicken sheds during the day and stated
that they never bite man. Messersmith (1965)
trapped large numbers of engorged females in
poultry houses in Virginia. Smith and Varnell
(1967) reported females of C. arboricola biting
man in Florida. Childers and Wingo (1968) took
large numbers of this species in a light trap
near a chicken coop in Missouri. Hair and
Turner (1968) reported biting of birds and man;
Humphreys and Turner (1973) on rabbits, in
Virginia. We have one record of C. arboricola
biting man in Gainesville, on a screened porch
at 11 p.m.

SEASONAL DISTRIBUTION: Jamnback
(1965) reported that C. arboricola breeds con-
tinuously in the south in warm weather.
Messersmith (1966) trapped it in Virginia from
July to September with a numerical peak in
August. Khalaf (1969) stated that C. arboricola
is a rare species in Louisiana, present from ear-
ly April through the summer. In Florida, Beck
(1952, 1958) gave records from 7 February to
14 December; it was taken in large numbers on-
ly in a Jackson County light trap, with a
population peak in July.

DISTRIBUTION: Eastern United States
from Minnesota and Texas to Connecticut and
Florida. This species is more common in the
southern part of its range. Williams (1955)
found it comprised 13% of his light trap cat-
ches in Baker County, Georgia, while
Messersmith (1965) counted it only 5.4% of his
collections in Virginia.






























00#OP


Fig. 40. Florida distribution of Culicoides arboricola.

FLORIDA DISTRIBUTION (fig. 40): We
have recorded C. arboricola from the entire
state, from Escambia County to the Keys,
where it has been common in all of our good
light trap locations. Records as follows:
Alachua Co., Gainesville, 15-V-64, T. Walker,
LT, 2 females; 19-V-65, SBH, LT, 1 female;
20-IV-67, WWW, LT, 1 male, 4 females;
V-VI-67, FSB, LT, 2 females; 1-V-73, R. I.
Sailer, biting man 11pm on screen porch, 3
females. Baker Co., Olustee, IX-71, FSB, LT, 1
female. Collier Co., Everglades City, 27-V-49,
Davidson, LT, 3 males, 2 females. Citrus Co.,
Crystal River, 7-VI-49, Hudson, LT, 1 female.
Dade Co., Homestead, 9-IX-68, Baranowski,
LT, 1 female. Dade Co., Orchid Jungle,
V-VI-69, Baranowski, LT, 1 male, 1 female.
Escambia Co., Bratt, 27-VIII-64, Blanton and
Broce, LT, 1 female. Escambia Co., Innerarity
Point, 29-IV-49, Rathert, LT, 1 male, 4
females. Escambia Co., Molino, 12-VIII-69,
FSB, LT, 10 females. Flagler Co., N. Flagler,
VIII-61, SBH, LT, 2 females. Franklin Co.,
Wright Lake near Sumatra, IV-73, G. B. Fair-
child, LT, 5 females. Gilchrist Co., Suwannee
River, 15-VI-3-VII-61, G. K. Hicks, LT, 1 male,
1 female. Hamilton Co., Jasper, IV-61, SBH,
LT, 1 male. Indian River Co., Vero Beach,
IX-59, ERC, LT, 1 female. Jefferson Co., Mon-
ticello, IV,VI,IX,X-69, W. H. Whitcomb, LT,
10 females. Leon Co., 3 mi N Tallahassee, V-70,
FSB, LT, 100 females. Leon Co., Tallahassee,
III-60, M. A. Kohn, at light,' 1 female. Leon
Co., Tall Timbers, IV-X-68, Bhaktar and
Baker, LT, 100's. Levy Co., Gulf Hammock,


Wacassassa Fish Camp, VIII-71, FSB, LT, 25
females. Liberty Co., Torreya State Park,
30-VI-57, FSB, LT, 3 females; V-71, FSB, LT, 1
female; 4-VII-65, H. V. Weems, Jr., LT, 1 male;
22-IV-67, WWW, LT, 1 male, 1 female. Marion
Co., Juniper Springs, 28-IV-70, WWW, LT, 2
males, 12 females. Monroe Co., Islamorada,
7-IX-48, Smith, LT, 3 males, 2 females. Monroe
Co., Marathon, Vaca Key, 27-VIII-47, W.
Buren, LT, 1 female. Orange Co., Lake
Magnolia Park, 6-VIII-70, E. Irons, LT, 1
male. Orange Co., Rock Springs, 21-VI-70,
WWW, LT, 2 females. Palm Beach Co., Lake
Worth, 9-VIII-51, WWW, LT, 1 male, 1 female.
Putnam Co., Lon's Lake, V,VI,VIII,IX-71,
FSB, LT, 80 females. Taylor Co., Blue Springs,
9-V-71, G. Butler, LT, 1 female. Wakulla Co.,
Ochlockonee River State Park, 29-IV-70,
WWW, LT, 4 females.


Culicoides barbosai Wirth and Blanton
(fig. 41, 42, 132, 160, 209)


Culicoides barbosai Wirth and Blanton,
1956a:161 (male, female; Panama, Florida,
Bahamas, Ecuador; fig.); Beck, 1958:10
(Florida); Lewis, 1958:721 (Jamaica; inter-
nal anatomy; fig.); Wirth and Blanton,
1959:400 (redescribed; Panama; fig.);
Breeland, 1960:164 (Panama; habits);
Davies, 1964:33 (biology; Jamaica); Linley,
1965a:58 (pupa; Jamaica; fig.); Linley,
1966b:1 (ovarian cycle; Jamaica); Davies,
1967:39 (Jamaica; larval habitat);
Gutsevich, Garcia, and Gonzales, 1969:4
(Cuba; notes); Kettle and Linley, 1969a:729
(Jamaica; biting habits); Kettle, 1969b:21
(Jamaica; biting habits); Kettle, 1969c:241
(Jamaica; biting habits); Linley and Davies,
1971:264 (biology and control; Florida,
West Indies); Wirth and Blanton, 1974a:26
(redescribed; fig.; West Indies distribution).

TYPE: Holotype, female, Mojinga Swamp,
Panama Canal Zone, January 1953, F. S. Blan-
ton, light trap (USNM 63157).

FEMALE: Wing length 0.88 mm.

HEAD: Eyes (fig. 41d) nearly contiguous,
bare. Antenna (fig. 41a) with lengths of
flagellar segments in proportion of
18-13-13-13-13-13-13-13-15-18-20-20-28; AR
0.93; sensory pattern 3,7-10, rarely absent on
7. Palpal segments (fig. 41c) with lengths in
proportion of 9-17-21-8-8; PR 2.2; 3rd segment
moderately swollen, with a small deep sensory
pit. Proboscis moderately short, P/H Ratio
0.68; mandible with 16 teeth.







THORAX: Mesonotum (fig. 41e) pruinose gray
with pattern of dark brown punctiform dots at
bases of mesonotal hairs, these dots irregularly
fused in some areas, principally in 2 sublateral
longitudinal bands. Scutellum narrowly dark
in middle, yellowish on sides. Legs (fig. 41g)
dark brown; knee spots blackish, all femora
with subapical, all tibiae with subbasal, and
hind tibia with apical, narrow pale rings; hind
tibial comb with 4 spines, the one nearest the
spur longest.
WING (fig. 41b, 247): Pattern as figured; 2nd
radial cell dark to tip; large yellowish anterior
spots at wing base and over r-m crossvein; 2
longitudinally elongate poststigmatic pale
spots in cell R5 narrowly fused to form an
hourglass-shaped spot; distal pale spot in cell
R5 large, rounded and broadly meeting
anterior wing margin; 2 pale spots in cell Ml,
the proximal one streak-like and lying adjacent
to vein M2, the distal one usually connected by
a narrow pale line to wing margin; veins M1


and M2 very faintly pale-margined on distal
halves; a pale line running through cell M2 to
the pale spot at apex of cell M2; large pale spot
in cell M4; 2 pale spots, more or less coalesced,
in distal part of anal cell. Macrotrichia
numerous on distal /2 of wing and a few in anal
cell; CR 0.58. Halter dark.

ABDOMEN: Dark brown, cerci pale. Sper-
mathecae (fig. 41f) 2, ovoid, subequal, each
measuring 0.045 by 0.030 mm, the bases of the
ducts sclerotized a short distance.

MALE GENITALIA (fig. 41i): Ninth ster-
num with broad caudomedian excavation, the
ventral membrane not spiculate; 9th tergum
long, the apicolateral processes moderately
long and slender. Basistyle with slender, foot-
shaped ventral root, dorsal root slender;
dististyle long and slender, with bent apex.
Aedeagus with basal arch extending to about
0.7 of total length, basal arms slender and


c cIDWa
a


0--


b g


* ,.
'

e


, \, / / \





(I


Fig. 41. Culicoides barbosai: a, antenna; b, wing; c, palpus; d, eye separation; e, mesonotum; f, spermathecae; g, hind
femur and tibia; h, male parameres; i, male genitalia, parameres removed.


d


f


Cn


e


'






slightly curved, distal portion tapered to
slender, rounded point with an indistinct
lateral pair of pointed hyaline processes near
juncture with the arch. Parameres (fig. 41h)
separate; each with strongly sclerotized basal
knob, stem slender and curved near base, distal
portion gradually more swollen, with a distinct
low ventral lobe, then distally becoming quite
slender, tapering to a fine point with a few
minute lateral spines.

DISCUSSION: C. barbosai closely
resembles C. furens (Poey) and before 1956 was
confused with that species in southern Florida
and the western Caribbean. C. furens is a
slightly larger species with more conspicuous
wing markings, in which the species may be
easily recognized by the presence of a small
round pale spot in cell R5 between the
poststigmatic pale spots and the irregular pale
spot in the distal part of the cell.

IMMATURE STAGES: Linley (1965a)
described and figured the pupa of C. barbosai
in Jamaica.

PUPA: Respiratory horn (fig. 132) pale,
darkened on apical 1/a; with 4-5 apical and 2-3
lateral spiracular openings, lateral openings
each on a prominent projection; median 1/ with
transverse convolutions, spines lacking; widest
nearest base, somewhat tapering distally; L/W
Ratio 7.0. Operculum (fig. 160) with long
spines abundant along margins and on disc,
spines denser and longer than in C. furens; am
seta long and stout, about 0.4 as long as max-
imum width of operculum. The d tubercles as
in C. furens, tubercles 1-3 not in line, seta 1 and
2 closer than 2 and 3, seta 1 and 2 rather short,
generally not overlapping, thoracic surface be-
tween them and adjacent to tubercles with few
papillae, nearly smooth, Abdomen with fine
nodules especially along anterior and posterior
margins of segments; Ipm tubercles 1 and 2
with seta flanked by 2 spines, tubercle 3 (up-
permost) with 3 spines ( in C. furens all lpm's
have a seta flanked by 2 spines). Caudal seg-
ment (fig. 209) encircled anteriorly by a band of
small spinules, no spines on caudal apicolateral
processes, the latter dark-tipped, directed
laterally at an angle of 70 to longitudinal axis
of body.

LARVAL HABITAT: Breeland (1960)
reared C. barbosai numerous times in Panama
from coastal mangrove areas, where it showed
a preference for coral sand habitats near the
ocean shore line. Linley (1965a) reared the
species from the banks of a small spit projec-
ting into a mangrove swamp at Reading, near


Fig. 42. Florida distribution of Culicoides barbosai.

Montego Bay, Jamaica. Larvae were found in
yellow mud, much drier than that yielding C.
insignis, more toward the landward side of the
spit. C. furens was also found breeding at this
spot. Davies (1967) reported that C. barbosai in
Jamaica is almost entirely confined to the tidal
waters of the seaward edge of mangrove
swamps and is not numerous in the red
mangrove habitat (Rhizophora mangle Lin-
naeus). The emergence rate is inversely propor-
tional to the mean tides, and outbreaks may be
expected in April-May and October-November
when the sea levels are minimal. C. barbosai
habitats are generally at about 0.2 foot lower
elevation than those of C. furens.

FEEDING HABITS: Studies on the biting
habits of C. barbosai in Jamaica were reported
by Kettle and Linley (1969a). This species was
found to prefer the arm to the leg, when biting
humans. Kettle (1969b) found that biting ac-
tivity was crepuscular and nocturnal with a
peak at the dawn hour, and (Kettle, 1969c) that
moderately high wind speed (above 3 mph)
reduced the biting rate.

SEASONAL DISTRIBUTION: Beck (1958)
gave records from Lee County, Florida, show-
ing sizeable populations from April through
July and in October, with the peak in May (10
April 21 December).


DISTRIBUTION: Bahamas,
Cayman Islands, Cuba, Jamaica,
Costa Rica, Panama, Ecuador.


Florida,
Mexico,







FLORIDA DISTRIBUTION (fig. 42): C.
barbosai is especially abundant in the
mangrove swamps in Monroe and Collier coun-
ties. It has been taken on the East Coast as far
north as Indian River County and on the West
Coast north to Lee County. One female of C.
barbosai in the USNM is labeled Mayport, 14
December 1925, W. E. Dove, indicating the
species may occur as far north as Duval Coun-
ty, but this record needs confirmation. Records
as follows: Collier Co., Caxambus, 13-XI-25,
W. E. Dove, 12 females. Colli o. oer Co., Collier
Seminole State Park, 17-V-73, WWW, LT, 8
females. Collier Co., Fakahatchee Island,
VIII-72, FSB, LT, 1 male, 1000 females. Collier
Co., Marco, 15-XI-25,-W. E. Dove, 2 females.
Dade Co., Miami Beach, IV-2, J. E. Porter,
sticky trap, 1 male, 10 females. Indian River
Co., Vero Beach, 7-III-61, ERC, LT, 2 females.
Lee Co., Bonita Beach, 17-IV-70, WWW,
biting man, 1 pm, 15 females; 19-V-73, WWW,
biting man, a, am, 100's. Lee Co., Sanibel
Island, 28-11-61, C. Woodring, LT, 7 females;
II,VIIVIII-61, SBH, LT, 3 females; 11-V-73,
WWW, LT, 10 females. Lee Co., St. James,
1-X-61, SBH, LT, 1 female. Monroe Co., Big
Pine Key, 3-VI-71, D. G. Young, LT, 1 female;
10-IV-70, WWW, LT, 1 female. Monroe Co.,
Key Largo, 16-111-65, S. Kemp, LT, 3 females.
Monroe Co., Marathon, Vaca Key, 16-XII-47,
W. Buren, LT, 5 females. Monroe Co., Rock
Harbor, 3-XII-48, Brown, LT, 1 female; VI-61,
SBH, LT, 2 females. Monroe Co., Stock Island,
3-XII-63, F. A. Buchanan, LT, 3 females. Palm
Beach Co., Lantana, X-51, R. Curran, LT, 1
female. St. Lucie Co., Pepper Park, 2 mi N Ft.
Pierce Inlet, 1961, SBH, LT, 1 female.


Culicoides baueri Hoffman
(fig. 43, 44, 133, 161, 188)


Culicoides baueri Hoffman, 1925:297 (female;
Maryland; fig.); Root and Hoffman,
1937:163 (notes; fig.; distribution); Beck,
1952:104 (Florida); Foote and Pratt,
1954:16 (notes; distribution; fig.); Williams,
1955a:33 (Georgia; larval habitat);
Messersmith, 1966:93 (Virginia; seasonal
incidence); Khalaf, 1969:1159 (Louisiana;
seasonal incidence); Battle and Turner,
1970:426 (North Carolina; larval habitat);
Gazeau and Messersmith, 1970a:32
(Maryland; habitat); Battle and Turner,
1971:24 (redescribed; Virginia; fig.);
Cochrane, 1974a:315 (compared with C.
bergi; distribution).


FEMALE: Wing length 1.03 mm.

HEAD: Eyes (fig. 43d) narrowly separated,
bare. Antenna (fig. 43a) with lengths of
flagellar segments in proportion of
21-14-14-15-15-15-15-16-23-25-26-27-39; AR
1.12; sensory pattern 3,7-10, sometimes also on
4,5,6,11. Third palpal segment (fig. 43b) great-
ly swollen, PR 2.0; sensory pit deep with large
opening. Proboscis moderately short, P/H
Ratio 0.64; mandible with 12 teeth.

THORAX: Mesonotum with distinct pattern.
Legs (fig. 43f) with knee spots dark, distinct
pale bands on each side of knees; hind tibia
with 4 spines, the 1st longest.

WING (fig. 43c): Pattern as figured; 1 pale
spot in distal part of anal cell; cell R5 with
transverse double pale spot in distal portion;
pale spot straddling midportion of vein M2 and
one lying in front of basal portion of vein Ml;
veins Ml, M2, and Cul with pale spot at wing
margin and vein M3+4 pale adjacent to the
pale spot in cell M4; CR 0.56. Halter pale.

ABDOMEN: Dark brown. Spermathecae (fig.
43e) 2, oval with long slender necks; unequal,
measuring 0.056 by 0.040 mm and 0.048 by
0.035 mm; sclerotized ring and rudimentary
3rd spermatheca present.

MALE GENITALIA (fig. 43h): Ninth ster-
num with shallow caudomedian excavation,
ventral membrane not spiculate; 9th tergum
with short, pointed, apicolateral processes.
Basistyle with ventral root foot-shaped, the
posterior "heel" long; dististyle slender with
bent, pointed tip. Aedeagus with rounded
basal arch extending to more than / of total
length; a pair of sublateral, pointed, posterior-
ly directed processes on shoulders of arch;
distomedian process tapering to slender tip.
Parameres (fig. 43g) separate; each with large
basal knob, midportion slender, ventral lobe
absent; tapering to recurved slender tip bear-
ing several lateral fringing spines.

DISCUSSION:The double distal pale spot in
cell R5 of C. baueri leads to confusion with C.
arboricola and other small individuals of the
guttipennis group, but all those species have 3
pale spots in the anal cell and differ in other
particulars. This species has been recorded
from the northern and western states, but
Cochrane (1974a) has shown that these records
pertain to a distinct species, C. bergi Cochrane.


TYPE: Holotype, female,' Baltimore, IMMATURE STAGES: Not described. The
Maryland, 16-17 September 1923, W. A. pupa described by Jones (1961b) and Jamn-
Hoffman, reared (USNM 27276). back (1965) as C. baueri, represents a distinct,






closely related species, C. bergi Cochrane. We
have included figures of the pupa of C. bergi
(fig. 133, 161, 188) for comparison.

LARVAL HABITAT: Williams (1955)
reared C. baueri from stream and spring
margins in Georgia. Gazeau and Messersmith
(1970a) reared the species from dead leaves and
mud in a swamp in Maryland. Battle and
Turner (1970) reared C. baueri from mud beside
a small stream in North Carolina.

FEEDING HABITS: We have 2 records of
this species biting man (see below).

SEASONAL DISTRIBUTION: Messer-
smith (1966) collected C. baueri in Virginia
from May through September with a peak in
June. Khalaf (1969) stated that C. baueri is a
rare species in Louisiana appearing in numbers
in late April and early May and persisting until
fall. Most of our Florida material was collected
in April and May.


DISTRIBUTION: Southeastern United
States from Tennessee and Louisiana to
Maryland and Florida.

FLORIDA DISTRIBUTION (fig. 44): In
Florida, our records of C. baueri are from Bratt
in Escambia County, and in a series of north
Florida counties from Liberty to Duval and
Orange counties. It was taken in numbers only
in Orange County where McDuffie collected 20
females biting man 15-XI-51 at Rock Springs.
Records as follows: Alachua Co., Gainesville,
5-V-62, SBH, LT, 1 female; 10-V, 20-VII,
11-VIII-67, FSB, LT, 4 females; 27-IV-70,
WWW, LT, 4 females; 3-V-73, WWW, LT, 10
males, 15 females; 4-V-73, WWW, swept from
wooded area at Hogtown Creek, 2 males, 1
female. Duval Co., Arlington, 30-V-50, VI-56,
E. C. Beck, LT, 1 male, 3 females. Duval Co.,
Jacksonville, 1-IV-67, C. G. Alvarez, biting
man, 1 female. Escambia Co., Bratt, VI-68,
FSB, LT, 1 female. Jackson Co., Florida
Caverns State Park, 26-V-73, WWW, LT, 1
male, 4 females. Levy Co., Williams Camp,
VII-60, G. K. Hicks, 2 females. Liberty Co.,


cr--^>cr-x~rncr~cr^0000oo00i
a


h\


d


(C


f


Fig. 43. Culicoides baueri: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f, hind femur and tibia; g,
male parameres; h, male genitalia, parameres removed.





























0PA s0


Fig. 44. Florida distribution of Culicoides baueri.

Torreya State Park, 15-IV-57, V-71, FSB, LT,
2 females; 20-V-66, H. V. Weems, Jr., LT, 8
females; 1-IX-67, FSB, LT, 2 females. Marion
Co., Citra, V-50, SBH, LT, 1 female. Orange
Co., Rock Springs, 15-XI-51, W. E. McDuffie,
biting man, 20 females; 21-IV-70, WWW, LT, 1
female. Polk Co., Mulberry, III, VI-55, SBH,
LT, 2 males, 1 female. St. Johns Co., St.
Augustine, V-56, SBH, LT, 1 female. Taylor
Co., Blue Springs, 9-V-71, G. Butler, LT, 2
females. Wakulla Co., Ochlockonee River State
Park, 29-IV-70, WWW, LT, 1 female.


Culicoides beckae Wirth and Blanton
(fig. 45, 46)


Culicoides beckae Wirth and Blanton, 1967:213
(male, female; Florida; fig.); Battle and
Turner, 1971:26 (redescribed; fig.; Virginia).

TYPE: Holotype, female, Gainesville,
Alachua Co., Florida, 23 July 1963, F. S. Blan-
ton, light trap (USNM 69485).

FEMALE: Wing length 1.15 mm.

HEAD: Eyes (fig. 45e) narrowly separated;
bare. Antenna (fig. 45a) with lengths of
flagellar segments in proportion of 36-30-30-
30-30-30-33-36-69-75-75-75-96; AR' 1.53; sen-
sory pattern 3-15. Third palpal segment (fig.
45b) moderately swollen on midportion, with


broad, shallow, sensory pit; PR 2.5. Proboscis
moderately long, P/H Ratio 0.83; mandible
with 15 teeth.

THORAX: Dark brown; mesonotum with
prominent pattern of large pruinose gray pat-
ches. Legs (fig. 45f) dark brown; knee spots
blackish; all femora with subapical and tibiae
with subbasal, narrow pale rings; hind tibia
with broad apical pale band; tibial comb with 5
spines, the two nearest the spur longer, sub-
equal.

WING (fig. 45c): With prominent pattern; key
features include double distal pale spot in cell
R5, pale spots straddling base of vein Ml and
midportion of vein M2; pale spots restricted,
pale spot over r-m crossvein small, not ex-
tending into costal cell or cell M2; apex of vein
Cul not pale at wing margin; macrotrichia
abundant and coarse; CR 0.61. Halter knob
dark.

ABDOMEN: Dark brown. Spermathecae (fig.
45d) 2 plus rudimentary 3rd and sclerotized
ring; functional spermathecae ovoid with short
necks; equal, each measuring 0.051 by 0.035
mm.

MALE GENITALIA (fig. 45h): Ninth ster-
num with broad, shallow, caudomedian excava-
tion, ventral membrane not spiculate; 9th
tergum moderately long with stout, angular,
apicolateral processes. Basistyle with ventral
and dorsal roots slender, subequal; dististyle
nearly straight, tapering to slender, pointed,
slightly bent tip. Aedeagus with basal arch ex-
tending to 1/2 of total length; distal process
slender with simple pointed tip. Parameres
(fig. 45g) each with basal knob directed laterad,
a constriction at base of stem; main portion
moderately swollen proximally, slender distal-
ly, tapering to slender distal filament.

DISCUSSION: C. beckae resembles C.
villosipennis and C. ousairani in its restricted
wing spots, but it differs from both in its dark
halteres and pale-banded hind femur. It differs
from C. villosipennis in its shorter distal anten-
nal segments and sensorial pattern, and from
C. ousairani in its mesonotal pattern and
shallow palpal pit. The genitalia of C. beckae
are most similar to those of C. arboricola, but
the parameres are much more slender distally.

IMMATURE STAGES: Unknown.

LARVAL HABITAT: Unknown; presumed
to be tree holes as in other members of the gut-
tipennis group.













_,-1

~~C


cDCz~iiiii


Fig. 45. Culicoides beckae: a, antenna; b, palpus; c, wing; d, spermathecae; e, eye separation; f, hind femur and tibia; g,
male parameres; h, male genitalia, parameres removed.


Fig. 46. Florida distribution of Culicoides beckae.


FEEDING HABITS: Unknown.

SEASONAL DISTRIBUTION: Battle and
Turner (1971) collected C. beckae in Virginia in
July and August. Our records are from April
through October in northern Florida.

DISTRIBUTION: Maryland, Virginia,
Alabama, Florida.

FLORIDA DISTRIBUTION (fig. 46): C.
beckae is a rare species known only from 7
north Florida counties. Our records: Alachua
Co., Gainesville, Chantilly Acres, IX-67, FSB,
LT, 1 female. Baker Co., Olustee, VII-71, FSB,
LT, 3 females. Escambia Co., Molino,
12-VIII-69, FSB, LT, 1 female. Gulf Co., St.
Joseph State Park, 1-V-70, WWW, LT, 1
female. Holmes Co., Bonifay, IX-61, SBH, LT,
2 females. Liberty Co., Telogia, X-61, SBH,
LT, 2 females. Liberty Co., Torreya State Park,
4-VII-65, H. V. Weems, Jr., LT, 1 female.
Wakulla Co., Ochlockonee River State Park,
29-IV-70, WWW, LT, 2 females.


'-~-~,~~c_~~~~3s~Os3~30~88~i







Culicoides bermudensis Williams
(fig. 47, 48, 134, 162, 189, 261)


Culicoides bermudensis Williams, 1956a:298
(female; fig.; Bermuda; habitat); Williams,
1956b:300 (Bermuda; habitat); Beck,
1956:134 (male; Florida); Williams, 1957:62
(Bermuda; habitats); Beck, 1958:10
(Florida; seasonal incidence); Jones and
Wirth, 1958:86 (Texas); Williams, 1961:116
(parthenogenesis in Bermuda); Jones,
1961a:703 (Texas; larval habitat); Jamn-
back, 1965:44 (New York; redescribed; all
stages; fig.); Khalaf, 1966a:229 (redes-
cribed; Louisiana); Khalaf, 1966b:882 (Loui-
siana; seasonal incidence); Khalaf,
1969:1159 (Louisiana; seasonal incidence);
Kline and Axtell, 1977:545 (larval habitat,
biology; North Carolina).

TYPE: Holotype, female, Trott's Pond,
Hamilton Parish, Bermuda, 12 June 1955, R.
W. Williams, recovery cage (USNM 71485).


FEMALE: Wing length 0.97 mm.

HEAD: Eyes (fig. 47d) broadly separated,
bare. Antenna (fig. 47a) with lengths of
flagellar segments in proportion of
20-15-15-15-15-14-14-14-18-20-22-25-35; AR
1.00; sensory pattern 3,13-15, sometimes
3,11-14. Palpus (fig. 47b) with lengths of
segments in proportion of 12-18-22-10-12; PR
2.2; 3rd segment short and slightly swollen,
with a small, shallow, round pit. Proboscis
short, P/H Ratio 0.67; mandible with 7
vestigial, very weakly sclerotized teeth.

THORAX: Uniformily dull grayish brown,
some specimens yellowish brown; with dense
pale grayish pollinosity. Legs (fig. 47f)
uniformly yellowish brown; tibial comb with 4
spines, that nearest the spur longest.

WING (fig. 47c, 261): Relatively narrow, with
inconspicuous pattern of diffuse paler streaks
between the brownish veins; generally rather
dark grayish brown. Macrotrichia sparse but
evenly distributed over most of wing; CR 0.60;


Q:CIDI3L\


a


b


--~~t~akr~~ ..
~u~a~*rr~~~mu~l,:'.i' :s
''- ~t~t~h.,
7.~.- ---..`--r
c. ,,
rt~~-. -
'c,
~....~
C
'


Fig. 47. Culicoides bermudensis: a, antenna; b, palpus; c, wing; d, eye separation; e, spermatheca; f, hind femur and
tibia; g, male parameres; h, male genitalia, parameres removed.


c"~,~~--?to~ooooO







radial cells narrow, especially distal portion of
2nd. Halter brownish infuscated.

ABDOMEN: Pale brown. Spermatheca (fig.
47e) 1, unusually large and lightly sclerotized,
oval with short slender neck, measuring 0.134
by 0.088 mm.

MALE GENITALIA (fig. 47h): Ninth ster-
num with moderately broad and deep caudome-
dian excavation, the ventral membrane with
fine spicules; 9th tergum long and tapering,
with long, slender, divergent, apicolateral pro-
cesses, the caudal margin between them slight-
ly cleft mesally. Basistyle with ventral root
vestigial, dorsal root short and angular;
dististyle rather stout proximad, gently
curved to slender, bent, apical point. Aedeagus
short and stout, basal arch extending to 1/ of
total length, basal arms short and stout;
distomedian process broad, bluntly rounded
distally, with strong lateral sclerotization on
each side forming a distinct ventral channel on
distal '/. Parameres (fig. 47g) separate, each
with large basal knob bearing a long anterior
process; midportion distinctly swollen prox-
imally, tapering distally to slender, simple
point abruptly bent ventrolaterad.

DISCUSSION: This species is distinguished
from the other members of the crepuscularis
group by its small size, uniformly brownish
mesonotum, narrow grayish wing with in-
distinct pale markings, its poorly developed
mandible teeth, short proboscis, antennal sen-
sory pattern, and very large, poorly sclerotized
spermatheca. The broad tip and strong lateral
sclerotizations on the male aedeagus are also
characteristic.
Khalaf (1966a) questioned the identity of the
Louisiana and Florida populations, in which
males are common, with the typical population
of Bermuda where males are unknown. How-
ever Williams (1961) discussed the evolu-
tionary mechanisms whereby an island popula-
tion might lose its need for blood-feeding and
sexual reproduction and could reproduce par-
thenogenetically.

IMMATURE STAGES: Jamnback (1965)
described and figured the larva and pupa of C.
bermudensis from New York.

PUPA: Respiratory horn (fig. 134) pale except
slightly darkened apically with 5 apical and 2-3
lateral spiracular openings; not narrowed nor
convoluted near middle, surface smooth with
only a few very inconspicuous scattered small
spines, L/W Ratio 5.6. Operculum (fig. 162)
with long stout spines moderately abundant
and confined mainly to the disc, but extending


farther posteriorly along lateral margins; am
setae short, less than /3 as long as maximum
width of operculum. The d tubercles 1-3 in line,
equidistant, setae 1 and 2 subequal, moderate-
ly long, area between d tubercles with a few
fine papillae. Abdomen with fine spines confin-
ed mostly to anterior margins of segments.
The Ipm tubercles (fig. 189) rounded or weakly
ridged. Caudal segment without patch of
spines on disc; caudal apicolateral processes
with small spines present and apical /4 darken-
ed, directed posteriorly at a 450 angle to
longitudinal axis of body.

MATURE LARVA: Frontoclypeus 0.091 mm
long; comb with 8-9 teeth on each side; total
comb width 0.038 mm.

LARVAL HABITAT (fig. 10): Williams
(1957) took C. bermudensis in recovery cages in
Bermuda in marshes with a salinity range from
1.2 to 31.5 parts per 1,000. Vegetation at one
site consisted of mangrove (Rhizophora), while
at other sites there were spike rush
(Eleocharis), prickly sedge (Mariscus), beard
grass (Polypogon), seashore rush grass
(Sporobolus), and Cape weed (Lippia). Jones
and Wirth (1958) and Jones (1961a) reported
hearings from margins of salt water pools in
Texas. Jamnback (1965) reared it from
saltmarsh sod beneath Spartina alterniflora in
New York. In coastal North Carolina, Kline
and Axtell (1977) found that C. bermudensis
larvae were restricted to areas of salt grass
(Distichlis spicata) and black needle rush Jun-
cus roemerianus). These areas were flooded by
tides only 6.4% to 11.4% of the time. In spite
of strict controls they reared a greater number
of adults from their larval rearing chambers
than the number of larvae added to the media,
and speculated that this phenomenon might be
accounted for by assuming larval
paedogenesis. Paedogenesis is a rare
phenomenon in the Diptera and has never
before been reported in ceratopogonids. They
reared only females of this species and never
found males in their emergence traps or adult
collections in North Carolina. They called for
further investigations on the biology of C. ber-
mudensis.

FEEDING HABITS: The short proboscis
and reduced mandibular teeth indicate that
this species probably does not take a blood
meal. Williams (1961) reported that it is
autogenous and parthenogenic in Bermuda
where males are unknown.

PARTHENOGENESIS: Williams (1961)
found no males of C. bermudensis in Bermuda
in 3 summers collecting by light traps and







emergence traps. In 1960 about 20% of the
females taken in emergence traps were gravid.
Larvae reared in the laboratory produced only
adult females. Some reared females produced
eggs which hatched into larvae. "It was thus
demonstrated that C. bermudensis had
developed an obligate thelytoky type of par-
thenogenesis, in which males are completely
absent, that this species is autogenous, and
that no form of nourishment, even plant
sugars, need be ingested prior to the develop-
ment and deposition of the first batch of eggs."
The occurrence and mechanism of par-
thenogenesis and autogeny in this species on
the mainland coasts needs investigation.
Males are taken in significant numbers along
with the females in most mainland localities
where C. bermudensis has been collected.

SEASONAL DISTRIBUTION: Collected in
June in Bermuda (Williams, 1956b); April,
May, and August in Texas (Jones and Wirth,
1958). Peak abundance in Florida light traps in
August (Beck, 1958), with collections from 13
March to 21 August. Khalaf (1966b, 1969)
trapped it in November and from March to
August in Louisiana.

DISTRIBUTION: Bermuda, seacoast of
eastern U.S.A. from New York to Florida and
Texas.


Fig. 48. Florida distribution of Culicoides bermudensis.


FLORIDA DISTRIBUTION (fig. 48): Our
records of C. bermudensis are scattered along
the Florida coast from 7 counties: Duval,
Volusia, Indian River, Dade, Collier, Gulf, and
Walton counties. The species was fairly
numerous in a grassy saltmarsh (fig. 10) at St.
Joseph State Park, Gulf County, 1-3-V-70,
where Wirth swept 4 males and 4 females in a
short period of collecting. Other records: Col-
lier Co., Fakahatchee Island, VIII-72, Blanton
and Meyer, LT, 1 female. Duval Co., Mayport,
16-XII-25, W. E. Dove, 20 females. Indian
River Co., Vero Beach, 5-III-58, VII-58, ERC,
LT, 4 females. Monroe Co., Key Largo,
16-III-65, S. Kemp, LT, 1 female. Volusia Co.,
New Smyrna Beach, 111-39, B. V. Travis, 2
females. Walton Co., Santa Rosa, 26-VII-55,
Peterson, LT, 1 male, 3 females.


Culicoides bickleyi Wirth and Hubert
(fig. 49, 50, 135, 163, 190, 210, 215, 240)


Culicoides bickleyi Wirth and Hubert, 1962:188
(male, female; Maryland; fig.); Jamnback,
1965:46 (redescribed, all stages; fig.; New
York; biology); Khalaf, 1966a:230 (Loui-
siana; notes); Khalaf, 1966b:881 (Louisiana;
seasonal incidence); Khalaf, 1967a:151
(Louisiana; seasonal incidence); Khalaf,
1967c:52 (Louisiana; notes); Khalaf,
1969:1159 (Louisiana; seasonal incidence);
Battle and Turner, 1971:28 (redescribed;
fig.; Virginia); Humphreys and Turner,
1973:82 (feeding preference; Virginia).

TYPES: Holotype, female, allotype, male,
Cranesville Swamp, Garrett County,
Maryland, 6 May 1960, W. W. Wirth, reared
from sphagnum (USNM 65717).

FEMALE: Wing length 1.07 mm.

HEAD: Eyes (fig. 49f) narrowly separated,
bare. Antenna (fig. 49a) with lengths of
flagellar segments in proportion of
19-12-12-12-12-12-13-13-19-20-24-27-28; AR
1.20; sensory pattern 3,13-15. Palpus (fig. 49b)
with lengths of segments in proportion of
9-20-22-10-13; PR 2.3; 3rd segment only slight-
ly swollen with a rather small, shallow, sensory
pit. Proboscis short, P/H Ratio 0.62; mandible
with 12 teeth.

THORAX: Dark brown, mesonotum with
grayish pruinosity leaving a sublateral pair of
dark brown patches. Legs (fig. 49g) pale brown;
all tibiae with narrow subbasal pale rings,
knees dark, fore and mid femora with in-
distinct subapical pale rings; tibial comb with






4 spines, the 2 nearest the spur longest, sub-
equal.

WING (fig. 49c, 240): With prominent exten-
sive pattern typical of the piliferus group;
poststigmatic pale spot occasionally involves
apex of 2nd radial cell. Macrotrichia long and
moderately dense over most of wing, area from
base of wing to anal angle and r-m crossvein
nearly bare; CR 0.60. Halter pale.

ABDOMEN: Pale brown. Spermathecae (fig.
49d) 2 plus rudimentary 3rd and sclerotized
ring; functional spermathecae long oval,
without sclerotized necks, unequal, measuring
0.075 by 0.047 mm and 0.051 by 0.034 mm.

MALE GENITALIA (fig. 49h): Ninth ster-
num with broad, moderately deep, caudome-
dian excavation, ventral membrane not
spiculate; 9th tergum moderately long and


tapering, apicolateral processes long and
slender, the caudal margin between them near-
ly straight. Basistyle with well developed
"heel" and "toe" on the foot-shaped ventral
root, dorsal root slender; dististyle gently curv-
ing, slender distally with bent, pointed tip.
Aedeagus with high, broad basal arch extend-
ing to 0.6 of total length, basal arms curved
and slender; distomedian process moderately
stout with subparallel sides and truncated tip.
Parameres (fig. 49g) separate; each with strong
basal knob, main body slender, curved at base,
sinuate in midportion, and tapering to slender,
pointed tip abruptly bent ventrad and bearing
lateral fringing spines.

DISCUSSION: This species can be distin-
guished from other members of the piliferus
group by its extensive pale wing pattern, nar-
rowly separated eyes, and antennal sensory
pattern 3,13-15.


a


b


e h



f g


Fig. 49. Culicoides bickleyi: a, antenna; b, palpus; c, wing; d, spermathecae; e, male parameres; f, eye separation; g,
hind femur and tibia; h, male genitalia, parameres removed.


~9~L~d~KdUL.
-~:'R~."'~-"
.M~ L-~~k~t~
f-l- Z.
'~~V~
.'
"'
.T, 'r
~~-~_ ~:~L*_ 5f
Y -- ~;LL L
G
:: -`L-' .*
h ~-L .5 *
~~.~
a, , -
.2. .i .,
s ;.c
lil I
Ilm, mn~sln~~~ii? ~. ..~.*n Iil C
i


Ca
dB


I







IMMATURE STAGES: Jamnback (1965)
described and figured the larva and pupa.

PUPA: Exuviae brownish. Respiratory horn
(fig. 139) dark brown, with 5 apical and 3
lateral spiracular openings; median 0.6 with
coarse spines, lacking transverse convolutions;
horn widest near base, L/W Ratio 6.0. Oper-
culum (fig. 163) dark brown; with distinct
spines along lateral margins, disc with
papilliform spines; am setae long, 0.4 as long
as maximum width of operculum. The d
tubercles 1-3 in line, 1 and 2 closer than 2 and
3, setae 1 and 2 short and stout; thoracic sur-
face between and adjacent to tubercles with
microscopic papillae. Abdomen with numerous
fine spines confined mostly to anterior margins
of segments, the surface with sparse spines on
posterior portions of segments; Ipm tubercles
(fig. 190) usually bifid, often with 1 of the
points weaker than the other. Caudal segment
(fig. 210) with patch of spines on disc;
apicolateral processes with small spines pres-
ent, distal 4 darkened, directed posteriorly at
an angle of 300 to longitudinal axis of body.

MATURE LARVA: Head capsule brownish
yellow, frontoclypeus slightly darker brown.
Frontoclypeus 0.197 mm long; comb with 7 un-
equal teeth on each side; total comb width
0.029 mm. Thorax (fig. 215) with distinct dark
markings; dorsum of prothorax with a
transverse bar near anterior margin;
mesothorax with transverse bar and oval
lateral markings; metathorax with longer and
wider lateral markings.

LARVAL HABITAT: Wirth and Hubert
(1962) reared C. bickleyi from sphagnum in a
larch swamp in Maryland. Jamnback (1965)
reared the species from soft mud taken 2
inches below the water surface in a small
woodland stream, from mixtures of decaying
hay, grass roots, and humus at 2 swamp sites,
and from thick sphagnum on the margins of a
swamp in New York.

FEEDING HABITS: Wirth and Hubert
(1962) reported this species biting man in
Massachusetts and West Virginia. Battle and
Turner (1971) state that the reduced sensillar
pattern indicates a preference for mammals.
Humphreys and Turner (1973) collected 39
females from goats and 1 from a turkey in
Virginia.


SEASONAL DISTRIBUTION: According
to Jamnback (1965) C. bickleyi is an early
spring species with adults most abundant in
June in the northeastern states and as early as
March in the south. Khalaf (1969) reports that
in Louisiana C. bickleyi reaches its population
peak in March and disappears by the last week
in May. Our Florida records are in April.

FLORIDA DISTRIBUTION (fig. 50): C.
bickleyi is rare in Florida. We have records
from: Highlands Co., Sebring, Highlands Ham-
mock State Park, 5-V-70,WWW, LT, 5 females.
Orange Co., Rock Springs, 21-IV-70, WWW,
LT, 1 female. Wakulla Co., Ochlockonee River
State Park, 29-IV-70, WWW, LT, 8 females.


Fig. 50. Florida distribution of Culicoides bickleyi.


69






Culicoides biguttatus (Coquillett)
(fig. 51, 52, 136, 164, 191, 250)


Ceratopogon biguttatus Coquillett, 1901:604
(female; D.C.).
Culicoides biguttatus (Coquillett); Kieffer,
1906:54 (combination); Malloch, 1915:308
(Illinois); Hoffman, 1925:283 (redescribed;
fig.; distribution); Root and Hoffman,
1937:170 (notes; fig.; distribution);
Thomsen,1937:70 (pupa; fig.); Beck,
1952:103 (Florida); Wirth, 1951:318
(Virginia; larval habitat); Foote and Pratt,
1954:16 (redescribed; fig.; distribution);
Williams, 1955b:115 (Michigan; habitats);
Beck, 1956:133 (Florida); Murray, 1957:77
(Virginia; habitats and seasonal distribu-
tion); Snow, Pickard, and Moore, 1957:22
(Tennessee Valley; larval habitats); Snow
and Pickard, 1958:3 (Tennessee); Jamn-
back, 1965:48 (redescribed; all stages; fig.;
biology; New York); Das Gupta and
Hansens, 1965:159 (redescribed; New
Jersey); Khalaf, 1966:881 (Louisiana;
seasonal distribution); Hair, Turner and


Messersmith, 1966:202 (Virginia; larval
habitats); Childers and Wingo, 1968:10
(Missouri; habitat; habits); Khalaf,
1969:1159 (Louisiana; seasonal incidence);
Gazeau and Messersmith, 1970b:32
(Maryland; habitat); Battle and Turner,
1971:30 (redescribed; fig.; Virginia); Kar-
datzke and Rowley, 1971:216 (Iowa; larval
habitats).

TYPE: Holotype, female, Washington, D. C.,
6 June, H. S. Barber (USNM 5473).

FEMALE: Wing length 1.30 mm.

HEAD: Eyes (fig. 51d) narrowly separated,
bare. Antenna (fig 51a) with lengths of flagellar
segments in proportion of 30-23-25-25-25-28-
28-28-40-40-40-46-56; AR 1.15; sensory pattern
3,11-15, series usually extending proximad
through some or all of segments 5-10. Palpus
(fig. 51b) with lengths of segments in propor-
tion of 15-55-53-18-15, 2nd segment unusually
long; PR 2.7; 3rd segment slightly swollen with
an irregular sensory pit. Proboscis moderately
long, P/H Ratio 0.85; mandible with 15 teeth.


a


' b

C'


Fig. 51. Culicoides biguttatus: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f, hind femur and
tibia; g, male parameres; h, male genitalia, parameres removed.






THORAX: Dark brown; mesonotum with
uniform, dense, grayish pollinosity. Legs (fig.
51f) brownish; knee spots blackish; tibiae with
faint basal pale rings; tibial comb with 4
spines, that nearest the spur longest.

WING (fig. 51c, 250): Uniformly dark grayish
brown, with pattern restricted to 2 small pale
spots, 1 over r-m crossvein, and the other on
anterior margin just past 2nd radial cell.
Macrotrichia unusually dense and coarse, ex-
tending to base of anal cell and nearly to base
of cell M2; CR 0.64; radial cells with broad
lumens. Halter infuscated.

ABDOMEN: Dark brown. Spermathecae (fig.
51e) 2 plus rudimentary 3rd and sclerotized
ring; functional spermathecae ovoid with stout
tapering necks, heavily sclerotized dark brown;
equal, each measuring 0.075 by 0.052 mm.

MALE GENITALIA (fig. 51h): Ninth ster-
num with broad, moderately deep, caudome-
dian excavation, the membrane with well
developed spicules; 9th tergum long and distal-
ly rather broad, with large, bluntly pointed,
slightly divergent, apicolateral processes, the
caudal margin between them deeply cleft.
Basistyle with ventral and dorsal roots simple,
the dorsal root stouter; dististyle moderately
curved, tapering distally to slender, bent,
sharp-pointed tip. Aedeagus short and stout,
basal arch extending to i of total length, the
basal arms nearly straight, sides of distal por-
tion nearly straight, tapering to blunt tip.
Parameres (fig. 51g) separate, basal arms
directed laterad with small anterior extension
at lateral ends; each with main body short,
nearly straight proximad, distally tapering
and gently curved ventrolaterad to moderately
stout-pointed, simple tip.

DISCUSSION: C. mulrennani Beck is close-
ly related to C. biguttatus and very similar, but
can be distinguished by its smaller size, nar-
rower, paler wing with much fewer
macrotrichia, paler yellowish brown body, and
more lightly sclerotized spermathecae with
slenderer necks.

IMMATURE STAGES: Thomsen (1937)
described and figured the pupa of C. bigut-
tatus. Jamnback (1966) described and figured
the larva and pupa.

PUPA: Respiratory horn (fig. 136) uniformly
brown, darker than rest of pupa, with 4-6
apical and 2-3 lateral spiracular openings; horn
not narrowed or convoluted near'the middle;
with abundant scale-like spines on median %/,
widest near base, L/W Ratio 5.6. Operculum


(fig. 164) with coarse short spine-like tubercles
over most of disc, extending farther back along
the lateral margins; am setae moderately long,
stout, less than /2 as long as maximum width
of operculum. The d tubercles in line, equidis-
tant, setae 1 and 2 subequal, short; thoracic
surface between and adjacent to tubercles with
fine papillae. Abdomen with fine spines along
anterior margins of segments and sparsely
over rest of segments. The Ipm tubercles (fig.
191) rounded or weakly ridged, occasionally
weakly bifid. Caudal segment with patch of
spines on disc; apicolateral processes with
spines, apex not darkened, directed posteriorly
about parallel to longitudinal axis of body.

MATURE LARVA: Head capsule yellow; fron-
toclypeus 0.187-0.211 mm long. Comb with 7-9
teeth on each side; total comb width 0.032 mm.

LARVAL HABITAT: Wirth (1951),
Williams (1955), Murray (1957), and Snow et
al. (1957) reported hearings of C. biguttatus
from the margins of lakes, pools, streams, and
moist leaf depressions, with moist mud, sand,
or decaying leaves, Jamnback (1965) reared
this species from a semipermanent woodland
pool, a cedar bog, and 4 grassy marsh sites,
with black mud beneath, or grass roots,
humus, or clayey soil. Snow et al. (1957) found
pupae of C. biguttatus commonly in flooded
pools near top reservoir levels, and in recently
flooded wooded bottomlands in the Tennessee
Valley. Hair et al. (1966) recovered this species
from shaded stream and pool margins in
Virginia. Childers and Wingo (1968) reared C.
biguttatus from a temporary pool margin in
Missouri. Gazeau and Messersmith (1970)
reared it in Maryland from dead leaves and
mud in a swamp. Kardatzke and Rowley (1971)
found C. biguttatus in Iowa commonly
breeding in wooded, flowing, freshwater
habitats.

FEEDING HABITS: Jamnback (1965)
reported that C. biguttatus rarely bites man,
and that many literature records may be er-
roneous. Hoffman (1925) reported it "bad" on
cows; Pickard and Snow (1955) feeding heavily
on horses; Downes (1958b) on horses, cattle,
and chickens; Jellison and Philip (1933),
Downes (1958b) and Judd (1959) from birds'
nests; Murray (1957) biting man in Virginia,
and Snow and Pickard (1958) biting man dur-
ing daylight hours in forested bottomlands in
Alabama and Tennessee. Hair and Turner
(1968) regarded it as a general feeder. Childers
and Wingo (1968) collected 1 specimen biting
man in central Missouri.






SEASONAL DISTRIBUTION: C. bigut-
tatus was recorded by Jamnback (1965) from
May to October in New York, where it was
most common from early June through
August. Snow et al. (1957) recorded the species
from June through August in the Tennessee
Valley. In Virginia, Murray (1957) detected a
peak population of C. biguttatus from mid May
to mid July, and Messersmith (1966) collected
adults between 18 April and 22 July with a
population peak in June. Khalaf (1969) found
C. biguttatus abundant along the Gulf Coast of
Louisiana and Mississippi, appearing after the
middle of March, reaching a peak from the 1st
of April to the middle of May, and disappear-
ing by the end of June.

DISTRIBUTION: Eastern North America
from Wisconsin to Nova Scotia and south to
Louisiana and Florida.

FLORIDA DISTRIBUTION (fig. 52): C.
biguttatus is a common species in northern and
western Florida, where it has been collected in
nearly every county from Escambia to Baker
and Putnam counties. All our records are from
light traps. For some reason or other it does
not seem to occur in the Peninsular Florida
Uplands below Gainesville. Records as follows:
Alachua Co., Gainesville, IV-64, T. J. Walker,
LT, 1 female; 7-V-67, FSB, LT, 2 females.
Alachua Co., Hawthorne, 27-IV-68, FSB, LT, 1
female. Alachua Co., O'Leno State Park,
26-IV, 5-14-V-66, W. W. Smith, LT, 3 males, 10


Fig. 52. Florida distribution of Culicoides biguttatus.


females. Baker Co., Glen St. Mary, IV,V-71,
FSB, LT, 7 females. Bay Co., (Beck, 1952).
Escambia Co., Innerarity Point, 29-IV-47,
Rathert, LT, 4 females. Escambia Co., Walnut
Hill, VI-60, FSB, LT, 1 female. Franklin Co.,
Wright Lake near Sumatra, IV-71, G. B. Fair-
child, LT, 50 females. Holmes Co., Bonifay, V,
SBH, LT, 1 female. Jackson Co., Florida
Caverns State Park, 4-V-70, WWW, LT, 3
males, 9 females. Jackson Co., Marianna,
VI-55, SBH, LT, 1 male; V-71, FSB, LT, 5
females. Jackson Co., Sneads, 2-IV-54, SBH,
LT, 1 male. Jefferson Co., Monticello, IV,V-69,
W. H. Whitcomb, LT, 35 females. Leon Co., 3
mi N Tallahassee, V-70, FSB, LT, 4 females.
Leon Co., Tall Timbers, 29-V-73, WWW, LT, 1
female. Liberty Co., Torreya State Park,
15-IV-57, FSB, LT, 2 females; 15-V-71, G. B.
Fairchild, LT, 1 male; 22-IV-67, WWW, LT, 2
males, 12 females. Putnam Co., Welaka,
9-IV-64, H. A. Denmark, LT, 3 females. Santa
Rosa Co., Blackwater River State Forest,
21-V-73, WWW, LT, 1 female. Santa Rosa Co.,
Jay, V-62, T. W. Boyd, LT, 5 females. Taylor
Co., Steinhatchee, 29-IV-49, SBH, LT, 1
female. Wakulla Co., Ochlockonee River State
Park, 29-IV-70, WWW, LT, 1 male, 39 females.
Walton Co., Santa Rosa, 5-V-49, Peterson, LT,
2 females.


Culicoides chiopterus (Meigen)
(fig. 53, 54, 137, 165, 216, 229)


Ceratopogon chiopterus Meigen, 1830:263
(Europe).
Culicoides chiopterus (Meigen); Kieffer,
1925:76 (combination); Root and Hoffman,
1937:156 (male; Maryland; fig. genitalia);
Downes and Kettle, 1952:66 (male; Britain;
fig. genitalia); Kettle and Lawson, 1952:447
(larva, pupa; Scotland; fig.); Foote and
Pratt, 1954:18 (male; Maryland; fig.
genitalia); Downes, 1958b:803 (Ontario);
Jamnback and Wirth, 1963:187 (redescrib-
ed; fig.; North American distribution);
Jamnback, 1965:50 (all stages redescribed;
fig.; biology; New York); Khalaf, 1969:1160
(Louisiana; redescribed, female was juddi
misident.); Battle and Turner, 1971:32
(redescribed; fig.; Virginia); Humphreys
and Turner, 1973:82 (feeding preference;
Virginia).

TYPES: Europe; male syntypes from Win-
them collection in Vienna Museum.

FEMALE:Wing length 0.91 mm.






HEAD: Eyes (fig. 53d) contiguous, bare.
Antenna (fig. 53a) with lengths of flagellar
segments in proportion of 19-12-12-13-13-14-
14-15-21-22-24-24-37;AR 1.15; sensory pattern
3,11-15. Palpus (fig. 53b) with 3rd segment
short and stout, PR 1.9; sensory pit shallow
with small round opening. Proboscis short,
P/H Ratio 0.65; mandible with 6-9 well
developed teeth.

THORAX: Dull, unmarked, brownish black
with faint greenish pollinosity. Legs (fig. 53f)
without distinct pale bands; tibial comb with 5
spines, that nearest the spur longest.

WING (fig. 53c, 229): Pale spots faint, distal
1/ of 2nd radial cell included in a pale spot.
Macrotrichia sparse and confined to wing tip,
vein M3+4 usually with fewer than 8
macrotrichia and cell M4 usually without
macrotrichia excluding those bordering the
wing margin; CR 0.59. Halter pale.

ABDOMEN: Pale brown. Spermathecae (fig.
53e) 2, ovoid, tapering to distinct necks; sub-


equal in size, each measuring 0.049 by 0.036
mm; rudimentary 3rd spermatheca and
sclerotized ring present.

MALE GENITALIA (fig. 53h): Ninth ster-
num bearing broad, shallow, caudomedian ex-
cavation, the ventral membrane not spiculate;
9th tergum with short, apically rounded, wide-
ly spaced, apicolateral processes. Basistyle
with dorsal and ventral roots slender, simple;
dististyle with slightly expanded, rounded tip.
Aedeagus with basal arch extending nearly to
'/2 of total length, basal arms slender; distal
median process tapering to slender tip, a
distinct internal sclerotized point at base of
distal process. Parameres (fig. 53g) separate;
each with stout basal arm directed
anterolaterad, mid-portion moderately stout,
tapering distally to a slender, bare point.

DISCUSSION: C. chiopterus can be
distinguished from the closely related species,
C. obsoletus and C. sanguisuga by its slightly
smaller size, less conspicuous wing pattern and
shorter, stouter, 3rd palpal segment. The


a


621Zjji~cQL~X


,, . ..* " ..- ..... .








d( \








e 4


Fig. 53. Culicoides chiopterus: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f, hind femur and
tibia; g, male parameres; h, male genitalia, parameres removed.






shape of the rounded apicolateral processes of
the male 9th tergum also are distinctive.
Khalaf (1969) redescribed C. chiopterus from
Louisiana. His females, which had vestigial
mandibular teeth, are C. juddi. Khalaf's males
were identical with C. chiopterus, and it cannot
be determined at this point whether he actually
had C. chiopterus or whether the male of C.
juddi is identical with that of C. chiopterus.

IMMATURE STAGES: Kettle and Lawson
(1952), Jamnback and Wirth (1963), and Jamn-
back (1965) described and figured the larva and
pupa.

PUPA: Respiratory horn (fig. 137) uniformly
medium to dark brown, darker than rest of
pupa, with 3-5 apical and 3-5 lateral spiracular
openings; horn without spines, slightly nar-
rowed medially, widest near base, L/W Ratio
5.0. Operculum (fig. 165) with very long, hair-
like spines confined to lateral margins and
disc; am setae long and slender, about 0.75 as
long as maximum width of operculum. The d
tubercles 1 to 3 nearly in line, 1 and 2 much
closer than 2 and 3; setae of the 1st 2 long and
overlapping, 2 more than twice as long as 1; d
tubercles often with several separate points;
thoracic surface between and adjacent to
tubercles nearly smooth. Abdomen with fine
spines confined mostly to anterior margins of
segments; Ipm tubercles with a single point
and a subapical seta. Caudal segment with a
transverse band of spines across disc;
apicolateral processes reduced in size, with
scales on inner margin, tips sometimes darken-
ed, directed posteriorly nearly parallel to
longitudinal axis of body.

Note: C. chiopterus pupae can be distin-
guished from those of C. sanguisuga and C. ob-
soletus by the dark respiratory horn, more
spinules on the lateral margin of the oper-
culum, and the relatively longer ad setae, the
longer of which is longer than the respiratory
horn.

MATURE LARVA: Head capsule pale yellow;
dorsal head length 0.118 mm. Comb with 9
teeth on each side; total comb width 0.019 mm.
Each thoracic segment (fig. 216) with lateral
pigmented oval spot about halfway between
anterior and posterior margins, that on
metathorax more elongate than those on pro-
and mesothorax. The larva of C. chiopterus can
be distinguished from that of C. obsoletus and
C. sanguisuga by its shorter head capsule.

LARVAL HABITAT: Kettle' and Lawson
(1952) collected C. chiopterus from pats of cow
manure in pastures in Scotland. Jamnback


(1965) reared it from moist straw and moist
soil mixed with chicken manure in New York.

FEEDING HABITS: Remm (1956) reported
this species to be a bloodsucking pest of man in
Estonia, especially in the spring and fall.
Messersmith (1965) took 8 females in light
traps operated in poultry houses in Virginia.
Humphreys and Turner (1973) collected 2
females from galliform birds in Virginia.

SEASONAL DISTRIBUTION: Jamnback
(1965) took C. chiopterus from May until Oc-
tober in New York. Messersmith (1966)
trapped adults in Virginia from 24 May to 2
September, with a peak in July.

DISTRIBUTION: Europe, northern Asia,
North America from California and Alaska to
Quebec and Florida.

FLORIDA DISTRIBUTION (fig. 54): C.
chiopterus is a rare species in Florida. We have
records from: Alachua Co., Gainesville,
16-III-67, W. W. Smith, LT, 1 male; IX,XI-67,
FSB, LT, 2 males. Alachua Co., O'Leno State
Park, 13-II-66, W. W. Smith, 1 female. Hardee
Co., Ona, VII-70, E. Irons, LT, 1 male, 1
female. Jefferson Co., Monticello, VII,IX-69,
W. H. Whitcomb, LT, 1 male, 1 female. Liberty
Co., Torreya State Park, 20-V-66, H. V.
Weems, Jr., LT, 1 male; 22-IV-67, WWW, LT 1
male.


Fig. 54. Florida distribution of Culicoides chiopterus.






Culicoides crepuscularis Malloch
(fig. 55, 56, 138, 166, 192, 217, 258)


Culicoides crepuscularis Malloch, 1915:303
(male, female; fig.; Illinois); Hoffman,
1925:298 (female; fig.; records); Root and
Hoffman, 1937:159 (redescribed; fig.;
distribution); Thomsen, 1937:70 (immature
stages); Wirth, 1952a:167 (redescribed; fig.;
California); Beck, 1952:103 (Florida);
Khalaf, 1952:356 (Oklahoma); Foote and
Pratt, 1954:19 (redescribed; fig.; distribu-
tion); Williams, 1955:115 (Michigan;
habitats); Wirth and Bottimer, 1956:263
(Texas; seasonal distribution); Williams,
1956b:303 (Bermuda); Murray, 1957:77
(Virginia; habitat and seasonal
distribution); Williams, 1957:62 (Bermuda;
habitats); Robinson, 1961:898 (host of
filarial worms); Jones, 1961a:703 (larval
habitats); Frost, 1964:155 (record;
Highlands Co., Florida); Jamnback,
1965:52 (redescribed; all stages; fig.;
biology; New York); Das Gupta and
Hansens, 1965:159 (New Jersey; descrip-
tive notes; fig.); Atchley, 1967:972 (redes-
cribed; fig.; New Mexico); Childers and
Wingo, 1968:11 (Missouri; biology; fig.);
Khalaf, 1969:1158 (Louisiana; seasonal in-
cidence); Battle and Turner, 1970:426
(North Carolina; larval habitats); Battle
and Turner, 1971:34 (redescribed; fig.;
Virginia); Kardatzke and Rowley, 1971:216
(Iowa; larval habitats); Rowley and Corn-
ford, 1972:1208 (palpal sensilla by scanning
electron microscopy).

TYPE: Holotype, male, Dubois, Illinois, 24
April 1914, J. R. Malloch (Illinois Natural
History Survey, Urbana).

FEMALE: Wing length 1.30 mm.

HEAD: Eyes (fig. 55d) narrowly separated;
bare. Antenna (fig. 55a) with lengths of
flagellar segments in proportion of
23-15-15-15-15-15-15-15-33-33-35-35-55; AR
1.40; sensory pattern 3-15. Palpus (fig. 55b)
with lengths of segments in proportion of
10-22-35-12-14; PR 2.2; 3rd segment greatly
swollen, with a large, deep, round, sensory pit.
Proboscis moderately long, P/H Ratio 0.80;
mandible with 14 teeth.

THORAX: Brownish; mesonotum grayish
brown pollinose, with dark brown markings
forming a narrow median line terminating at
the transverse suture in a diamond shaped
marking, and a pair of crescent shaped lateral
patches at sutural level; the brown patches fre-


quently become reduced in extent. Legs (fig.
55f) brown; fore femur with subapical pale ring
and all tibiae with subbasal pale rings; tibial
comb with 4 spines, that nearest the spur
longest.

WING (fig. 55c, 258): Second radial cell in a
dark area; pattern of conspicuous oval or
rounded pale spots centered between the veins;
a large pale spot over r-m crossvein extending
broadly to anterior wing margin; cell R5 with a
large poststigmatic pale spot and a large,
usually quadrate pale spot in distal portion of
cell, usually broadly meeting wing margin well
before tip of cell; cell M1 with 2 pale spots, the
distal spot not reaching wing margin; cell M2
with a small pale spot behind mediocubital
fork and 2 pale spots in distal portion of cell,
the distal spot not reaching wing margin; cell
M4 with a large rounded pale spot touching
wing margin posteriorly; anal cell with a large
pale area at base and a large double pale spot in
distal portion of cell. Macrotrichia long and
dense, covering all of wing to base of cell M2
and anal cell; CR 0.56; radial cells complete.
Halter pale.

ABDOMEN: Pale brown. Spermatheca (fig.
55e) 1, oval with short, slender neck; measur-
ing 0.115 by 0.062 mm.

MALE GENITALIA (fig. 55h): Ninth ster-
num with broad, shallow, caudomedian excava-
tion, the ventral membrane spiculate; 9th
tergum long and slightly tapering, with long,
slender, apicolateral processes, the caudal
margin between them slightly cleft. Basistyle
with ventral root inconspicuous, dorsal root
slender, dististyle nearly straight, with
slender, bent, pointed tip. Aedeagus short and
broad, basal arch extending to nearly 0.6 of
total length, the basal arms rather stout, and
slightly curved; distal portion short, tapering
to blunt, moderately stout, median process.
Parameres (fig. 55g) separate; each with small
basal knob bearing a distinct anterior process;
midportion nearly straight, slightly swollen
proximad and tapering distally to slender, sim-
ple, median point abruptly twisted ventrad.

DISCUSSION: Populations of C.
crepuscularis from the Atlantic and Gulf
Coastal Plain and from Bermuda have a high
incidence of more extensive pale wing spots,
especially a broadly quadrate distal pale mark-
ing in cell R5, and the dark brown mesonotal
markings are not as extensive as those from
the remainder of the country. Because these
populations appear to intergrade irregularly
with the typical form, we have been unable to
work out any readily apparent infraspecific









a


b


__ vsna -iseS r^-.- '- ..
SR
ZT


Fig. 55. Culicoides crepuscularis: a, antenna; b, palpus; c, wing; d, eye separation; e, spermatheca; f, hind femur and
tibia; g, male parameres; h, male genitalia, parameres removed.


taxa. We believe an analysis of variation in this
species using biosystematic procedures and
numerical taxonomy such as used by Atchley
(1970) on the subgenus Selfia, might prove
quite productive.
C. crepuscularis is one of the commonest and
most widespread species in North America and
is especially dominant in grassy, sunlit
habitats outside the deep forest. It could be a
prime suspect in the search for vectors of many
pathogenic organisms, but has not proved easy
to colonize or to handle in the laboratory, and
the exact details of its larval and adult habits
remain virtually unknown. Beck (1958) noted
the occurrence of several gynandromorphs of
C. crepuscularis in light trap collections in
Jackson County, Florida. The abdominal cavi-
ty was always occupied by a mermithid worm.
We have noted frequent parasitism of this
species by mermithid nematodes, and
biological control by this parasite may be
responsible for the failure of C. crepuscularis
to build up extremely dense populations such
as have been observed in C. varipennis, occurr-
ing at times in the same habitats as C.
crepuscularis.


IMMATURE STAGES: Thomsen (1937)
and Fox (1942) described and figured the pupa.
Jamnback (1965) described and figured the lar-
va and pupa.

PUPA: Exuviae brownish. Respiratory horn
(fig. 138) much darker apically, with 6-8 apical
and 3-4 lateral spiracular openings; horn nar-
rowed near middle but without transverse con-
volutions, with well developed transverse
scale-like spines on median %; horn widest near
base, L/W Ratio 6.7. Operculum (fig. 166) dark
brown, densely covered with short stout
spines; am setae short and stout, less than 0.3
as long as maximum width of operculum. The d
tubercles in line, about equidistant; d setae 1
and 2 moderately long, 1 slightly longer than 2,
not overlapping; thoracic surface between and
adjacent to tubercles rough, covered with
numerous papillae. Abdomen with fine spines
confined mostly to anterior margins of
segments, sparse elsewhere; 1pm tubercles (fig.
192) rounded, some weakly pointed or bifid.
Caudal segment with a patch of spines on disc;
apicolateral processes with spines, distal 1/


P
r) 11 Id ~ Li~ 11 lr-41 11- r


7'_







dark brown; directed posteriorly at an angle of
30" or less to the longitudinal axis of body.

MATURE LARVA: Head capsule pale yellow;
dorsal head length 0.172 mm. Comb with 8
unequal teeth on each side; total comb width
0.026 mm. Dorsum of all thoracic segments
(fig. 217) lightly and rather uniformly
pigmented reddish brown; meso- and
metathorax with more heavily pigmented
round lateral spots at midlength of each seg-
ment.

LARVAL HABITAT: Wirth and Bottimer
(1956) reared C. crepuscularis in Texas from
mud at pond margins, puddles at stock tank
overflows, and in septic tank effluents.
Williams (1956b, 1957) in Bermuda reported C.
crepuscularis breeding in a variety of marsh
habitats, with wide range of salinity, from
freshwater to that of seawater, but not in the
shade of mangroves. Snow et al. (1957) found
pupae among rushes in a rainfilled roadside
ditch and in a seepage ditch with marginal
grass in Tennessee. Jones (1959) reported light
breeding of C. crepuscularis in household sep-
tic tank effluents in Texas, along with much
higher populations of C. uariipennis. Jones
(1961a) stated that C. crepuscularis will breed
in most freshwater soil habitats suitable for
Culicoides breeding.
Jamnback (1965) reared C. crepuscularis in
New York from cattle hoofprints in marshy
meadows, from stream margins, a lagoon
margin, and a marshy drainage ditch with
substrate usually of clayey mud with grass
roots and humus. Hair et al. (1966) found the
species in Virginia extremely abundant in
polluted mud near livestock watering troughs
and pools in open sunlight with little vegeta-
tion. Childers and Wingo (1968) gave notes on
larval habitats in central Missouri; they were
found in sewage lagoon effluent areas, edges of
temporary and permanent ponds, a roadside
drainage ditch in a forested area, the edge of a
sand bank of an isolated pool, and spring
seepage areas. Larvae and pupae were most
abundant in exposed or nearly exposed mud
flats. Battle and Turner (1970) reared C.
crepuscularis from 5 breeding sites in North
Carolina: mud at a ditch margin, mud at a
creek margin, mud at the edge of a hog pond,
sandy mud at a river margin, and mud at a lake
margin. Kardatzke and Rowley (1971) found C.
crepuscularis the most abundant Iowa species
in flowing freshwater habitats, preferring non-
wooded, herbaceous-mud situations.


(1957) from birds' nests. Edmunds and Keener
(1954) report the species as a pest of man in
Nebraska, but elsewhere it bites man rarely
(Snow et al. 1957) or not at all (Downes, 1958).
Snow (1955) compared diurnal versus noctur-
nal feeding habits and ground level versus tree
canopy distribution in a bottomland forest in
Tennessee; C. crepuscularis was nocturnal and
did not penetrate the forest, preferring the
canopy level at the forest edge. Pickard and
Snow (1955) recorded the species biting man in
Tennessee. Bennett (1960) and Fallis and Ben-
nett (1960, 1961) took C. crepuscularis biting a
variety of birds in Ontario, where it was in-
criminated as an intermediate host of a
Haemoproteus blood parasite of crows and
purple finches. Robinson (1961) took this
species in starling traps and was able to infect
it with a filarial parasite from a starling. Hair
and Turner (1968) stated that C. crepuscularis
was ornithophilic. Messersmith (1965) trapped
large numbers of engorged females in poultry
houses in Virginia. Jamnback (1965) did not
collect C. crepuscularis biting or annoying man
in New York.

SEASONAL DISTRIBUTION: Beck (1958)
reported peak populations of C. crepuscularis
in March in Florida. Wirth and Bottimer (1956)
reported increased numbers in both spring and
fall in Texas. Snow et al. (1957) recorded an ear-
ly June peak in Tennessee. Lewis (1959)
reported a peak in late July and another in ear-
ly September in Connecticut. Jamnback (1965)
stated that in New York C. crepuscularis pro-
duced more than 1 generation per year with
adult populations highest in mid to late sum-
mer. The species was more abundant in the
warmer parts of the state than in the cooler
Adirondacks. Childers and Wingo (1968) gave
notes on seasonal emergence from breeding
sites in central Missouri; at a sewage lagoon
there was a peak emergence in mid April and a
2nd in mid July. In light traps they recorded 3
population peaks: the 1st or overwintering
generation peaked 17-24 April, the second 9-15
May, and the 3rd 13-19 June. They trapped
adults as late as mid October and believed that
C. crepuscularis had 3-5 generations in their
area. Khalaf (1969) considered C. crepuscularis
an aestival species in Louisiana, with a high
population in March which it maintained until
October.

DISTRIBUTION: Widely distributed in
North America from southern Alaska and
Canada to northern Mexico, Florida, and Ber-
muda.


FEEDING HABITS: Hoffman (1925)
recorded C crepuscularis feeding on chickens; FLORIDA DISTRIBUTION (fig. 56): C.
Jellison and Philip (1933) and Snow et al. crepuscularis is a common and widespread






species in Florida, where we have collected it in
light traps in nearly every county from Escam-
bia and Duval to Monroe counties. We have
Florida records from: Alachua Co., Gainesville,
V,VI-60, G. K. Hicks, LT, 10 males, 10
females; 10-VI-65, 11-IV-66, W.W.Smith, LT, 2
females; 20-IV-67, WWW, malaise trap, 1
male, 7 females; 27-IV-70, FSB, LT, 6 females;
3-V-73, R. I. Sailer, LT, 3 females. Alachua Co.,
Island Grove, 23-V-50, Jones, LT, 1 female;
4-VII-61, 1-VIII-62, FSB, LT, 1 male, 1 female.
Alachua Co., Newnans Lake, 22-II-65, W. W.
Smith, LT, 1 female. Baker Co., Macclenny,
VI-71, FSB, LT, 1 female. Baker Co., Olustee,
1-XI-68, R. E. Woodruff, LT, 2 females;
VI,VII,VIII,IX-71, FSB, LT, 12 females. Bay
Co., Panama City Beach, 8-VIII-50, SBH, LT,
2 females. Charlotte Co. (Beck, 1952). Citrus
Co., Crystal River, V,XII-49, 3-11-50, 8-IV-54,
Hudson, LT, 1 male, 4 females. Clay Co.,
Keystone Heights, 6-IV-54, Davis, LT, 1 male.
Duval Co., Arlington, 25-VI-50, SBH, LT, 1
female. Duval Co., Jacksonville, 28-111-50,
SBH, LT, 1 female. Escambia Co., Innerarity
Point, 31-V-54, Rathert, LT, 4 females. Escam-
bia Co., Molino, 12-VIII-69, FSB, LT, 1 female.
Gilchrist Co., Suwannee River, 26-V-62, G. K.
Hicks, LT, 3 females. Gilchrist Co., Trenton,
X-61, SBH, LT, 3 females. Gulf Co., St. Joseph
State Park, 1-V-70, WWW, LT, 1 female.
Hardee Co., Wauchula, 13-VI-60, W. Jernigan,
LT, 1 male. Highlands Co., Highlands Ham-
mock State Park, 15-IV-70, WWW, LT, 3
females. Highlands Co., Lake Placid, 19-IV-70,


Fig. 56. Florida distribution of Culicoides crepuscularis.


J. Layne, LT, 1 male. Hillsborough Co., David
Island, VIII-61, SBH, LT, 1 male.
Hillsborough Co., Gibsonton, 4-11-49, SBH,
LT, 1 female. Indian River Co., (Beck, 1952).
Jackson Co., Florida Caverns State Park,
26-V-73, WWW, LT, 1 male. Marianna,
21-VIII-54, Calloway, LT, 1 female. Sneads,
2-IV-54, Howell, LT, 1 gynandromorph;
15-III-55, Howell, LT, 1 female. Jefferson Co.,
Monticello, IV-69, W. H. Whitcomb, LT, 1,000
females. Lake Co., Goose Prairie, 29-1-54,
SBH, LT, 1 female. Lake Co., Leesburg,
20-VI-49, 31-1-50, Braddock, LT, 2 males. Lee
Co., (Beck, 1952). Lee Co., (Beck, 1952). Leon
Co., 3 mi N Tallahassee, V-70, FSB, LT, 50
males, females. Levy Co., Gulf Hammock,
V-58, SBH, LT, 1 female. Liberty Co., Torreya
State Park, 20-V-66, H. V. Weems, Jr., LT, 1
male, 2 females. Manatee Co., (Beck, 1952).
Marion Co., Citra, 22-VIII-50, Fowler, LT, 1
male; 20-IV-54, Clemmons, LT, 2 males, 1
female. Marion Co., Juniper Springs, 28-IV-70,
WWW, LT, 4 females. Monroe Co. (Beck,
1952). Orange Co., Lake Magnolia Park,
6-VIII-70, E. Irons, LT, 2 males, 1 female.
Pasco Co., St. Leo, 7-III-50, SBH, LT, 1
female. Pinellas Co., Largo, 26-VI-50, Hickey,
LT, 1 male. Polk Co., Lakeland, 30-III-54,
Emerson, LT, 1 female. Putnam Co., Lon's
Lake, V,VI,VII-71, FSB, LT, 30 females. Put-
nam Co., Welaka, 9-IV-64, H. A. Denmark, LT,
1 male, 6 females. Santa Rosa Co., Blackwater
River State Forest, 25-V-71, G. B. Fairchild,
LT, 1 female; 25-V-73, WWW, LT, 2 males, 1
female. St. Johns Co., (Beck, 1952). Sumter
Co., Center Hill, 15-VIII-50, Scholevat, LT, 1
male. Sumter Co., Coleman, no date, Bridges,
LT, 1 male. Taylor Co., Steinhatchee, 20-V-50,
29-IV-54, Wood, LT, 3 females. Walton Co.,
Santa Rosa, 5-V-50, Peterson, LT, 2 females.


Culicoides debilipalpis Lutz
(fig. 57, 58, 248)


Culicoides debilipalpis Lutz, 1913:60 (female;
Brazil; fig. wing); Costa Lima, 1937:415
(Brazil; notes; fig. palpus); Macfie, 1937:7
(female redescribed; Trinidad); Forattini,
1957:383 (redescribed; fig.; distribution);
Wirth and Blanton, 1959:442 (redescribed;
fig.; Panama); Messersmith, 1964:339
(Virginia; breeding site); Smith, 1965a:65
(Florida; reared from tree holes);
Messersmith, 1966:94 (Virginia);Khalaf,
1967a:151 (descriptive notes; Louisiana);
Smith and Varnell, 1967:520 (Florida; tree
holes); Wirth and Blanton, 1971a:34
(redescribed; distribution); Wirth and Blan-











%~D6~iiI~


.,-.


~~C


Fig. 57. Culicoides debilipalpis: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f,
tibia; g, male parameres; h, male genitalia, parameres removed.


ton, 1971b:75 (synonym: khalafi; compared
with torreyae).
Culicoides khalafi Beck, 1957:104 (male,
female; Florida; fig. male genitalia).

TYPES: Syntypes from Brazil: Anhemby,
Sao Paulo; Formoso, Serra de Bocaina; Saltos
de Iguassu, Parana (in Instituto Oswaldo Cruz,
Rio de Janeiro).

FEMALE: Wing length 0.80 mm.

HEAD: Eyes (fig. 57d) moderately separated,
with short interfacetal pubescence. Antenna
(fig. 57a) with lengths of flagellar segments in
proportion of 28-20-22-24-26-26-26-26-26-
28-30-32-44; AR 0.83; sensory pattern 3,8-10.
Palpus (fig. 57b) with 3rd segment long and
slender, PR 2.2; sensory pit small and deep
with pit deeper than diameter of pore opening.
Proboscis moderately long, P/H Ratio 1.00;
mandible with 15 teeth.

THORAX: Brown; mesonotum with moderate-
ly prominent dark brown sublateral patches.
Legs (fig. 57f) with narrow pale rings


hind femur and


subapically on fore and mid femora, subbasal
pale rings on all tibiae, and apex of hind tibia
pale; tibial comb with 4 spines, that nearest the
spur longest.

WING (fig. 57c, 248): Second radial cell dark
to tip; 2 poststigmatic pale spots in cell R5
usually well separated, the posterior spot
located slightly proximad of the other; distal
pale spot in cell R5 transverse, not filling space
between anterior wing margin and vein Ml; 1
distal pale spot in distal part of anal cell; pale
spot lying behind medial fork distinct, but
none lying immediately in front of
mediocubital fork. Wing moderately hairy,
macrotrichia extending to base of cell M2 in 2
rows; CR 0.65. Halter infuscated.

ABDOMEN: Dark brown. Spermathecae (fig.
57e) 2 plus rudimentary 3rd and sclerotized
ring; ovoid with long, rather stout necks;
slightly unequal, measuring 0.060 by 0.039
mm and 0.050 by 0.032 mm.

MALE GENITALIA (fig. 57h): Ninth ster-
num with broad, shallow, caudomedian excava-


~_~=~3~a~=~6=~=~3~~r3~






tion, ventral membrane not spiculate; 9th
tergum long and tapering with short,
triangular, apicolateral processes. Basistyle
with foot-shaped ventral root, dorsal root
slender, dististyle slender and curving.
Aedeagus with basal arch extending to more
than % of total length, mesal apex of arch nar-
row, basal arms straight; distal process
pointed apically without apparent subapical
projections. Parameres (fig. 57g) separate; each
with knobbed base, stem bent near base,
slender and straight in midportion, with well
developed ventral lobe; distal portion not
greatly elongated, with slender pointed tip and
4-6 lateral fringing spines.

DISCUSSION: Earlier records and descrip-
tions of C. debilipalpis were much confused by
inability to discriminate between various
members of the debilipalpis group of species,
most of which are readily distinguishable only
by minute structural features such as the male
genitalia, spermathecae, antennal sensorial
pattern, and nature of the 3rd palpal segment
and its sensory pit. Wirth and Blanton (1971a)
pointed out the characters by which C.
debilipalpis could accurately be distinguished,
and described C. hoffmani and C. eadsi, with
which it would most easily be confused. In
another paper they (1971b) pointed out the
synonymy of C. khalafi Beck.

IMMATURE STAGES: Undescribed.

LARVAL HABITAT: Williams (1964)
reared C. debilipalpis from rotting cocoa pods
and bamboo stumps in Trinidad. Messersmith
(1964) reared the species from wet debris in a
tree cavity in Virginia. Smith (1965b) reared it
from wet tree holes and stump holes in Florida,
where Smith and Varnell (1967) reported that
it preferred substrates with pH less than 8.7.

FEEDING HABITS: Messersmith (1966)
collected 11 specimens in chicken houses in
Virginia, 3 of which were engorged. This
species is reported to bite man and horses in
South America.

SEASONAL DISTRIBUTION: Messer-
smith (1966) trapped C. debilipalpis from mid
June to the 1st week of August with a peak in
July in Virginia.

DISTRIBUTION: Southeastern United
States from Maryland and Kentucky to
Florida and Louisiana, in Central America
from Honduras to Panama, and in South
America to Trinidad, Brazil, ard Argentina. In
Texas and Mexico C. debilipalpis is replaced by
C. eadsi, and the West Indies by C. hoffmani.


FLORIDA DISTRIBUTION (fig. 58): C.
debilipalpis is abundant in Florida only locally
where its tree hole habitats are available, but
we have taken it in 10 scattered counties from
one end of the state to the other. Our records:
Alachua Co., Gainesville, 22-1-65, W. W.
Smith, reared from tree hole, 1 female; VIII-66,
W. W. Smith, LT, 1 female; VI,VII,
VIII,IX-68, FSB, LT, 20 males, 50 females.
San Felasco Hammock, 14-V-66, 26-1-66, W.
W. Smith, LT, 5 males, 10 females; 16-III-65,
W. W. Smith, reared from tree hole, 1 male.
Wacahoota, 19-1-66, W. W. Smith, reared from
tree hole, 2 males, 1 female. Calhoun Co.,
Blountstown, IX-68, FSB, LT, 2 females. Col-
lier Co., Fakahatchee Island, VIII-72, Blanton
and Meyer, LT, 6 females. Dade Co., Miami,
17-II-61, A. S. Mills, LT, 1 female. Highlands
Co., Highlands Hammock State Park,
15-IV-70, WWW, LT, 1 male, 3 females.
Jackson Co., Florida Caverns State Park,
26-V-73, WWW, LT, 1 male, 3 females.
Jackson Co., Marianna, V-55, SBH, LT, 1
male, 1 female (types of khalafi Beck). Jeffer-
son Co., Monticello, VII,XI,X-69, W. H. Whit-
comb, LT, 10 females. Leon Co., 3 mi N
Tallahassee, V-70, FSB, LT, 25 females. Leon
Co., Tall Timbers, VII-68, Bhaktar and Baker,
LT, 3 females. Liberty Co., Torreya State Park,
7-VII-65, H. V. Weems, Jr., LT, 3 females;
30-VI-57, V-71, FSB, LT, 10 males, 20 females;
22-IV-67, WWW, LT, 1 female; 15-V-71, G. B.
Fairchild, 1 female.


Fig. 58. Florida distribution of Culicoides debilipalpis.






Culicoides edeni Wirth and Blanton
(fig. 59, 60)


Culicoides edeni Wirth and Blanton, 1947b:23
(male, female; Florida; fig.).
Culicoides haematopotus Malloch (in part, mis-
identified); Beck, 1952:106 (in part, south
Florida records); Foote and Pratt, 1954:23
(in part, south Florida records); Beck,
1956:134 (in part, south Florida records);
Frost, 1964:155 (record; Highlands County,
Florida).

TYPES: Holotype, female, allotype, male,
Rock Springs, Orange Co., Florida, 21 April
1970, W. W. Wirth, light trap (USNM 71483).

FEMALE: Wing length 1.12 mm.

HEAD: Eyes (fig. 59d) narrowly to moderate-
ly separated, bare. Antenna (fig. 59a) with
lengths of flagellar segments in proportion of
25-16-17-18-20-20-20-20-45-50-53-57-72; AR
1.78; sensory pattern variable, usually 3,10-15,


sensoria sometimes also present on 5,7,9.
Palpus (fig. 59b) with lengths of segments in
proportion of 15-27-50-12-20; 3rd segment
moderately swollen, PR 2.2; sensory pit
shallow with large round opening. Proboscis
moderately long, P/H Ratio 0.83; mandible
with 15 teeth.

THORAX: Yellowish brown, with heavy pale
gray pollinosity; mesonotum with pattern of
narrow, irregular, dark brown, longitudinal
markings present. Legs (fig. 59f) pale yellowish
brown, knee spots blackish, no distinct dark
brown bands present; tibial comb with 4
spines, that nearest the spur longest.

WING (fig. 59c): Pattern nearly identicalwith
that of C. haematopotus, but not quite as
distinct, the ground color of the wing
somewhat paler; in addition to the pale mark-
ings found in C. haematopotus, wing is
characterized by a pale area across costal cell
from r-m crossvein to costal margin, and a
distinct pale spot near wing margin in prox-
imal portion of anal cell. Macrotrichia long and


a


b


^^.^--- *^-:^^:::-





'****'^;;7 .** 3 ^^~-" C
/' "'",,',iK1! !!!'_ . .. .:' __ .. .i'l^ *.|il .ll .*' il"" :' . _.',


Fig. 59. Culicoides edeni: a, antenna; b, palpus; c, wing; d, eye separation; e, spermathecae; f, hind femur and tibia; g,
male parameres; h, male genitalia, parameres removed.


''
g







numerous, extending nearly to bases of medial
and anal cells; CR 0.58. Halter pale.

ABDOMEN: Pale yellowish. Spermathecae
(fig. 59e) 2 plus rudimentary 3rd and promi-
nent sclerotized ring of characteristic shape;
functional spermathecae unequal, measuring
0.061 by 0.038 mm and 0.054 by 0.032 mm in-
cluding the long sclerotized necks; sper-
mathecae slightly broader in proportion to
length than in C. haematopotus.


MALE GENITALIA (fig. 59g,
distinguishable from those
haematopotus.


h):
of


In-
C.


DISCUSSION: C. edeni is very similar to C.
haematopotus and was confused with it for
many years. C. haematopotus, which is
widespread in North America, can be
distinguished from C. edeni by its darker color,
with dark brown thorax and legs, and by the
wing markings, in which the pale spot over the
r-m crossvein does not extend to the costal
margin, and the proximal pale spot near the
wing margin in the anal cell is lacking or in-
distinct. The pupa of C. haematopotus is larger
and darker in color than in C. edeni, with spines
better developed, with the area between the am
tubercles bearing moderate to distinct spine-
like markings ranging to distinct spines.

IMMATURE STAGES: Jones (1961b) gave
notes on the pupa, from Wekiwa Springs,
Florida. It differs from C. haematopotus:
"Smaller, color light brown, less spinose.
Respiratory trumpet not darkened so exten-
sively, apex narrowly dark; both trumpets
with five apical and three basal spiracular
openings. Operculum disc with few uniformly
medium-sized spines, most of these confined to
lateral row on each side; area between and
beyond a.m.'s bare. Area between d's devoid of
distinct markings".

LARVAL HABITAT: Wirth and Keller
reared edeni from mud at the margin of the
stream below Wekiwa Springs in Orange Coun-
ty in 1951. Wirth reared several specimens
from pupae collected on the sloping, sandy
margins of a canal on Congress Road near Lan-
tana in August 1951. Wirth collected pupae
from sandy humus at the margins of small
spring areas at Rock Springs in Orange Coun-
ty in 1970 (all unpublished data).

FEEDING HABITS: Unknown.

SEASONAL DISTRIBUTION: Our Florida
collections are distributed from April to
December.


Fig. 60. Florida distribution of Culicoides edeni.

DISTRIBUTION: Florida, Bahamas.

FLORIDA DISTRIBUTION (fig. 60): C.
edeni seldom has been taken in West Florida,
with records from Escambia and Wakulla
counties, but from Jacksonville and
Gainesville southward it becomes abundant
and replaces its close relative C. haematopotus
entirely in south Florida. Frost (1964) recorded
this species (as C. haematopotus) from
Highlands County in December.
Our records: Alachua Co., Gainesville,
V,VI,VII,VIII-67, FSB, LT, 3 males, 1 female;
27-IV-70, WWW, LT, 1 female. Brevard Co.
(Beck, 1952, as C. haematopotus). Charlotte
Co. (Beck, 1952). Citrus Co., Crystal River,
18-IX-50, Hudson, LT, 3 females. Collier Co.,
Collier Seminole State Park, 17-V-73, WWW,
LT, 2 males, 2 females. Collier Co., Ochopee,
IX-71, FSB, LT, 1 female. Duval Co. (Beck,
1952). Escambia Co., Innerarity Point,
29-IV-49, Rathert, LT, 3 males, 16 females.
Gilchrist Co., Trenton, V-61, SBH, LT, 1
female. Glades Co., Palmdale, 14-VII-70, E.
Irons, LT, 1 female. Gulf Co., St. Joseph St.
Park, 1-V-70, WWW, LT, 13 females. Hardee
Co., Ona, VII-70, E, Irons, LT, 3 females.
Highlands Co., Archbold Biol. Sta., 3-19-IV-70,
WWW, LT, 2 females. Highlands Co.,
Highlands Hammock State Park, 15-IV-70,
WWW, LT, 1 male, 2 females. Highlands Co.,
Lake Placid, 19-IV-70, J. N. Layne, LT, 1 male,
2 females. Hillsborough Co., Gibsonton, IV-49,
SBH, LT, 1 male. Lee Co., Fort Myers,
20-XII-49, Keith, LT, 2 females. Lee Co.,




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